Pleurotus placentodes, originally described from Sikkim, rediscovered after 164 years

Abstract

Although many species of Pleurotus are commonly known to be important edible mushrooms, the species delimitation of the genus is often controversial due to phenotypic plasticity and morphological stasis. In the present paper, Pleurotus placentodes, a conspicuous species originally described from Sikkim by M.J. Berkeley in 1852 and so far known only from the type collection, is documented based on specimens recently gathered in the eastern Himalayas and Hengduan Mountains, southwestern China. The morphological and molecular phylogenetic data indicate that this species is an independent taxon, and accordingly the previously proposed synonym of Pleurotus djamor is rejected. Pleurotus placentodes differs from the majority of other related species assigned to Pleurotus by the ellipsoid to subovoid basidiospores with a lower ratio of length/ width and the geographical distribution in the subalpine habitat of the Himalayan Mountains.

Full text

Phytotaxa 267 (2): 137–145
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Genevieve Gates: 17 Jun. 2016; published: 4 Jul. 2016
http://dx.doi.org/10.11646/phytotaxa.267.2.6
137
Pleurotus placentodes, originally described from Sikkim, rediscovered after 164
years
XIAO-BIN LIU1,2, JING LI1,2,4, EGON HORAK3 & ZHU L. YANG1*
1 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kun-
ming 650201, Yunnan, China
2 University of Chinese Academy of Sciences, Beijing 10049, China
3 Nikodemweg 5, Innsbruck, Austria
4 Key Laboratory of Conservation and Utilization for Bioresources and Key Laboratory of Microbial Diversity in Southwest China,
Ministry of Education, Yunnan University, Kunming 650091, Yunnan, China
*e-mail: fungi@mail.kib.ac.cn
Abstract
Although many species of Pleurotus are commonly known to be important edible mushrooms, the species delimitation of
the genus is often controversial due to phenotypic plasticity and morphological stasis. In the present paper, Pleurotus plac-
entodes, a conspicuous species originally described from Sikkim by M.J. Berkeley in 1852 and so far known only from the
type collection, is documented based on specimens recently gathered in the eastern Himalayas and Hengduan Mountains,
southwestern China. The morphological and molecular phylogenetic data indicate that this species is an independent taxon,
and accordingly the previously proposed synonym of Pleurotus djamor is rejected. Pleurotus placentodes differs from the
majority of other related species assigned to Pleurotus by the ellipsoid to subovoid basidiospores with a lower ratio of length/
width and the geographical distribution in the subalpine habitat of the Himalayan Mountains.
Key words: Agaricales, edible mushroom, Himalayas, Pleurotaceae, taxonomy
Introduction
Many species of Pleurotus (Fr.) P. Kumm. (Fries 1821: 178) Kummer (1871: 24) have great economic, dietary and
ecological importance. Therefore, numerous studies on the taxonomy of the saprobic species assigned to the genus
have been carried out (Pilát 1935; Corner 1981; Hilber 1982; Singer 1986; Vilgalys et al. 1993; Vilgalys & Sun 1994;
Segedin et al. 1995; Petersen & Krisai-Greilhuber 1996, Li & Yao 2004, 2005; Zervakis et al. 2004; Huang et al. 2010;
Li et al. 2014).
Due to phenotypic plasticity and morphological stasis, the delimitation of species in Pleurotus has often been
controversial, as there are often many synonyms under a single accepted species on one hand and also names formally
and validly published but overlooked on the other hand. Therefore, besides the morphological features, sexual
intercompatibility tests and cultural characters have been used for species delimitation (Petersen 1992, 1995; Petersen
& Hughes 1993, 1997; Vilgalys & Sun 1994). In the last twenty years, molecular phylogenetic data have also been
employed in understanding the generic limit and relationships of the species in Pleurotus (Vilgalys & Sun 1994;
Albertó et al. 2002; Moncalvo et al. 2002; Zervakis et al. 2004; Li & Yao 2004; Huang et al. 2010; Liu et al. 2015).
During the last few years, we have conducted macrofungal diversity surveys in the Eastern Himalayas and the
Hengduan Mountains, southwestern China, and collected five collections of an interesting Pleurotus. A survey of the
relevant literature and morphological examination of the materials suggested that it should be Pleurotus placentodes
(Berkeley 1852: 104) Sacc. (1887: 359), a taxon originally described from Sikkim, India, which Albertó et al. (2002)
considered to be a synonym of Pleurotus djamor (Fries 1821: 185) Boedijn (1959: 292). Our morphological observations
and molecular phylogenetic results, however, indicated that P. placentodes distinctly differs from P. djamor. In the
present paper, we characterize this half-forgotten species using both morphological and molecular evidence.

LIU ET AL.
138 • Phytotaxa 267 (2) © 2016 Magnolia Press
Materials and methods
Specimens and morphological descriptions
Specimens were collected from both Tibet and Yunnan during summers 2009–2010 in subalpine forests dominated by
Betula, Abies and Picea and are deposited in the Herbarium of Cryptogams, Kunming Institute of Botany of the Chinese
Academy of Sciences, China (HKAS). The macro-morphological characters of fresh specimens were described based
on field notes and photographs. Color codes are according to Kornerup & Wanscher (1981). Microscopic observations
and interpretation of basidiospore data follow Yang (2000). Terminology for descriptive terms follows Bas (1969) and
Albertó et al. (2002). The type of Pleurotus placentodes in the herbarium of the Royal Botanic Gardens, Kew (K) was
examined by one of the authors (EH).
DNA extraction, PCR and sequencing
Genomic DNA was extracted from silica-gel dried or herbarium material using the CTAB method Doyle & Doyle
(1987). The internal transcribed spacer (ITS) region was amplified with primer pair ITS1/ITS4 White et al. (1990)
in an ABI 2720 thermal cycler (Applied Biosystems, Foster City, CA, USA). The PCR program was conducted as
follows: pre-denaturation at 94 °C for 4 min followed by 35 cycles of denaturation at 94 °C for 40 s, annealing at 50
°C for 50 s, elongation at 72 °C for 50 s; afterwards, a final elongation at 72 °C for 8 min was included. PCR products
were purified with the Gel Extraction & PCR Purification Combo Kit (Spin-column, Bioteke, Beijing, China), and
then sequenced on an ABI-3730-XL sequence analyzer (Applied Biosystems, Foster City, CA, USA) using the same
primer combinations as for the PCR. Forward and reverse sequences were assembled and edited with SeqMan (DNA
STAR package; DNAStar Inc., Madison, WI, USA). Sequences retrieved from GenBank and obtained by us in this
study are listed in Table 1.
TABLE 1. Taxa used in molecular phylogenetic analyses and their GenBank accession numbers of ITS sequences
Species Specimens or strains Location GenBank acc. #
H. auriscalpium T-082 United Kingdom EF409725
H. mastrucata T-025 Canada EF409737
P. abieticola TENN52359 Russia AY450348
P. abieticola HKAS45720 China: Tibet KP771696
P. abieticola TENN58284 Russia AF345656
P. albidus Duke327 Brazil AF345658
P. albidus BAFC 50.261 Argentina AF345659
P. australis VT1953 Australia AY315758
P. australis PDD59215 New Zealand AY315761
P. australis PDD87/021XP New Zealand AY315764
P. calyptratus CCRC 36211 - AY265814
P. calyptratus C-1 - JQ837485
P. citrinopileatus TFM-M-E793 - AB115043
P. citrinopileatus HMAS63344 China: Jilin AY696301
P. cornucopiae H-14 - JQ837484
P. cornucopiae TENN55191 Austria AY450341
P. cystidiosus AG55 USA FJ608592
P. cystidiosus IFO30607 Japan AY315778
P. cystidiosus subsp. abalonus CBS80391 China AY315806
P. cystidiosus subsp. abalonus VT2476 USA: Hawaii AY315802
...Continued on next page

PLEUROTUS PLACENTODES Phytotaxa 267 (2) © 2016 Magnolia Press • 139
TABLE 1. (Continued)
Species Specimens or strains Location GenBank acc. #
P. djamor HKAS94069 Sri Lanka KX061788
P. djamor HKAS94070 Sri Lanka KX061789
P. djamor SP445789 Brazil KF280329
P. djamor IB36 Peru KJ831854
P. djamor MEL:2382775 Australia KP012957
P. djamor ECS-0176 Mexico GU722275
P. djamor MTK1189 Kenya KJ754115
P. eryngii HIK154 China: Beijing HM998841
P. eryngii HIK139 Iran: Kordestan HM998837
P. eryngii 32PL Greece FJ904755
“P. cf. eryngii”C24 Iran FJ514570
P. eryngii var. tuoliensis HIK138 Iran: Esfahan HM998836
P. eryngii var. tuoliensis HIK152 China HM998839
P. eryngii var. tuoliensis CCMSSC01433 China: Beijing KP867912
P. fossulatus HIK127 Armenia HM998828
P. fossulatus ATCC 52666 India AY265833
P. nebrodensis HIK125 Italy: Apulia HM998826
P. nebrodensis UPA6 Italy: Madonie Mt. HM998816
P. nebrodensis HIK137 Iran: Kordestan HM998835
P. ostreatus TENN 53662 Austria (epitype) AY854077
P. ostreatus CCMSSC06141 China KP867915
P. ostreatus HKAS84903 Germany KP867913
P. placentodes HKAS57781 China: Yunnan KR827694
P. placentodes HKAS57145 China: Tibet (epitype) KR827693
P. placentodes HKAS94408 China: Yunnan KX061786
P. placentodes HKAS94409 China: Yunnan KX061787
P. placentodes HKAS94410 China: Tibet JQ283967
P. populinus ATCC 90083 - AY368667
P. populinus TENN56749 USA AY450346
P. pulmonarius HMAS76672 China: Heilongjiang AY696299
P. pulmonarius HKAS76382 China KP867916
P. pulmonarius ECS-0158 Mexico GU722283
P. purpureo-olivaceus PDD91632 New Zealand GQ411523
P. purpureo-olivaceus ICMP17077 New Zealand GQ411512
P. tuberregium PtWat Cameroon AF109988
P. tuberregium PTV2 Ghana AF109978
P. tuberregium Pt1 Nigeria AF109983
* Sequences KR827693, KR827694 and KX061786–KX061789 were generated in this study.

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140 • Phytotaxa 267 (2) © 2016 Magnolia Press
Phylogenetic analyses
A total of 55 ITS sequences form the dataset, of which 48 were retrieved from GenBank and 7 are newly generated
sequences from this study. Hohenbuehelia auriscalpium (Maire 1930: 220) Singer (1951: 255) and H. mastrucata
(Fries 1818: 229) Singer (1951: 255) were chosen as outgroup according to the recent phylogenetic studies by Menolli
et al. (2014). The dataset was then aligned using MAFFT v7.130b Katoh & Standley (2013) and manually optimized
in Bioedit v7.0.9 Hall (1999).
Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were applied using RAxML v7.2.6 Stamatakis
(2006) and MrBayes v3.1.2 Ronquist & Huelsenbeck (2003), respectively. For the phylogenetic analysis, GTR + G
was chosen as the best fit model for the dataset by using Mrmodeltest 2.3 Nylander (2004). Statistical supports were
calculated using nonparametric bootstrapping with 1000 replicates in ML analysis. For BI analysis, using selected
models and 4 chains were conducted and stopped until the standard deviation of the split frequencies fell below 0.01
and ESS values >200. Tracer v1.5 (http://tree.bio.ed.ac.uk/software/tracer/) was used to monitor the chain convergence.
Trees were sampled every 100 generations. Subsequently, trees were summarized and statistical supports were obtained
by using the sumt command complemented in MrBayes by discarding the first 25% generations as burn-ins.
FIGURE 1. Phylogenetic analysis of Pleurotus species inferred from Maximum Likelihood (ML) analysis of ITS sequences. Bootstrap
values (ML, > 50%) are shown above or beneath individual branches. Bayesian posterior probabilities (BI, > 0.90) are indicated with thick
branches. Taxonomic position of P. placentodes is emphasized in bold face.

PLEUROTUS PLACENTODES Phytotaxa 267 (2) © 2016 Magnolia Press • 141
Results
Molecular phylogeny
In the alignment of the dataset, 349 characters are constant, while 294 characters are variable, of which 258 are parsimony
informative. The topology of phylogenetic trees based on the dataset generated from ML and BI analyses are almost
identical, while statistical supports show slight differences. Our results indicate that P. placentodes is related to the P.
ostreatus (Jacquin 1774: 3) Kummer (1871: 105) and P. pulmonarius (Fries 1821: 187) Quélet (1872: 113) complex,
including P. fossulatus (Cooke 1888: 121), P. eryngii var. tuoliensis (Mou et al. 1987: 153), P. ostreatus, P. populinus
(Hilber 1993: 61), P. albidus (Berkeley 1843: 633) Pegler (1983: 219), P. pulmonarius and P. abieticola (Petersen &
Hughes 1997: 175) with high statistical support (Bootstrap values 100%, and Bayesian posterior probabilities 1) (Fig.
1). Quotation marks were placed around the names in the cladogram and in the table in case the monophyly of the taxa
with the same name is questionable.
FIGURE 2. Basidiomes of Pleurotus placentodes in situ. a. HKAS 57781; b. HKAS 57145. Bars = 2 cm Photos by B. Feng and G. Wu.

LIU ET AL.
142 • Phytotaxa 267 (2) © 2016 Magnolia Press
Taxonomy
Pleurotus placentodes (Berk.) Sacc., Syll. fung. (Abellini) 5: 359 (1887) Figs. 2–3
Basionym: Agaricus placentodes Berk., Hooker’s J. Bot. Kew Gard. Misc. 4: 104 (1852).
Synonym: Dendrosarcus placentodes (Berk.) Kuntze. Rev. Gen. Plant. 3: 464 (1898).
FIGURE 3. Microscopic characters of Pleurotus placentodes. a. Basidia at different stages of development and subhymenium elements
(HKAS 57781); b. Basidiospores (HKAS 57781); c. Broadly clavate to subcylindrical cheilocystidia (HKAS 57781); d. Mucronate
cheilocystidia (HKAS 94410); e. Pileipellis (HKAS 57781); f. Basidiospores (HKAS 57145); g. Mediostratum of lamellar trama (HKAS
57781). Bars 1, 3, 4, 5 and 7 = 20 µm, 2 and 6 = 10 µm. Drawings by Zhu L. Yang.
Description:—Basidiome medium-sized. Pileus flabelliform to orbiculate, occasionally obovate, 40–70 mm from
point of attachment to margin, 40–80 mm in diam.; surface pale ochre, greyish brown to brownish grey (2A2, 3B2–3),
becoming ochre to biscuit-colored (6E6–8) in dry conditions, glabrous, smooth, faintly innately streaked towards
margin, slightly viscid when wet, finely cracked when dry; margin paler colored, inrolled when young, becoming
straight at maturity; context white (1A1), unchanging, relatively thin (up to 8 mm wide near base). Lamellae decurrent,
crowded to subdistant, narrow (up to 3 mm wide), white (1A1), occasionally branched or anastomosing; lamellar edges
entire and concolorous; lamellulae 2–3 tiers. Stipe lateral, 5–10 × 5–8 mm, subcylindric, whitish to greyish (1A1, 3B3,
4B3). Odor pleasant, orange-like or indistinct; taste mild. Spore print white to cream-colored (1A1, 3A2).
Basidiospores [100/4/4] (5.5) 6–8 (9) × (3.5)4–5 µm, Q = (1.35–) 1.45–1.75 (–1.80) (Q = 1.58 ± 0.10), ellipsoid to
subovoid, sometimes elongate, colorless and hyaline, thin-walled, smooth, inamyloid, non-dextrinoid. Basidia 25–30
× 6.5–7.5 µm, narrowly clavate to clavate, thin-walled, 4-spored; sterigmata 3–4 µm long. Cheilocystidia 15–20 × 3–7
µm, not well-differentiated, clavate to broadly clavate, sometimes nearly cylindrical, occasionally mucronate, colorless
and hyaline, thin-walled. Pleurocystidia absent. Lamellar trama monomitic, composed of subregularly to irregularly
arranged thin- to thick-walled (up to 1 µm diam.), colorless and hyaline, branching, filamentous hyphae 2–8(12) µm wide;
terminal cells 50–100 µm long, non-skeletalized. Subhymenium composed of frequently branching hyphae, 3–5 µm

PLEUROTUS PLACENTODES Phytotaxa 267 (2) © 2016 Magnolia Press • 143
wide. Pileipellis a 20–50 µm thick cutis composed of repent, radially arranged, yellowish to brownish, thin- to slightly
thick-walled (up to 0.5 µm diam.) filamentous hyphae, 3–6(8) µm wide; suprapileipellis slightly gelatinized, often with
coralloid to finger-like hyphal tips, 5–20 × 2–4 µm. Pileitrama monomitic, composed of radially to irregularly arranged
thin- to slightly thick-walled (up to 1 µm diam.), colorless and hyaline, filamentous hyphae, 3–15 µm wide. Stipititrama
composed of more or less irregularly arranged, frequently branching and septate, slightly thick-walled (0.5–1.5 µm
diam.), colorless and hyaline monomitic hyphae, 3–12 µm wide. Clamp connections abundant at all septa.
Habitat and known distribution:—Growing on decaying wood of Betula (reported to be the host substrate of the
type material) and Picea in subalpine forests dominated by Abies, Betula, Picea and Rhododendron; during summer
(June to September) at 3000–4200 m elev., in southwestern China. Originally described from Sikkim, India.
Specimens of Pleurotus placentodes examined: INDIA. Sikkim, on Betula wood, 3353 m elev., 1849, Hooker
f. 16 (K, holotype). CHINA. Yunnan Province, Yulong County, Shitou, 3000 m elev., 5 September 2009, G. Wu 249
(HKAS 57781); Dali City, locality unknown, Y.C. Zhao (HKAS 94408 and 94409, dried strains); Tibet Autonomous
Region, Linzhi County, Sejilashan, 4200 m elev., on Betula sp., 29 Jun. 2009, B. Feng 416 (HKAS 57145, epitype
designated here); same locality, 4100 m elev., 6 August 2010, WSX 2010806 (HKAS 94410).
Specimens of Pleurotus djamor examined: SRI LANKA. Kandy, Mt. Hanthana, 11 IX 2011, S.C. Karunarathna
SK-76 (HKAS 94069); Knuckels Range, 16 IX 2011, S.C. Karunarathna SK-83 (HKAS 94070).
Discussion
Based on the presented molecular data P. placentodes is phylogenetically basal to all other species of the P. ostreatus
- P. pulmonarius complex, including P. fossulatus, P. eryngii, P. nebrodensis, P. eryngii var. tuoliensis, P. ostreatus,
P. populinus, P. albidus, P. pulmonarius and P. abieticola (Fig. 1). The species of this complex are characterized by
monomitic basidiomes (type I of Albertó et al. 2002). It is noteworthy that the basidiospores of the genus Pleurotus are
usually cylindric to elongate, while those of P. placentodes are ellipsoid to subovoid, and therefore not “typical” for
Pleurotus. The mucronate cheilocystidia were not observed in HKAS 57145 and 57781 but were relatively common
in HKAS 94410.
Morphological characters of our collections correspond well both with the description and illustration of P.
placentodes Pegler (1977a) and the holotype material re-examined by one of the authors of the present paper (EH).
Pleurotus placentodes significantly differs from the other taxa in the aforementioned P. ostreatus - P. pulmonarius
complex by its shorter and proportionally wider basidiospores with lower Q and the occurrence in montane forests
dominated by broadleaf trees and conifers. The basidiomes from southwestern China usually possess a short stipe (Fig.
2), while the stipe is absent in the type Pegler (1997a). We treat the length of the stipe as a variable character and should
not be over-weighted.
Several names of Pleurotus, including P. placentodes, were listed as synonyms of P. djamor Albertó et al. (2002),
probably due to the variability in basidiome color and stature of species accommodated in the P. djamor complex
(Corner 1981; Pegler 1986; Nicholl & Petersen 2000). However, P. djamor is a pantropical taxon with cylindrical
basidiospores (Pegler 1977b; Corner 1981; Pegler 1986; Li et al. 2014; cf. also our observation on the collections of P.
djamor from Sri Lanka; Fig. 1), while P. placentodes with ellipsoid to subovoid basidiospores (Berkeley 1852; Pegler
1977a; also confirmed by re-examination of the holotype material) is restricted to alpine to subalpine areas in the
Himalayas at about 3000 m elev. Furthermore, P. djamor possesses a dimitic hyphal system with developing terminal
and tapering skeletal hyphae (Corner 1981; Pegler 1986; type III of Albertó et al. 2002). The results of molecular
phylogenetic analysis presented in this contribution place P. djamor and P. placentodes in two different major clades
(Fig. 1), indicating that they are not closely related.
To our knowledge, although Pegler (1977a) provided a description based on a study of the type collection, P.
placentodes was largely overlooked since its first publication by Berkeley (1852). The holotype of P. placentodes
(India, Sikkim, on Betula wood, 3353 m elev., 1849, Hooker f. 16, K) is now in very poor condition. In order to present
an updated taxonomic species concept, the recently collected specimen (HKAS 57145), is designated here as the
epitype.
Morphologically, Pleurotus placentodes is similar to the sympatric yet ill-defined species P. anserinus Sacc.
(1850: Syll. Fung. 2: 83) which is also reported by J.D. Hooker from Jillapahar, near Darjeeling, Sikkim. However,
the two species can be distinguished by the color of pilei and the shape of the basidiospores (Berkeley 1850; Pegler
1977a).

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144 • Phytotaxa 267 (2) © 2016 Magnolia Press
Acknowledgements
The authors are very grateful to Mr. S.C. Karunarathna (Mae Fah Luang University), Dr. Y.C. Zhao (Yunnan Academy
of Agricultural Sciences), Dr. S.X. Wang (Beijing Academy of Agriculture and Forestry Sciences), Dr. B. Feng and Dr.
G. Wu (Kunming Institute of Botany, Chinese Academy of Sciences) for providing specimens for this study. We also
thank Dr. B. Feng and Dr. G. Wu for providing pictures of P. placentodes. The anonymous reviewers are also gratefully
acknowledged for their comments and suggestions. This study is supported by the National Basic Research Program
of China (973 Program, No. 2014CB138305).
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