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First report of Polyporus ciliatus in Japan, and taxonomic re-evaluation of synonymous species described from Japan

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First report of Polyporus ciliatus in Japan, and 

taxonomic re-evaluation of synonymous species 

described from Japan

Toshinori MATOZAKI

1)

, Tsutomu HATTORI

2)

, Tomohiro KUWAHARA

3)

Sophon BOONLUE

4)

, Nitaro MAEKAWA

5)

, Akira NAKAGIRI

5)

, Naoki ENDO

5)

and Kozue SOTOME

5)

*

1)

The United Graduate School of Agricultural Sciences, Tottori University, 4-101 Koyama, Tottori 680-8553, Japan

2)

Forestry and Forest Products Research Institute, Forest Research and Management Organization, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan

3)

Faculty of Agriculture, Tottori University, 4-101 Koyama, Tottori 680-8553, Japan

4)

Department of Microbiology, Faculty of Science, Khon Kaen University, Khon Kaen, 40002, Thailand

5)

Fungus/Mushroom Resource and Research Center, Faculty of Agriculture, Tottori University, 4-101 Koyama, Tottori 680-8553, Japan

(Received 16 August 2019 / Accepted 4 December 2019)

Abstract

Polyporus ciliatus (Polyporales, Basidiomycota) is reported as a new record to Japan with a new Japanese name (“Ezono-amisugi-take”) based on both morphological and phylogenetic analyses of specimens from Hokkaido. Phylogenetic analyses revealed that P. ciliatus collections from Japan, China and Denmark had identical nuclear large ribosomal subunit sequences and formed a distinct clade with P. longiporus. Examinations of cultural characters revealed that P. ciliatus produce subglobose to fusiform, rarely clavate chlamydospores. We also re-evaluated the holotype of P. saitoi collected from Japan. Polyporus saitoi has been treated as a synonym of P. ciliatus, but the holotype has corky context, a reddish-brown pileus

surface when dry, and larger pores [(3–) 4–5 pores/mm] than P. ciliatus, and it is morphologically distinct from

P. ciliatus. We conclude that the status of P. saitoi as a valid species is dubious name because the holotype lacks microscopic characters besides skeletal-binding hyphae which are insufficient for species identification.

Key words: Lentinus, Polypore, Taxonomy, Type specimens, Wood decay fungi

Regular Paper

Introduction

Polyporus Fr. (Polyporales, Basidiomycota) was considered to contain white-rot fungi having a more or less stipitate basidiocarp, poroid hymenophore, dimitic hyphal system with arboriform skeletal-binding hyphae,

and smooth cylindrical basidiospores1-3). However, recent

evaluations of the genus revealed that Polyporus is polyphyletic, and that Polyporus spp. producing centrally stipitate basidiocarps with stipes lacking a dark crust formed a clade together with species of a lamellate genus, Lentinus Fr., which includes many edible fungi that are

highly valued in South East Asian countries4-8).

Among the species of Polyporus s.l. defined by Núñez

and Ryvarden2, 3), P. ciliatus Fr. is characterized by having

a centrally stipitate basidiocarp, light brown to umber and circular pileus (up to 10 cm diameter) often covered by squamules, light cream to tan pores (5-7 pores/mm), cylindrical stipe (up to 7-8 cm long), dimitic hyphal system with clamped generative hyphae and skeletal hyphae, and allantoid to cylindrical basidiospores (5-7 × 1.5-2 μm)2, 9, 10)

. Polyporus ciliatus is widely distributed

in the temperate regions of Eurasia, including China and Japan3, 11-13)

, where it usually occurs on a wide variety of hardwood species and occasionally on conifer species in the genera Abies Mill., Acer L., Alnus Mill., Betula L.,

Cornus L., Eucalyptus L’Hér., Fagus L., Fraxinus L., Ilex L.,

Juglans L., Populus L., Picea A.Dietr., Pinus L., Prunus L., Salix L., and Tilia L.10, 11)

.

Ryvarden12)

concluded that P. saitoi Lloyd, described from Japan in 1924, was synonymous with P. ciliatus without further comment. Therefore, the record of

P. saitoi documented by Lloyd14)

can be treated as the first

record of P. ciliatus from Japan. Núñez and Ryvarden2)

mentioned the distribution of this fungus in Japan, but

no referable voucher was indicated. Sotome15)

questioned whether P. ciliatus was distributed in Japan.

Recently, a Polyporus fungus resembling P. ciliatus was collected from a cool-temperate area in the northern part of Japan (Sapporo, Hokkaido). In this study, we carried out morphological examinations and phylogenetic analyses based on partial sequences of the internal transcribed spacer (ITS) and nuclear region large subunit (nrLSU) regions of the ribosomal RNA genes to conform the phylogenetic position of the specimen. Morphological comparison of the specimen against an

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authenticated specimen of P. saitoi clarified that P. saitoi and P. ciliatus were distinct species. We herein present a detailed description of P. ciliatus as the first record of this fungus from Japan.

Materials and Methods

1. Morphological characteristics

All fresh basidiocarps were collected in a mushroom workshop convened by the Sapporo Mushroom Club at Mt. Sakurayama in May 2017 and 2019. Fresh basidiocarps were dried after isolation using a dryer (DF612, Yamato Science Co., Ltd., Tokyo, Japan) set at 45℃ for 48 hours. Macroscopic characteristics were described based on fresh and dry specimens. Color descriptions were based on the Munsell System (http:// munsell.com/). Microscopic characters were based on dried specimens, which were sectioned by hand using a sharp razorblade. Sections were mounted in 3% (w/ v) KOH solution after staining with 1% (w/v) phloxine solution or in Melzer’s reagent. The abbreviation IKI- is used for chemical reactions indicating both nonamyloid and nondextrinoid. Measurements and line drawings of microscopic elements were made using a Nikon 80i microscope with a Microscope Zoom Drawing arm (Nikon, Tokyo, Japan). Basidiospore measurements were made from material mounted in Melzer’s reagent. The following abbreviations are used for basidiospore measurements: L = mean basidiospore length, W = mean basidiospore width, R = the length/width ratio of a basidiospore, r = arithmetic mean of R, (n = x/y) = x numbers of basidiospores from y specimens. Herbaria holding specimens are abbreviated according to Holmgren et al.16)

.

Mycelial isolates were established on Malt extract agar media using tissue from basidiocarps or germinate basidiospores. Cultural characters were studied on malt

extract agar plates (1.25% BD BactoTM

Malt Extract, 2.0% BD BactoTM

Agar) at 25℃ and described according to Nobles17)

.

The mycelial growth rate Kr at 25℃ was calculated as

follows: Kr = (R1-R0)/(t1-t0), where R1 = colony radius at

time t1, R0 = colony radius at time t0. Extracellular oxidase

reactions were tested according to Kӓӓrik18)

. All cultures have been deposited at the Fungus/Mushroom Resource and Research Center (FMRC), Faculty of Agriculture, Tottori University, Tottori, Japan.

2. Molecular phylogeny

DNA wa s ex t r a c t e d f r om c u lt u r e d myc el i a

following Hosaka and Castellano19)

or using Maxwell®

16 Tissue DNA Purification Kit (Promega, WI, USA)

with the automatic DNA extraction by the Maxwell®

16 Instrument (Promega). Large subunit (LSU) and ITS regions were amplified and sequenced with the following

primers: ITS4/ITS520)

for ITS; and LR0R/LR5 for nrLSU21)

. Amplifying ITS and nrLSU were performed as described

by Sotome et al.22)

. PCR products were purified using ExoSAP-IT (GE Healthcare, Tokyo, Japan) or MonoFas

DNA Purification Kit (GL Sciences, Tokyo, Japan). DNA

sequences were determined using a BigDye® Terminator

3.1 Cycle Sequencing Kit (Applied Biosystems, CA, USA) with an ABI 3130 DNA sequencer (Applied Biosystems).

The ITS and nrLSU sequences were aligned using

MAFFT v. 7 (2019/May)23)

employing an L-INS-i strategy. Trailing ends of the alignments were trimmed, and

alignments were edited manually in BioEdit v. 724)

when necessary. Dataset congruence was tested using the

incongruence length difference (ILD) test25)

as implemented

in PAUP* 4.0b10 (‘partition-homogeneity test’)26)

. A P-value of < 0.05 was considered statistically significant. After testing for congruence, the individual gene datasets were combined, and phylogenetic analyses were conducted. Phylogenetic analyses of the combined dataset were performed using the maximum likelihood (ML) and

maximum parsimony (MP) methods in MEGA 727)

. Best-fit substitution models for ML were estimated by MEGA 7. MP analysis was performed by treating gaps as missing data. The most parsimonious trees were generated using heuristic searches with 1,000 random addition sequences, with maxtrees set to auto increase and TBR branch swapping. The ML and MP trees were tested statistically

with bootstrap values (BS) from 1,000 replicates28)

. Dentocorticium sulphurellum (Peck) M.J. Larsen & Gilb. and Lopharia cinerascens (Schwein.) G. Cunn. were used as outgroups because these species placed at basal clades corresponding to Polyporaceae in the phylogenetic trees published by Binder et al.29)

and Justo et al.30)

. The aligned dataset and resulting trees were deposited in TreeBASE under the accession number S24603.

Results

1. Phylogenetic analyses

In total, four sequences for the ITS and nrLSU regions were generated and submitted to the DDBJ database (Table 1). Additional ITS and nrLSU sequences for 24 species obtained from publicly accessible databases (Table 1) were selected with reference to the phylogenetic

trees reconstructed by Seelan et al.8)

and Sotome et al.31)

. An ILD test showed no significant incongruence between the ITS and nrLSU datasets (P = 0.14). The total evidence dataset, combining the ITS and nrLSU sequence data, had an aligned length of 1,108 sites including gaps, of which 227 characters were parsimoniously informative. The MP analysis yielded six equally most parsimonious trees of 647 steps (CI = 0.5, RI = 0.6). The ML tree based on the total evidence dataset was constructed using the General Time Reversible evolutionary model with Gamma-distributed rates and Invariable sites (GTR+G+I), which was selected as the best-fit model. The ML analysis of the total evidence dataset resulted in a best scoring tree with a likelihood of InL = -4556.47. The ML and MP trees based on the combined dataset showed consistency in well-supported clades (bootstrap support

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trees, the three collections of Polyporus from Hokkaido and P. ciliatus from Denmark and China formed a distinct clade with strong support (MLBS/MPBS = 100/100) and these placed as a sister clade of P. longiporus Audet, Boulet & Sirard (MLBS/MPBS = 92/95) in the Polyporus- Lentinus clade (Fig. 1).

All sequences of the partial LSU region from the Hokkaido samples were identical to those of P. ciliatus from China (Wei1582) and Denmark (CulTENN10167 SBI 9). The partial ITS sequences from the Hokkaido samples and P. ciliatus from China (Weil 582) were also found to

be identical, whereas the partial ITS sequence similarity between Hokkaido samples and P. ciliatus from Denmark (CulTENN10167 SBI 9) and were 97%.

2. Morphological comparison among P. ciliatus, Hokkaido specimens and the holotype of P. saitoi

Morphological characteristics of P. ciliatus, Hokkaido specimens and the holotype of P. saitoi are summarized in Table 2. All typically produce small- to middle-sized and centrally stipitate basidiocarps with a circular pileus. However, P. ciliatus and the Hokkaido specimens share a similar light brown pileus with cilia at the margin,

Table 1. Species used in phylogenetic analyses, their strain or specimen numbers, and their DDBJ accession numbers

Species Strain (Specimen) nos. DDBJ accession nos.

ITS nrLSU

Amauroderma rugosum ML56 LC412115 AB368061 Daedaleopsis confragosa WD747 LC471201 AB368062 Datronia mollis WD794 AB587623 AB368063 Dentocorticium sulphurellum T609 JN165015 JN164815 Favolus emerici WD2379 AB587628 AB587619 Ganoderma lingzhi WD565 AB462322 AB368068 Lentinus badius (JS0094) KP283478 KP283512 Lentinus crinitus TFB11196 GU207296 AY615978 Lentinus polychrous BCC_29606 AB478882 LC052215 Lentinus sajor-caju (FRI62056) KP283492 KP283509 Lentinus squarrosulus C500W AB478883 LC052216 Lentinus tigrinus MUCL22821 AB478881 AB368072 Lignosus rhinocerotis PEN94 JQ409359 AB368074 Lopharia cinerascens FP105043sp JN165019 JN164813 Neofavolus alveolaris WD2358 AB587624 AB368079 Polyporus arcularius WD2138 AB478874 AB368081 Polyporus brumalis WD2372 AB478877 AB368084 Polyporus ciliatus CulTENN10167 SBI 9 AB070882 AJ487943 Polyporus ciliatus (Wei1582) KU189767 KU189798 Polyporus ciliatus TUFC 1016302

LC4867713

LC4867733

Polyporus ciliatus TUFC 1016312

LC4867723

LC4867743

Polyporus longiporus (DAOM 229479)1

AB478880 LC052217 Polyporus thailandensis (MSUT_6734)1 LC052221 LC052219

Polyporus tricholoma 9591 AB478884 AB368100 Pseudofavolus cucullatus WD2157 AB587637 AB368114 Pycnoporus cinnabarinus WD741 AB735965 AB735945 Trametes orientalis WD1660 AB735966 AB735946

1 Holotypes.

2 Voucher specimens collected from Sapporo, Hokkaido, Japan. 3 Newly generated sequences.

Table 2.Morphological comparisons among P. ciliatus, the Hokkaido specimens, and the holotype of P. saitoi

P. ciliatus9, 10)

Hokkaido specimens P. saitoi (holotype) Basidiocarps Annual, centrally stipitate Annual, centrally stipitate Annual, centrally stipitate Shape and size of

pileus

Circular, convex, margin ciliate, 6-10 cm wide, up to 4-7 mm thick

Circular, convex, margin ciliate, 1.3-5 cm wide, up to 4.5 mm thick

Circular, margin convex, depressed in the center, margin entire, up to 2.5 cm wide, up to 2.5 mm thick Pileus surface Minutely squamulose or glabrous,

light brown, then umber

Minutely squamulose or glabrous, cream to yellowish brown, then dark brown

Glabrous, reddish brown

Stipe surface Fine tomentum, light ochraceous to dirty brownish

Fine tomentum, cream to yellowish brown, then bark brown

Smooth, reddish brown Pores Round, 5-7 pores/mm Round to angular, 5-7 pores/mm Angular, (3-) 4-5 pores/mm Shape and size of

basidiospores

Cylindrical slightly curved,

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Fig. 1. Maximum likelihood tree generated using the General Time Reversible model with Gamma-distributed rates and Invariable sites (GTR+G+I) from the combined ITS and nrLSU sequence dataset. Bootstrap values from maximum likelihood and maximum parsimony analyses (MLBS/MPBS) greater than 50% are indicated above the branches.

Fig. 2. Basidiocarps of Polyporus ciliatus (TUMH 63619). Fig. 3. Micromorphology of Polyporus ciliatus (TUMH 63620). A: Basidiospores. B: Basidia. C: Generative hyphae of the context. D: Skeletal-binding hyphae of the trama. E: Skeletal-binding hyphae of the con-text. F: Hyphal peg. Bars: 10 μm.

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minutely squamules on the pileus surface and a finely tomentose stipe. In addition, P. ciliatus and the Hokkaido specimens also resemble each other in the shape and size of pores (5-7 pores/mm) and basidiospores (about 5-7 ×

2 μm). On the other hand, P. saitoi differs from P. ciliatus

and the Hokkaido specimens by having a glabrous and reddish brown pileus without cilia at the margin, smooth stipes and smaller pores [(3-) 4-5 pores/mm)]. Since the morphological characteristics of the Hokkaido specimens are identical to P. ciliatus (Table 2), we concluded that these taxa are conspecific based on phylogenetic and morphological evidence.

 Taxonomy

Polyporus ciliatus Fr., Observ. Mycol. 1: 123 (1815)

Morphological characters (Figs. 2, 3): Basidiocarps annual, centrally stipitate, solitary. Pileus circular in outline, convex, 1.3-5 cm wide, up to 4.5 mm thick; pileus surface azonate, often minutely squamulose or glabrous, cream to yellowish brown (10YR4-9/4), then dark brown (10YR3/1-2); pileus margin thin, ciliate, slightly incurved or flat. Stipe cylindrical, equal, 1.5- 2.2 cm long, up to 5 mm wide; surface fine tomentum, cream to yellowish brown (10YR4-8/3), then dark brown (10YR3/1-2). Context fleshy when fresh, corky when dry, white when fresh, white to cream (10YR8-9/2-3) when dry, up to 6 mm thick. Pore surface white to cream (10YR9/2-3); pores round to angular, 5-7 pores/mm; dissepiments thin, slightly decurrent. Tubes concolorous with pore surface, up to 1.5 mm deep. Hyphal systems dimitic with generative hyphae and skeletal-binding hyphae. Contextual generative hyphae with clamp

connections, thin-walled, hyaline, IKI-, 2-8 μm in diam,

often inflating to 14 μm in diam. Contextual

skeletal-binding hyphae thick-walled to almost solid, dominant, occasionally branched, moderately interwoven, hyaline,

IKI-, 2-3 μm in diam, often inflating to 10 μm in

diam, with one to three lateral side branches. Tramal generative hyphae with clamp connections, thin-walled,

hyaline, IKI-, 2-3 (-6) μm in diam. Tramal

skeletal-binding hyphae dominant, thick-walled to almost solid, occasionally branched, moderately interwoven, hyaline,

IKI-, 2-3.5 μm in diam. Basidia clavate, (12-) 14-

19 × 4.5-5.5 μm, with four sterigmata. Basidiospores

cylindrical, slightly curved, thin-walled, smooth, (4.5

-) 5-7 (-7.5) × 2-2.5 μm, L = 5.94, W = 2.12, R = 2.29-

3.51, r = 2.81 (n = 90/3). Hyphal pegs sparse, consisting of

fascicle of thin-walled hyphae, up to 25 μm in diam at the

base, to 36 μm projecting from hymenium.

Specimens examined. JAPAN, Hokkaido, Sapporo, Minami-ku, Mt. Sakurayama, leg. the foray of Sapporo Mushroom Club, on bedlog of Lentinula edodes (Berk.) Pegler, 13 May 2017 (TUMH 63619); ibid., leg. the foray of Sapporo Mushroom Club, on decaying branch of a broad-leaved tree, 23 May 2017 (TUMH 63620); ibid., leg. the foray of Sapporo Mushroom Club, on decaying branch of a broad-leaved tree, 23 May 2017 (TUMH 63621); ibid., leg. the foray of Sapporo Mushroom Club, on decaying branch of a broad-leaved tree, 4 May 2019 (TUMH 63622).

Japanese name: Ezono-amisugi-take (newly proposed here)

Cultural characters (Fig. 4): Growth fast, 5.1-8.4 mm/ day, plates covered in one week. Advancing zone even, appressed, white. Mat white, aerial mycelium woolly to cottony. Reverse unchanged. Odor none. Hymenophore development not seen within six weeks. Generative hyphae from advancing zone thin-walled, hyaline, 2-5 (-7) μm in diam, with clamp connections. Generative hyphae from aerial mycelium thin- to thick-walled, hyaline, 2-3.5 μm in diam, with clamp connections. Fiber hyphae present, moderately to frequently branched, hyaline, thick-walled,

up to 3.5 μm in diam. Chlamydospores present, terminal

or intercalary, slightly thick-walled, abundant; intercalary chlamydospores subglobose to fusiform, hyaline, 6-22

× 4.5-12.5 μm; terminal chlamydospores subglobose to

fusiform, 7-18.5 × 4.5-12.5 μm or more rarely clavate,

20.5-25.5 × 7-8.5 μm.

Extracellular peroxidase activities; 1-naphthol, +; tyrosine, -.

Species code17)

: 2, 3, 8, 11, 34, 36, 38, 41, 54.

Cultures examined. TUFC 101630, TUFC 101631, TUFC 101632 and TUFC 101633 isolated from TUMH 63619, TUMH 63620, TUMH 63621 and TUMH 63622, respectively.

Fig. 5. Polyporus saitoi (BPI US0306339, holotype). A: Basidiocarp. B: Pores. Bars: A 1 cm; B 1 mm.

Fig. 4. Cultural characters of Polyporus ciliatus (TUFC 101631). A: Colony on malt extract agar plate at six weeks (upper surface). B: Chlamydospores. Bar: 10 μm.

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Polyporus saitoi Lloyd, Mycol. Writ. 7: 1269 (1924)

Morphological characters (Fig. 5): Basidiocarp annual, centrally stipitate, solitary. Pileus circular in outline, convex, depressed in the center, 2.5 cm wide, up to 2.5 mm thick; pileus surface azonate, glabrous, reddish brown (5YR3-5/3-6); margin incurved, thin, entire. Stipe cylindrical, equal, 2.8 cm long, up to 3 mm wide; surface smooth, reddish brown (10R3-4/3-7). Context hard corky, pale yellowish brown (10YR9/3), up to 1 mm thick. Pore surface yellowish brown (10YR5/6-8); pores angular, (3-) 4-5/mm; dissepiments thin, slightly decurrent. Tubes concolorous with pore surface, up to 1.5 mm deep. Hyphal systems dimitic: Generative hyphae not seen; Skeletal-binding hyphae thick wall to almost

solid, occasionally branched, 2.5-6 μm in diam. Basidia

and basidiospores not seen.

Specimen examined. JAPAN, Hokkaido, Sapporo, on Quercus grosseserrata Blume, leg. A. Saito, 1900 (holotype; BPI US0306339).

Discussion

Previous phylogenetic studies revealed that “Polyporus s.l.” in traditional usage is polyphyletic, and several species have been subsequently segregated from the

genus. Sotome et al. 32)

transferred “Polyporus spp.” with laterally stipitate basidiocarps without a crust on the stipe surface to Favolus Fr. and Neofavolus Sotome & T. Hatt.

Zmitrovich and Kovalenko33)

expanded the circumscrip-tion of Lentinus to accommodate “Polyporus spp.” and “Lentinus spp.” that share centrally stipitate basidiocarps without a crust on the stipe surface. These authors also proposed a new combination, L. substrictus (Bolton) Zmitr. & Kovalenko, for P. ciliatus because the name Lentinus ciliatus was previously validly published for the taxon L.

ciliatus Lév. On the other hand, Seelan et al.8)

preferred to use the genus name Polyporellus P. Karst. instead of

Lentinus as defined by Zmitrovich and Kovalenko33)

. In the present study, we prefer to tentatively retain P. ciliatus and related species, including P. saitoi, in Polyporus because of this nomenclatural confusion.

Morphologically and phylogenetically, the Polyporus species we examined here is identified as P. ciliatus. In contrast, P. saitoi has a hard, corky texture, reddish brown pileus when dry, and larger pores [(3-) 4-5 pores/ mm], which are inconsistent with the character states of P. ciliatus. Therefore, we conclude that the two species warrant recognition as distinct species. Thus, the present report is the first confirmed description of P. ciliatus from Japan. Polyporus saitoi is considered to be a member of the Polyporus-Lentinus clade because the species produces centrally stipitate and middle-sized basidiocarps with a dimitic hyphal system, but its morphological characters are inconsistent with any described species of this clade6, 31)

. Unfortunately, as noted in the original description of

P. saitoi by Lloyd14)

, basidiospores were not seen in the holotype and our present morphological observations

are not able to provide further microscopic characters for identification. We prefer to retain P. saitoi as a dubious name because its original description and holotype are insufficient for identification.

In Europe, basidiocarps of P. ciliatus usually appear

in late spring or summer from April to July2)

. All Japanese specimens of P. ciliatus used in the present study were collected in late spring in May in a cool-temperate area of Hokkaido. Based on morphological comparisons between European and Japanese P. ciliatus (Table 2), Japanese specimens have smaller and lighter colored pileus than those of Europe. These morphological differences may imply the presence of geographic variations within this species, but further critical examinations based on more Japanese specimens are necessary.

The present phylogenetic tree based on ITS and nrLSU sequence data indicated that P. longiporus is the closest relative of P. ciliatus, which is consistent with

previous studies by Sotome et al.6, 31)

. Both species are commonly found in May in cool-temperate areas of Hokkaido and produce similar middle-sized and centrally

stipitate basidiocarps with a convex pileus6)

. However, P. longiporus is distinguished from P. ciliatus by the glabrous, dark gray to almost black surface of the pileus, radially elongated pores (0.5-2 × 0.1-0.5 mm), and allantoid to cylindrical basidiospores [6.3-8.6 × 1.8-2.3 (-2.7) μm]6)

. Polyporus ciliatus is morphologically similar to P. brumalis (Pers.) Fr. in producing centrally stipitate basidiocarps, pileus often ciliate at the margin, angular pores, and similar-sized basidiospores (6-7 × 2-2.5 μm)2, 9, 10)

, but P. brumalis is distinguished from P. ciliatus in having a reddish brown to dark brown pileus surface

and larger pores [(1-) 3-4 pores/mm]2, 9, 10)

In the current study, we found that P. ciliatus produces subglobose to fusiform, or clavate chlamydospores (6-

22 × 4.5-12.5 or 20.5-25.5 × 7-8.5 μm) in culture on

malt agar (Fig. 4B), which is consistent with a number of related species in Polyporus and Lentinus [P. arcularius (Batsch) Fr., P. brumalis, and L. tigrinus (Bull.) Fr.

according to Nobles17)

and Stalpers34)

]. Maekawa35)

surveyed the productivity of anamorphs in cultures of 1,415 species of Agaricales (442 genera, 21 orders) and suggested that it is of value for taxonomic discrimination at the family, genus, and species ranks in the Agaricales. Further research focusing on the cultural characteristics of species allied to P. ciliatus may reveal any correlation between the clamydospore production and the molecular phylogeny of these taxa.

Acknowledgments We would like to express our thanks to the curator of U.S. National Fungus Collections, USDA-ARS (BPI) for the loan of the holotype. We also thank Mr. S. Habu and other participants in the mushroom workshop at Sakurayama by the Sapporo Mushroom Club for their kind provision of the specimens and photos. This research was partially supported

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by a JSPS Grant-in-Aid for Young Scientists (B) (Grant Number 15K18596 to KS), and a fund administered by the Institute for Fermentation, Osaka (to NM).

和 文 摘 要

Polyporus ciliatus の日本初報告 , 及び日本から

記載された異名種の分類学的再検討

的崎利規1) ・服部 力2) ・桑原知弘3) Sophon BOONLUE4) ・前川二太郎5) ・中桐 昭5) 遠藤直樹5) ・早乙女梢5) * 1) 鳥取大学大学院連合農学研究科  〒680-8553 鳥取県鳥取市湖山町南 4-101 2) 国立研究開発法人森林研究・ 整備機構森林総合研究所  〒305-8687 茨城県つくば市松の里 1 3) 鳥取大学農学部  〒680-8553 鳥取県鳥取市湖山町南 4-101 4)

Department of Microbiology, Faculty of Science, Khon Kaen University, Khon Kaen, 40002, Thailand

5)

鳥取大学農学部附属菌類きのこ遺伝資源センター  〒680-8553 鳥取県鳥取市湖山町南 4-101

北海道産 Polyporus 属種(Polyporales,Basidiomycota)

を形態学的,及び系統学的に検討し,日本新産種として P. ciliatus(新称:エゾノアミスギタケ)と同定した.これ らの nrLSU 領域の塩基配列は中国産およびデンマーク産 本種と同一で,系統樹上では,これらはと同じクレードを形 成し,さらに,本種は P. longiporus と最も近縁であった.本 種の培養性状を調査した結果,P. ciliatus は培地上に類球形 ~紡錘形,希に棍棒形の厚壁胞子を形成した.日本産標本に 基づき新種記載された P. saitoi の基準標本を再検討した結 果,基準標本の傘肉はコルク質で,傘表面は赤褐色,また孔 口は P. ciliatus と比較して大型 [(3-) 4-5 pores/mm] で あった.P. saitoi は,従来 P. ciliatus の異名として扱われて きたが,両者は形態学的に明らかに異なっていた.P. saitoi の基準標本からは骨格結合菌糸以外に種判別に重要な担子 胞子等が確認できないこと , 新種記載以降 P. saitoi の報告 例が無いことから,本種を疑問名と判断した. References

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Fig. 1.  Maximum likelihood tree generated using the General Time Reversible model with Gamma-distributed  rates and Invariable sites (GTR+G+I) from the combined ITS and nrLSU sequence dataset
Fig. 4. Cultural characters of Polyporus ciliatus (TUFC 101631). A:

参照

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