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Botanical Studies (2009) 50: 451-458.
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MICROBIOLOGY
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Ganoderma multipileum, the correct name for 'G. lucidunf in tropical Asia
Dong-Mei WANG1,2, Sheng-Hua WU3 *, Ching-Hua SU4, Jin-Torng PENG5, Ya-Hui SHIH3, and Lung-Chung CHEN6
1Guangdong Provincial Key Laboratory of Microbial Culture Collection and Application, Guangdong Institute of Microbiology, Guangzhou 510070, P.R. China
2Systematic Mycology and Lichenology Laboratory, Institute ofMicrobiology, Chinese Academy of Sciences, Beijimg
100101, P.R. China
3Department of Botany, National Museum of Natural Science, Taichung 404, Taiwan
4Department of Microbiology and Immunology, School of Medicine, Taipei Medical University, Taipei 110, Taiwan 5Agricultural Research Institute, Wu-feng, Taichung 413, Taiwan 6Department of Plant Pathology, National Chung Hsing University, Taichung 402, Taiwan (Received April 28, 2008; Accepted April 17, 2009)
ABSTRACT. Ganoderma lucidum (generic type), originally described from England, has been reported worldwide. In the Orient, an economically important fungus commonly known as 'ling-zhi' or 'chi-zhi' for more than 2000 years, has also been named G. lucidum. However, the identity of the Oriental fungus has been questioned in recent years. Earlier molecular studies suggested that G. lucidum sensu stricto may be restricted to Europe and that G. lucidum ' in Asia consists of at least two distinct species, one represented by material from mainland China and one by tropical Asian collections. This study attempts to clarify the identity of 'G. lucidum' in tropical Asia, with special emphasis on wild collections from Taiwan. The internal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) of related species and strains were sequenced, and phylogenetic analyses were conducted. The results confirmed that G. lucidum is a name mistakenly applied to Oriental collections. The forgotten name Ganoderma multipileum, presented over half a century ago from Taiwan, was found in this study as the earliest valid name for 'G. lucidum' known from tropical Asia.
Keywords: Forgotten name; Ganodermataceae; Molecular phylogeny; Taxonomy.
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INTRODUCTION
Ganoderma lucidum (Curtis) P. Karst., the type of Ganoderma P. Karst., was originally described based on a specimen collected from Peckham, London, UK. Currently, this species has been reported worldwide, i.e. Europe (Ryvarden and Gilbertson, 1993), Asia (Nunez and Ryvarden, 2000; Zhao and Zhang, 2000), Oceania (McKenzie and Foggo, 1989), Africa (Ryvarden and Johansen, 1980), and America (Bazzalo and Wright, 1982; Gilbertson and Ryvarden, 1986). In the Orient, a fungus commonly known as 'ling-zhi' or 'chi-zhi' for more than 2000 years, has also been named G. lucidum. This fungus is widely grown on a commercial scale for its medicinal properties. However, the identity of this important fungus has been questioned in recent years. Pegler and Yao (1996) mentioned that the morphology of ' G. lucidum' in the Orient differs from that of G. lucidum found in Britain and throughout Europe in having basidiocarps of a more-slender stature. Moncalvo et al. (1995a, b) concluded that
Corresponding authors: E-mail: shwu@mail.nmns.edu.tw; Tel: +886-4-23226940 ext. 503; Fax: +886-4-23258684.
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collections of G. lucidum from Asia and Europe belong to different species based on the analysis of DNA sequences derived from the internal transcribed spacer (ITS) and partial nuclear large subunit ribosomal DNA (LSU nrDNA) regions. Furthermore, collections of Oriental ' G. lucidum' from mainland China and Taiwan were separated into two clades in the molecular analyses of Moncalvo et al. (1995a, b). This suggests that at least two species are included. The correct respective names for them have yet to be determined.
Ganoderma lucidum is a temperate species, so far known with certainty from Europe only. In Taiwan, however, the fungus identified by this name is common in lowland tropical and subtropical belts. To clarify the identity of ' G. lucidum' in tropical Asia, we performed a detailed investigation of Ganoderma species in Taiwan using both morphological and molecular characters, with special emphasis on the fungus called G. lucidum. The ITS region, a gene marker useful in separating related species and strains of Ganoderma (Moncalvo et al., 1995a; Smith and Sivasithamparam, 2000) was sequenced for the purpose of a molecular analysis. This study confirms that G.
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lucidum is a name mistakenly applied to Oriental collections. To our knowledge, G. multipileum Hou, presented over half a century ago (Hou, 1950), represents the earliest valid name published for ' G. lucidum' in tropical Asia.
MATERIALS AND METHODS
Fungal materials and morphological observations
All studied specimens are deposited in the herbaria of the National Mu seum of Natural Science (TNM), Taichung, Taiwan, the National Taiwan Museum (TAIM), Taipei, Taiwan, the Forestry and Forest Products Research Institute (TF), Ibaraki, Japan, or in the National Museum of Nature and Science (TNS), Tokyo, Japan. Thin freehand sections from basidiocarps were cut and mounted in 5% KOH to ensure rehydration, and in Melzer's reagent to test for amyloid or dextrinoid reactions. At least 20 basidiospores were measured for each mature specimen. In the descriptions, basidiospore sizes are given both with and without the myxosporium, but only those with the myxosporium were used for comparisons among species. Line drawings of the basidiospores and cuticle cells were made with the assistance of a camera lucida.
DNA extraction, PCR amplification, DNA cloning and sequencing
The details of the molecular experiments, except for DNA cloning, were described in Wu et al. (2007). The primer pairs, ITS1/4 and ITS5/4 (White et al., 1990), were used in this study. For strains with intragenomic ITS heterogeneity, DNA cloning was performed using a yT&A cloning vector and competent ECOSTM 9-5 cells (Yeast-ern Biotech, Taipei, Taiwan). A single positive colony was picked for PCR amplification and DNA sequencing.
Sequence alignment and phylogenetic analysis
Sequences obtained from this study were compared with all Ganoderma sequences retrieved from GenBank and with those published by Smith and Sivasithamparam (2000, the sequences of which have not been deposited in GenBank and were retyped onto computer from the original publication) using Clustal X 1.83 (Thompson et al., 1997). After initial analyses of this broad sample, 29 sequences were chosen that include strains labeled ' G. lucidum' and closely related taxa, representative lineages of Ganoderma species, and two outgroup taxa of Am-auroderma Murrill and Tomophagus Murrill (Table 1). These selected sequences were realigned in Clustal X and adjusted by hand using BioEdit 7.0.4.1 (Hall, 1999). The optimized alignment (deposited in Treebase, accession no.: SN4381) was used for maximum-parsimony (MP) analysis in PAUP* 4.0b10 (Swofford, 2002). The analytical
parameter preferences are described in Wu et al. (2007). Bootstrap analysis (Felsenstein, 1985) was performed with 1000 replicates with random addition sequences to obtain estimates of the reliability of the nodes.
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RESULTS
Phylogenetic analysis
Amplification of the ITS region yielded PCR products of approx. 650 bp long. The final alignment of the 29 sequences included 673 positions. After exclusion of the conserved 5.8S region and ambiguous sites at both ends, 413 sites were used for the MP analysis. Sixteen most parsimonious trees (281 steps, Consistency Index (CI)= 0.662, Retention Index (RI) = 0.773) were retained in this analysis. Of the 413 included sites, 271 were constant, and 46 were variable, but parsimoniously was uninformative, and 96 were parsimoniously informative.
The topologies of the 16 trees were generally identical. One of them is presented in Figure 2. In this tree, collections labeled G. lucidum were separated into three distinct clades, and G. tropicum from tropical Asia formed another separate clade. These four clades are labeled A, B, C, and D in Figure 2.
Clade A is composed of six strains originally determined as G. lucidum, with three from Taiwan, two from India, and one from the Philippines. Changes across the sequences consisted of none to three substitutions. This clade received 83% support in the bootstrap analysis. Ganoderma steyaertanum, a recently presented species based on collections from Indonesia and Australia (Smith and Sivasithamparam, 2003), served as the sister group of clade A, and they were grouped together with high bootstrap support (90%).
Clade B represents ' G. lucidum' from mainland China and Japan, which was robustly supported in the bootstrap analysis (100%). There was no change or only a one-step change between the sequences of the strains. The relationship between the strains was not resolved.
Ganoderma tropicum from Taiwan formed clade C. This clade was very strongly supported by the bootstrap analysis (100%). Changes among the sequences of the strains consisted of none to three steps. It was clearly separated from clade A, where ' G. lucidum' from tropical Asia was seated.
Clade D, containing G. lucidum from Europe, the type locality, consisted of three strains respectively from the UK, France, and Norway. Changes among those sequences consisted of none to six steps. This clade was supported by a very high bootstrap value (99%). It was distantly related to the collections named G. lucidum in Asia.
Taxonomical description
Ganoderma multipileum Hou [as 'multipilea'], Q. J.
Taiwan Mus. 3: 101. 1950. Figure 1A-E
Basidiocarp annual, mostly stipitate, rarely sessile or only with a short base, corky. Pileus 1.5-9 x 2.8-16.5 cm, up to 2 cm thick, flabellate, reniform, dimidiate, sometimes with pilei growing from the lower pilei or growing together, up to 36 cm long and 54 cm wide in total; upper surface orange-yellow, orange-red to red-brown, weakly
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Figure 1. Morphology of Ganoderma spp. A-E, Ganoderma multipileum. (A) Basidiocarps (holotype); (B) Sections of cutis (holotype); (C) Basidiospores (holotype); (D) Basidiocarps (TNM F0012903); (E) Basidiocarps (TNM F0020891). F-H. Ganoderma tropicum (Wu 0407-2). (F) Basidiocarps; (G) Sections of cutis; (H) Basidiospores. Bars = 1 cm in A, E, and F; = 5 cm in D; = 10 (im in
B, C, G, and H.
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Table 1. Taxa used in this study, along with their strain/specimen numbers, origins, and GenBank accession numbers.
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aTaxa in bold indicate sequences from this study.
bTaxa without sequence accession nos. are those whose sequences published in Smith & Sivasithamparam (2000), but have not been submitted to GenBank.
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to strongly laccate, conspicuously sulcate or not, rugose or not; margin obtuse or not, white to orange-yellow. Pore surface cream when young, becoming straw or pale brown with age; tubes up to 1.1 cm long, pale brown or brown; pores circular or subcircular, 6-8 per mm, 60-220 [im in diam., dissepiments 25-110 [im thick. Stipe when present, 1-9 x 0.4-3 cm, flattened or subcylindrical, lateral, dorso-lateral or horizontally lateral, orange-yellow to red-brown, purple-black, strongly laccate or not. Context 0.1-1.8 cm thick, yellow-brown to dark brown, sometimes with melanoid substances, corky; generative hyphae 2-5 [im in diam., colorless, thin-walled, with clamp-connexions; skeletal hyphae 4-7 [im in diam., yellow-brown to red-brown in KOH, dextrinoid; binding hyphae of bovista-type, 1.2-2 [m in diam., few, colorless, thick-
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walled, much-branched. Basidiospores 8-13.5 x 5.5-7.5 [im (with myxosporium), 6.5-10.5 x 4.5-6.5 [im (without myxosporium), ovoid to ellipsoid, mostly truncate, brown, with a dark-brown eusporium bearing fine but slightly conspicuous echinulae, walls 0.5-0.8 [im thick. Cutis composed of clavate cells, 15-60 x 4-12.5 [im, dextrinoid to slightly or strongly amyloid.
Specimens examined. TAIWAN. TAICHUNG: By Taichung Park, on stump, 7 Sep 1 949, Y.F. Yu
(TAIMF000001-holotype of G. multipileum); National
Chunghsing University, 24°07' N, 120°41' E, on trunk of Delonix regia, 5 Sep 2003, M.C. Fan and S.A. Liu, FL0309-1 (TNM F0015492); National Museum of Natural
Science, 24°10' N, 120°39' E, elev. 100 m, on trunk base of D. regia, 23 Sep 1995, W.N. Chou, CWN 01269 (TNM
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F0004355); Shalu, on rotten wood, 17 Nov 2005, H.J.
Hou, WAN 1077 (TNM F0019380); Tatushan, 24°06' N, 120°38' E, elev. 100 m, on wood, 7 Sep 1995, T.T. Yuan, CWN 01246 (TNM F0004332); Veterans General Hospital, on dead trunk base, 9 Jun 2000, T.L. Li, CWN 04670 (TNM F0014622); Veterans General Hospital, on dead
trunk base of Sterculia nobilis, 11 Aug 2000, T.L. Li, CWN 04782 (TNM F0014624). NANTOU: Fruiting body
cultivated from BCRC 37033 in June 2006 in this study
(TNM F0020892; BCRC 37033 = TARI 88-1-30, collected from Chunghsinghsintsun). CHIAYI: On Acacia confusa, Jul 1916, K. Sasaki, 201.962a-c (TNS-F-201962 with 4 basidiocarps, 201.962a-d; basidiocarp 201.962d redetermined as G. tropicum (Jungh.) Bres. in this study); Shuishang, Kuohsing, on stump of Leguminosae sp., Oct 1986, P.H. Fu (TNM F0012903). PINGTUNG: Paoli, 22°04' N, 120°45' E, elev. 160 m, on rotten trunk base of
A. confusa, 6 Sep 1996, W.N. Chou, CWN 01740 (TNM
F0005258). INDIA. Fruiting body cultivated from BCRC
36123 in Jun 2006 in this study (TNM F0020891; BCRC 36123 = ATCC 32471 = FRI 55, collected from roots of Acrocarpus fraxinifolius).
DISCUSSION
Collections identified as ' G. lucidum' tested in this molecular analysis were separated into three clades: A,
B, and D (Figure 2). The Asian collections assigned to G. lucidum (clades A and B) were distantly related to G. lucidum from Europe, the type locality (clade D). This suggests that G. lucidum is a name mistakenly applied to Asian collections. Furthermore, ' G. lucidum' in Asia represents at least two distinct species, one from Taiwan, India and the Philippines, and another from mainland C hina and Japan. T his study is consistent with the conclusions of Moncalvo et al. (1995a, b).
Ganoderma multipileum, a species originally known only based on the type specimen from lowland Taiwan, morphologically resembles Taiwanese ' G. lucidum.' This species was originally described mainly based on three features: (1) two kinds of pilei, one from the stalk with some of the stipes and pilei growing together, and the other growing from the lower pilei; (2) a thin crust, composed of enlarged and bulbous ends of hyphae, 16.5 x 2.35-6.05 [ m; and (3) basidiospores 8.2-9.4 x 4.7 [ m, ovoid, truncate, with numerous and minute echinulae. The former two characteristics are less useful for circumscription of G. multipileum. The feature of multiple-pileate basidiocarp was proven to be unreliable in a cultivation study of G. multipileum by Chang (1983). The presence of enlarged hyphal ends in the crust is not diagnostic because this feature is common to all laccate species of Ganoderma. Hence, only the feature of basidiospores is meaningful in identification. The holotype of G. mutlipileum shares all of the fundamental morphological characteristics with collections of ' G. lucidum' from Taiwan, including the feature of basidiospores with fine echinulae. Their conspe-cific status was confirmed in this study.
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£a. lucidum XrS745aX7e766 G. (ucidtmt Xfl735iaxe73si G. iucidum BCRC37033 G. kJCiOum BCRC36123 Q. fttf^rm BCRC37043 G.hJddum CWN01710 G. stsyaertanum DAR 73780 G. steyaertanum DAR 73779 G. stayastianum QFRIS$47.1 Q. iucidum WD-5G5
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A
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C. mufiipHoum
(tropical fi&a)
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G. iucidum
G. iucidum X87354iX87364
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B 'G. fuciflvrrt'
(mainland China andJapan)
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G. iudCfi/m Z37050SZ37074 G philippii AJ60&714&AJ608715 G. f/ep^J Z37066&Z3708fl
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tropicum
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C
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G. tropicum BCRC37122
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G. tropicum Wu0407'3 Typel G. resinaceum G wsherianum
G subamboinanso var. fd^^ispOfum X7S736&X78757 G. fucitium Z37049&Z37tm
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G. /ucldum
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D
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G. fvckium Z37096&Z37073
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G./yC^fli AF094511AAF£H4490 G. incrass^tum 0AR73763 G. auslr^fe AJ657S90&AJ627591 G. sdspersum X7g742&X7&763 G. iobatum X7S740SX7&761
Amaurodsfwa rude vsi.intefmedfi/fn X767&3&X78774
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Tomophagus (Qanodarrna) colossus Z37071SZ37O91
Figure 2. One of the 16 most parsimonious trees derived from the ITS sequence data. The upper and lower numerals at the nodes denote the number of estimated substitutions and proportions of bootstrap replicates, respectively. Only bootstrap values > 50 % are shown.
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According to Moncalvo and Ryvarden (1997), nearly ninety names of laccate Ganoderma have been proposed from Asian and Pacific regions. Over 90% of these names were excluded as an older name than G. multipileum for the same species, due to their later publications, or representing different species as referring to studies of types or authentic specimens by the present authors (unpublished) or by other researchers (e.g. Steyaert, 1972, 1980; Ryvarden, 1977, 1983, 1985, 1990). The remaining taxa lack modern descriptions or require neotypification (see Moncalvo and Ryvarden, 1997). So far, G. multipile-um is the earliest valid name we can find for 'G. lucidum'' from tropical Asia, and is suggested herein as the correct name for this tropical fungus.
Chang and Chen (1986) stated that the culture from a specimen with multiple pilea, characteristic of G. mul-tipileum, is compatible with that of the Taiwanese ' G. lucidum' without multiple pilea; and these cultures were also compatible with two Indian isolates determined as G.
lucidum (ATCC 32471 and ATCC 32472). Therefore, they
concluded that G. multipileum is a synonym of G. lucidum. However, those two Indian isolates were grouped together with materials of G. multiplileum (clade A), and clearly separated from European G. lucidum (clade D) in the ITS
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phylogenetic tree (Figure 2). Further, the morphology of cultivated fruiting bodies from one of these two Indian strains, BCRC 36123 (= ATCC 32471) is very similar to that of G. multipileum, and the preference for Legumino-sae plants of Indian ' G. lucidum' (Sankaran et al., 2005) is also consistent with that of G. multipileum. In reality, those two isolates from India belongs to G. multipileum.
The material of ' G. lucidum ' from the Philippines (JMM P93.1) is identical to those of G. multipileum from Taiwan in ITS sequence except for one or two single-base substitutions, and embedded in the clade A where Taiwanese G. multipileum was nested (Figure 2). We re-determined this Philippine strain as G. multipileum owing to their high ITS similarity.
Some commercially grown strains of ' G. lucidum' in the
Orient, e.g. RSH RZ and RSH BLC, though not included
in this study, were also re-determined as G. multipileum because they were grouped together with strains ATCC 32471 and JMM P93.1 in previous studies (Moncalvo et
al., 1995a; Hseu et al., 1996).
Smith and Sivasithamparam (2003) described a new species, G. steyaertanum from Australian and Indonesian collections, which has been commonly mistaken for G. lucidum. They stated that a previous ITS sequence analysis by Smith and Sivasithamparam (2000) indicates the allopatric speciation of G. steyaertanum from a species from India and the Philippines, respectively, based on
ATCC 32471 and JMM P93.1 (both determined as G.
multipileum in this study). Furthermore, they concluded that the distribution of G. steyaertanum may not extend much further north of Indonesia. However, the morphology and hosts of G. steyaertanum (Smith and Sivasi-thamparam, 2003) are generally consistent with those of G. multipileum. The ITS sequences of these two species also exhibit high similarity. Although they formed two clusters in the phylogenetic tree (Figure 2), they were grouped together with strong bootstrap support (90%). The relationship between G. steyaertanum and G. multipileum merits further study.
' Ganoderma lucidum' from mainland China and Japan formed a distinct clade (clade B) separate from clade A where G. multipileum was nested (Figure 2). The morphology of ' Ganoderma lucidum' from Japan examined in this study differs from G. multipileum by having moderately echinulate basidiospores and a paler context. Currently, the true identity of this fungus in clade B is still unknown.
Ganoderma tropicum is another species widely distributed in lowland tropical Asia. It resembles G. multi-pileum in having a similar habitat and morphology (Figure 1F, and G). Ganoderma tropicum differs from G. multipi-leum in having strongly echinulate basidiospores (Figure 1H). These two species were also separated from each other in the molecular analysis (clades A and C, Figure 2).
Acknowledgments. We are grateful to the curator of TAIM for arranging the study of the holotype of G. multi-
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pileum, Dr. Tsuyoshi Hosoya (TNS) for the loan of a specimen of ' G. lucidum' collected from Chiayi, Taiwan and Dr. Tsutomu Hattori (TFM) for arranging the loan of two specimens of ' G. lucidum' collected from Ibaraki, Japan. We thank Ms. Shih-Yi Yu for her assistance in obtaining DNA sequence data for this study. This project was supported by the National Museum of Natural Science (ROC), Foundation of the National Museum of Natural Science (ROC), and a Postdoctoral Fellowship Grant of the National Science Council (ROC), no. NSC96-2816-B-178-001,
to the senior author. The senior author also appreciates the support of a project grant (30270006) from the National Nature Science Foundation of China for her initial Gano-derma work in mainland China through Dr. Y.-J. Yao.
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重傘靈芝(Ganoderma multipileum)熱帶亞洲赤芝 “G. lucidum” 的正確學名
王冬梅1,2 吳聲華3 蘇慶華4 彭金騰5 石雅惠3 陳隆鐘6
1廣東省微生物研究所廣東省菌種保藏與應用重點實驗室
2中國科學院微生物研究所真菌地衣系統學實驗室
3國立自然科學博物館植物學組
4台北醫學大學醫學系微生物與免疫學科
5台中農業試驗所
6國立中興大學植病學系
Ganoderma Ac'dww (靈芝屬之模式種）最早描述自英國，目前世界各地均有報導分布。兩千多年來
在東方被稱為「靈芝」或「赤芝」的重要經濟真菌也被鑑定為 G. lucidum 。然而，近年來這個東方赤芝 真菌的種類鑑定受到質疑。早些年的分子研究可看出狹義 G. lucidum 可能僅局限於歐洲，而亞洲的 “ G. lucidum”至少包括兩個種：中國大陸種和熱帶亞洲種，均非G. lucidum 。本文旨在澄清熱帶亞洲赤芝的 種類鑑別，且特別納入台灣的野生標本進行研究。定序相關種類和菌株的核核糖體內轉錄間隔區（ITS region)，運行系統發育分析。結果證實G. lucidum乃為東方赤芝真菌的錯誤命名。本研究並發現半世紀 前發表於台灣而被遺忘的「重傘靈芝」(G. multipileum)，是熱帶亞洲地區赤芝所最早提出的有效名稱。 |
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關鍵詞：被遺忘之名稱；靈芝科；分子系統學；分類學。
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