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1 Ganoderma (Ganodermataceae, Basidiomycota) Species from the Greater Mekong Ganoderma (Ganodermataceae, Basidiomycota) species from the Greater Mekong Subregion Thatsanee Luangharn ( [email protected] ) Mae Fah Luang University https://orcid.org/0000-0002-1684-6735 Samantha C Karunarathna Institute of Botany Chinese Academy of Sciences Arun Kumar Dutta West Bengal State Soumitra Paloi West Bengal State University Cin Khan Lian Yangon Le Thanh Huyen Hanoi University Hoang ND Pham Applied Biotechnology Institute Kevin David Hyde Mae Fah Luang University Jianchu Xu ( [email protected] ) Institute of Botany Chinese Academy of Sciences Peter E Mortimer ( [email protected] ) Institute of Botany Chinese Academy of Sciences Research Keywords: 2 new species, Biogeography, Ecological aspects, Lingzhi, Medicinal mushroom, Morphology Posted Date: July 24th, 2020 DOI: https://doi.org/10.21203/rs.3.rs-45287/v1 License: This work is licensed under a Creative Commons Attribution 4.0 International License. Read Full License 1 Ganoderma (Ganodermataceae, Basidiomycota) species from the Greater Mekong 2 Subregion 3 4 Thatsanee Luangharn1,2,3,4,5, Samantha C. Karunarathna1,3,4, Arun Kumar Dutta6, Soumitra 5 Paloi6, Cin Khan Lian8, Le Thanh Huyen9, Hoang ND Pham10, Kevin D. Hyde3,5,7, 6 Jianchu Xu1,3,4*, Peter E. Mortimer1,4* 7 8 1CAS Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute 9 of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China 10 2University of Chinese Academy of Sciences, Beijing 100049, China 11 3East and Central Asia Regional Office, World Agroforestry Centre (ICRAF), Kunming 12 650201, Yunnan, China 13 4Centre for Mountain Futures (CMF), Kunming Institute of Botany, Kunming 650201, 14 Yunnan, China 15 5Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, 16 Thailand 17 6Department of Botany, West Bengal State University, Barasat, North-24-Parganas, PIN- 18 700126, West Bengal, India 19 7Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu 20 District, Guangzhou 510225, P.R. China 21 8Ar Yone Oo social development association, Yangon, Myanmar 22 9Faculty of Environment, Hanoi University of Natural Resources and Environment, Hanoi, 23 Vietnam 24 10Applied Biotechnology Institute, Van Dat Tower, CN8 Street, Tan Binh Industrial Park, Tan 25 Binh District, Ho Chi Minh City, 700000, Vietnam 1 26 *Correspondence: [email protected]; [email protected] 27 28 Abstract 29 The cosmopolitan Ganoderma is an important pathogen on arboreal plant hosts. Ganoderma 30 is particularly diverse in tropical and temperate regions. It has long been used as traditional 31 medicine because of its beneficial medicinal properties and chemical constituents. In this 32 study, Ganoderma collections were made in tropical regions of Laos, Myanmar, Thailand, 33 Vietnam and temperate Yunnan Province, China. The specimens are described based on 34 micro-macro characteristics and phylogenetic analysis of combined ITS, LSU, TEF1α, and 35 RPB2 sequence data. In this comprehensive study, we report 22 Ganoderma species from 36 temperate and tropical regions of the Greater Mekong Subregion (GMS), namely G. 37 adspersum, G. applanatum, G. australe, G. calidophilum, G. ellipsoideum, G. flexipes, G. 38 gibbosum, G. heohnelianum, G. hochiminhense, G. leucocontextum, G. lingzhi, G. lucidum, G. 39 multiplicatum, G. multipileum, G. myanmarense, G. orbiforme, G. philippii, G. resinaceum, G. 40 sinense, G. subresinosum, G. williamsianum, and G. tsugae. Of these species, 12 were 41 collected from Yunnan Province, China; three species, were collected from Laos; three 42 species, 2 new records, and one new species were collected from Myanmar; 15 species, and 43 four new records were collected from Thailand; and one new species was collected from 44 Vietnam. Comprehensive descriptions, color photographs of macro and micro characteristics, 45 the distribution of Ganoderma in the GMS and worldwide, and a phylogenetic tree showing 46 the placement of all the Ganoderma reported from the GMS are provided. 47 48 Keywords: 2 new species, Biogeography, Ecological aspects, Lingzhi, Medicinal mushroom, 49 Morphology 50 2 51 INTRODUCTION 52 Ganoderma was established by Karsten (1881) with Polyporus lucidus (Curtis) Fr. (= 53 Ganoderma lucidum (Curtis) P. Karst.) as the type species (Moncalvo and Ryvarden 1997). 54 The genus has a worldwide distribution but is predominantly found in the tropical and 55 temperate regions, including Africa, America, Europe, and Asia (Pilotti 2005; Cao et al. 2012; 56 Wang et al. 2012; Cao and Yuan 2013). Most members of Ganoderma are pathogenic in 57 nature, causing various diseases to plants like white rot, butt rot, root rot, and stem rot as well 58 as wood-decay (Ryvarden 2004; Pilotti 2005; Dai et al. 2007; Cao and Yuan 2013; Coetzee et 59 al. 2015). The genus is circumscribed by sessile to stipitate basidiomata with double-walled 60 basidiospores, and interwall pillars (Karsten 1881; Moncalvo and Ryvarden 1997). Kirk et al. 61 (2008) mentioned that globally there are 80 species of Ganoderma, while two global fungal 62 databases viz. Index Fungorum (http://www.indexfungorum.org/; accessed date: 23 May 63 2020) and MycoBank (http://www.mycobank.org/; accessed date: 23 May 2020) list 456 and 64 398 records respectively. 65 In light of an earlier taxonomic framework, Ganoderma included three subgenera viz. 66 Ganoderma P. Karst., Elfvingia P. Karst., and Trachyderma Imazeki (Zhao and Zhang 2000). 67 The subgenus Ganoderma includes two sections. Ganoderma and Phaenema contain laccate 68 species (a cutis surface consisting of a palisade of inflated hyphal ends) with a shiny upper 69 surface, whereas, all non-laccate species (palisade is absent), which have a dull upper surface, 70 are under the subgens. Elfvingia (Smith and Sivasithamparam 2000; Richter et al. 2015). The 71 subgenus Trachyderma has been deemed illegitimate and is treated as a synonym of 72 Ganoderma due to the presence of a lichenised genus, Trachyderma (Richter et al. 2015). 73 Ganoderma mushrooms have long been used as traditional medicine in many Asian countries 74 (Cao et al. 2012). In the Pharmacopoeia of the People’s Republic of China, Ganoderma 75 species are recorded as being used over two millennia (Wu et al. 2013). Traditional Chinese 3 76 books proposed and classified Ganoderma species based on basidiocarp colouration (Szedlay 77 2002). Ganoderma species are good sources of natural bioactive compounds of high and low 78 molecular weight, especially polysaccharides, protein, sterols, and triterpenoids (Ahmadi and 79 Riazipour 2007; Chan et al. 2007; Chen and Seleen 2007; Richter et al. 2015). These 80 compounds are known to possess extensive therapeutic properties, such as antibacterial, 81 anticancer, antifungal, anti-hypotensive, anti-inflammatory, antioxidant, antitumor, and 82 antiviral agents (Liu et al. 2002; Teng et al. 2011; De Silva et al. 2012a, b; Richter et al. 2015; 83 Hyde et al. 2019). Moreover, these compounds also treat many immunological diseases (Teng 84 et al. 2011; Richter et al. 2015), making Ganoderma species a popular functional food (Singh 85 et al. 2013). 86 There is much controversy in the taxonomical efforts to resolve nomenclatural 87 confusion within Ganoderma. This is because of the presence of a high level of phenotypic 88 plasticity in Ganoderma species (Pilotti et al. 2004; Wang et al. 2009; Cao et al. 2012; Dai et 89 al. 2017; Loyd et al. 2018; Hapuarachchi et al. 2019a). Ganoderma lucidum was broadly 90 defined from the original concept (Dai et al. 2017), with variations occurring in macro- 91 characteristics of the basidiomes, resulting in a large number of synonyms and general 92 redundancy in the taxonomy of this species and genus (Dai et al. 2017; Xing et al. 2018). 93 94 Biogeography and ecological aspects of Ganoderma 95 Biogeography of Ganoderma 96 Ganoderma lucidum (Curtis) P. Karst., the type species of Ganoderma was described based 97 on a specimen collected from the UK. Moncalvo et al. (1995) concluded that G. lucidum was 98 distributed across northern and southern Europe and likely extended into China. However, 99 based on analyses of ITS and LSU ribosomal DNA sequences, the illustrated G. lucidum from 100 both Europe and mainland China were not conspecific (Moncalvo et al. 1995). Moncalvo et 4 101 al. (1995), Pegler and Yao (1996), as well as Hong and Jung (2004) confirmed that G. 102 lucidum is a temperate species, so far only known with certainty from Europe. The name 103 Ganoderma lingzhi (Lingzhi) has been adopted in placeof G. lucidum for specimens native to 104 East Asia (Cao et al. 2012). Among Ganoderma species, several species similar to G. lucidum 105 have also been described worldwide: G. multipileum (Hou 1950), G. sichuanense (Zhao et al. 106 1983), and G. lingzhi (Cao et al. 2012; Wu et al. 2012) from China; G. resinaceum from 107 Europe (Patouillard 1889); and G. oregonense, G. sessile, G. tsugae, and G. zonatum from the 108 US (Murrill 1902, 1908). These species are accepted as members of the G. lucidum complex. 109 Taxonomic concepts on Ganoderma species vary greatly due to the presence of 110 phenotypic plasticity (Moncalvo and Ryvarden 1995; Moncalvo 2000). Some Ganoderma 111 species are distributed worldwide, such G. applanatum and G. australe, while some species 112 are known from specific localities (Moncalvo and Ryvarden 1997; Douanla-Meli and Langer 113 2009). Ganoderma taxa are ambiguous due to the lack of holotypes and unclear geographical 114 distributions. Here, we summarize and update the global list of Ganoderma species with type 115 localities (Table 1) based on recent literature and our findings. Ganoderma species 116 distribution in the GMS is shown in Figure 1. 117 5 Ecological aspects of Ganoderma Ganoderma species feature a high degree of morphological variation, even within species (Hapuarachchi et al. 2019a). A high level of phenotypic plasticity of the basidiomes at the macroscopic level as well as uniformity of microscopic features has led to considerable confusion and frequent misinterpretations in the taxonomy of Ganoderma (Pilotti et al.
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