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Fungal Diversity (2018) 93:215–239 https://doi.org/10.1007/s13225-018-0415-7 (0123456789().,-volV)(0123456789().,-volV) Thailand’s amazing diversity: up to 96% of fungi in northern Thailand may be novel Kevin D. Hyde1,2 • Chada Norphanphoun2 • Jie Chen3 • Asha J. Dissanayake2 • Mingkwan Doilom1,4,5 • Sinang Hongsanan6,7 • Ruvishika S. Jayawardena2 • Rajesh Jeewon8 • Rekhani H. Perera2 • Benjarong Thongbai10 • Dhanushka N. Wanasinghe1,4 • Komsit Wisitrassameewong9 • Saowaluck Tibpromma1,2,4 Marc Stadler10 • Received: 8 October 2018 / Accepted: 14 November 2018 / Published online: 22 November 2018 Ó School of Science 2018 Abstract Fungi have been often neglected, despite the fact that they provided penicillin, lovastatin and many other important medicines. They are an understudied, but essential, fascinating and biotechnologically useful group of organisms. The study of fungi in northern Thailand has been carried out by us since 2005. These studies have been diverse, ranging from ecological aspects, phylogenetics with the incorportation of molecular dating, taxonomy (including morphology and chemotaxonomy) among a myriad of microfungi, to growing novel mushrooms, and DNA-based identification of plant pathogens. In this paper, advances in understanding the biodiversity of fungi in the region are discussed and compared with those further afield. Many new species have been inventoried for the region, but many unknown species remain to be described and/or catalogued. For example, in the edible genus Agaricus, over 35 new species have been introduced from northern Thailand, and numerous other taxa await description. In this relatively well known genus, 93% of species novelty is apparent. In the microfungi, which are relatively poorly studied, the percentage of novel species is, surprisingly, generally not as high (55–96%). As well as Thai fungi, fungi on several hosts from Europe have been also investigated. Even with the well studied European microfungi an astounding percentage of new taxa (32–76%) have been discovered. The work is just a beginning and it will be a daunting task to document this astonishingly high apparent novelty among fungi. Keywords Agaricus  Amanita  Colletotrichum  Cornus  Fungal diversity  Pandanaceae  Rosaceae  Rosa  Teak fungi Introduction Fungi are an incredibly understudied, but an essential, fascinating and biotechnologically useful group of organisms. The fungi of northern Thailand have been studied by Hyde and coworkers since 2005. The studies have been diverse, ranging across ecology, traditional taxonomy, phylogenetics, evolution, microbial community and chemotaxonomy (Thongkantha et al. 2008; Pinnoi et al. 2010; Phookamsak et al. 2015; Wurzbacher et al. 2017; Norphanphoun et al. 2018; Tedersoo et al. 2018), to & Mingkwan Doilom j_hammochi@hotmail.com growing novel mushrooms (Thongklang et al. 2014), molecular identification of endophytes and plant pathogens (Jayawardena et al. 2016b; Doilom et al. 2017b), and identification of entomophagous fungi (Xiao et al. 2017, 2018). Although there are many negative facets to fungi (see Hyde et al. 2018), they are an essential component of most ecosystems and without them there would be ecological imbalance, and possibly mankind would not survive on earth (Watkinson et al. 2015). They are major contributors to nutrient cycling, and the main organisms which can degrade lignocellulose in wood and leaves (Pointing et al. 2005; Bucher et al. 2004; Tang et al. 2005); without them we would live amongst mountains of dead trees (Gadd et al. 2007). Many species exist as symbionts with plants Extended author information available on the last page of the article 123 216 and promote plant growth, including crops in many environments (Abiala et al. 2013). Fungi occur in the guts of herbivores and help to digest the consumed grasses, while some species are also passed out with the faeces and degrade dung (Kruys et al. 2015; Paul et al. 2018). Some species infect and kill insects (Sung et al. 2007), others cause disease of humans, their hair and skin (Gostinčar et al. 2018; Hyde et al. 2018) and many fungi are extremely important plant pathogens causing major yield losses, and are thus of considerable agricultural and quarantine importance (Cai et al. 2011; Hyde et al. 2014). Fungi also have an incredible biotechnological potential (Pointing and Hyde 2001; Hyde et al. 2010). The fungi of northern Thailand have been studied by Hyde and coworkers for more than a decade and we have documented the biodiversity of both macro- and microfungi and more than 500 species have been introduced. Mushroom groups have revealed an amazingly high amount of novelty, the same result being apparent for the microfungal plant pathogens, saprobes, endophytes and epitypes. Other parts of Thailand have been less-well studied for fungi, however studies of BIOTEC, Hyde and coauthors, and others, have revealed an amazing noveltly (e.g. Pinnoi et al. 2006; Pinruan et al. 2007; Tibpromma et al. 2018). In this paper, the advances made in understanding the diversity of fungi in northern Thailand are presented together with details for eight examples presented from relatively conspicuous or important groups. This data shows that the species novelty in northern Thailand is amazingly between 55–96%. The data are being extended two host groups in Europe, which also show a surpising unexpected amount of novelty. Results and discussion Studies of fungi in the conspicuous macroscopic genera Agaricus, Amanita, and Lactarius, the well studied microscopic pathogenic genera Colletotrichum and Diaporthe, the less well-studied hosts Pandanaceae and Tectona grandis, and foliar epiphytes in northern Thailand have revealed a novelty of between 55–96% (Table 1). Although we provide examples from these few selected groups, we feel they are both important and are wellstudied elsewhere and thus may be representative of the majority of fungal groups. This astonishing novelty in such conspicuous and well studied genera and groups points to the overall fact that fungi are poorly studied and if one chooses an understudied genus such as Phaeosphaeria or poorly studied hosts such as ferns, one would expect to find an even higher percentage of novelty. 123 Fungal Diversity (2018) 93:215–239 Novelty in conspicuous mushrooms From our experience, it is easier and more common to collect larger and visible species; thus mushrooms (e.g. Agaricus, Russula) and larger ascomycetes (e.g. Xylaria, Hypoxylon) have generally been better studied (Daranagama et al. 2018). Because of this, we would expect to find a lower novelty in the commonly collected mushroom genera. However, we provide examples of the novelty of species found in three conspicuous genera, mostly in northern Thailand. We show that between 83–93% of these species are novel. Agaricus Species of Agaricus (Agaricales, Basidiomycota) are saprobes which grow in various habitats, such as grassland, forests, leaf litter, sand dunes and even occur in some arid areas (Parra 2008; Karunarathna et al. 2016). During the last decade, the number of known Agaricus species has increased rapidly especially in tropical regions where, thanks to the advances in molecular phylogenetics, many new species have been revealed. From January 2000 to September 2018, almost 200 new species have been described and more than 500 Agaricus species are now recognized (Chen et al. 2017; Karunarathna et al. 2016; Kerrigan 2016; Parra et al. 2018). Of these new species, 55% (102) were described from Asia, and most of these from China and northern Thailand (Ariyawansa et al. 2015; Bashir et al. 2018: Chen et al. 2012, 2015, 2016, 2017; Dai et al. 2016; Gui et al. 2015; He et al. 2017, 2018a, b; He and Zhao 2015; Hyde et al. 2017; Kaur et al. 2016; Karunarathna et al. 2014; Li et al. 2014, 2016; Liu et al. 2015; Mahdizadeh et al. 2018; Thongklang et al. 2014, 2016; Wang et al. 2015; Zhang et al. 2017b; Zhao et al. 2012a, b, 2016; Zhou et al. 2016). We have collected more than 500 Agaricus specimens since 2005, mainly in northern Thailand. To date 38 new species have been formally introduced (Tables 1, 2) with perhaps another 30 species awaiting description. In addition, the cosmopolitan species A. subrufescens, A. endoxanthus (‘‘great traveller’’) and A. microvolvatulus are confirmed new records for Thailand (Thongklang et al. 2014; Wisitrassameewong et al. 2012a; Zhao et al. 2012a), while two species bearing identical ITS sequences to the types of A. heterocystis and A. xanthosarcus need further morphological study to confirm their identity (Zhao et al. 2011). Thus, more than 93% of the Agaricus species collected in Thailand are new to science. This is remarkable novelty for a mushroom genus that is highly prized for its edible species. If only a few of the new Agaricus species could be cultivated, and were edible and Fungal Diversity (2018) 93:215–239 217 Table 1 Fungal numbers reported from Thailand discussed in this study Subject Number of species reported before our study Number of these species confirmed by molecular data Number of species collected Agaricus 17 0 75 38 32 5 93 a Number of described new species Number remaining undescribed New records of known species to Thailand Percentage novelty (%) Expected new species in future studies 40? 27 3 69 17 40 9 83 25? 0 0 23 17b 3 3 87 30? Colletotrichum 12 5 39 16 10 8 67 50? Diaporthe 26 0 26 13 12 1 96 100? Pandanaceae Tectona grandis 87 15 3 0 93 53 54 24 15 5 21 24c 74 55 100? 100 6 0 74 42 23 9 88 100 Amanita Lactarius subg. Russularia Foliar epiphytes a Three species recorded by Li et al. (2016); bOne species recorded by Verbeken et al. (2014); cTwo species recorded by Meeboon and Takamatsu (2017) tasty, then numerous new species could be introduced to international cuisine (Thawthong et al. 2014). Agaricus subrufescens is an important medicinal species (Wisitrassameewong et al. 2012a), and we predict that many of the other species could also be cultivated and used as functional foods and medicine. It must be noted that before 2011, only 17 species of Agaricus were recorded in Thailand and these were published in local reports and books (Zhao 2008; Chandrasrikul et al. 2011). However, of these 17 identifications, many were linked to species originally described from Europe, such as A. campestris, A. bisporus and A. bitorquis; these determinations are doubtful and cannot be confirmed due to a lack of herbarium material, detailed descriptions and molecular data. Amanita Amanita is an important genus of mushrooms and its section Phalloideae includes several species that are widely recognized as the most poisonous mushrooms in the world (Hyde et al. 2018). The genus also includes hallucinogenic species such as A. muscaria (type species of Amanita) and A. pantherina, as well as prized edible mushrooms, such as A. caesarea and other species of sect. Caesarea. Most species of Amanita are considered to be ectomycorrhizal and their distribution in forests and heaths including Betulaceae, Dipterocarpaceae, Fabaceae, Myrtaceae, Pinaceae, and Salicaceae, suggests that they play a critical role in forest ecosystems worldwide (Weiß et al. 1998; Yang 1997; Zhang et al. 2004). There are 36 taxa reported as non-ectomycorrhizal (Wolfe et al. 2012). As of July 2018, Amanita comprises just under 1000 taxa of which 600 have validly published names, 305 are known by provisional names or temporary codes, and the remainder have misapplied, invalid or illegitimate names (Tulloss and Yang 2018; Cui et al. 2018). Cui et al. (2018) dealt with the rearrangement of the Amanita, mainly based on Chinese materials. They recognised 156 taxa of Amanita in China and reported on several others, but it is anticipated that additional species remain to be formally named. Although they gave no update on the total number of species in Amanita. Cui et al. (2018) revisited its classification and recognised eleven sections and three subgenera (Amanita, Amanitina and Lepidella). Prior to the current study, 25 species (with one species affinis) of Amanita had been reported from northern Thailand (Sanmee et al. 2008). Since 2012, we have collected more than 250 specimens of Amanita, mainly in the north (Table 3). Of all Amanita species collected in Thailand, three are confirmed records of previously recorded taxa, nine are new records of known species, 17 are new species (three from Li et al. 2016) and 40 are species awaiting description. Thus, 83% of the collected species of Thai Amanita are new to science. Lactarius subg. Russularia Lactarius species are commonly known as milk-caps due to their latex exudation when the basidiomata are injured. They form ectomycorrhizal associations with diverse groups of terrestrial plants, both deciduous and coniferous. Some Lactarius species, such as L. deliciosus (L. Fr.) Gray, L. indigo (Schwein.) Fr. and L. hatsudake Tanaka are sought after mushrooms due to their pleasant taste. Of the 123 218 Fungal Diversity (2018) 93:215–239 Table 2 Agaricus species recorded from Thailand (novel species are in bold) Table 3 Amanita species recorded from Thailand (novel species are in bold) Taxon References Taxon References Agaricus albosquamosus Zhao et al. (2016) Amanita atrobrunnea Li et al. (2016) Agaricus angusticystidiatus He et al. (2018a) Amanita ballerina Thongbai et al. (2017a) Agaricus atrodiscus Ariyawansa et al. (2015) Amanita brunneitoxicaria Thongbai et al. (2017a) Agaricus badioniveus Chen et al. (2017) Amanita brunneoprocera Thongbai et al. (2018) Agaricus bisporiticus Thongklang et al. (2014) Amanita brunneosquamata Thongbai et al. (2018) Agaricus brunneolutosus Chen et al. (2017) Amanita brunneoumbonata Thongbai et al. (2018) Agaricus brunneosquamulosus Agaricus chiangmaiensis Chen et al. (2015) Karunarathna et al. (2014) Amanita castanea Amanita cinnamomea Thongbai et al. (2016) Thongbai et al. (2018) Agaricus endoxanthus Zhao et al. (2012a) Amanita concentrica Thongbai et al. (2016) Agaricus erectosquamosus Zhao et al. (2016) Amanita digitosa Li et al. (2016) Agaricus exilissimus Ariyawansa et al. (2015) Amanita esculenta Thongbai et al. (2018) Agaricus fimbrimarginatus Chen et al. (2017) Amanita flavidocerea Thongbai et al. (2018) Agaricus flammicolor Chen et al. (2017) Amanita flavidogrisea Thongbai et al. (2018) Agaricus flavicentrus Liu et al. (2015) Amanita fuligineoides Thongbai et al. (2017a) Agaricus flocculosipes Zhao et al. (2012b) Amanita gleocystidiosa Li et al. (2016) Agaricus fuscopunctatus Thongklang et al. (2014) Amanita hemibapha sensu lato Sanmee et al. (2008) Agaricus haematinus Ariyawansa et al. (2015) Amanita luteoparva Thongbai et al. (2018) Thongbai et al. (2017a) Agaricus hanthanaensis Liu et al. (2015) Amanita macrocarpa Agaricus heterocystis Zhao et al. (2011) Amanita cf. oberwinklerana Thongbai et al. (2017a) Agaricus inthanonensis Zhao et al. (2016) Amanita pyriformis Li et al. (2016) Agaricus leucocarpus Chen et al. (2017) Amanita pseudoporphyria Sanmee et al. (2008) Agaricus leucolepidotus Agaricus luteofibrillosus Zhao et al. (2016) Li et al. (2016) Amanita rimosa Amanita rubromarginata Thongbai et al. (2016) Thongbai et al. (2016) Agaricus luteopallidus Chen et al. (2017) Amanita rubrovolvata Sanmee et al. (2008) Agaricus megacystidiatus Karunarathna et al. (2014) Amanita cf. spissacea Thongbai et al. (2017a) Agaricus megalosporus Chen et al. (2012) Amanita strobilipes Li et al. (2016) Agaricus microvolvatulus Thongklang et al. (2014) Amanita suborientifulva Thongbai et al. (2018) Agaricus murinocephalus Zhao et al. (2012a) Amanita subovalispora Thongbai et al. (2018) Agaricus niveogranulatus Chen et al. (2015) Amanita zangii Thongbai et al. (2016) Agaricus parvibicolor Liu et al. (2015) Agaricus patris Chen et al. (2017) Agaricus pseudolangei Ariyawansa et al. (2015) Agaricus purpureofibrillosus Chen et al. (2017) Agaricus robustulus Chen et al. (2017) Agaricus sodalis Liu et al. (2015) Agaricus sordidocarpus Chen et al. (2015) Agaricus subrufescens Wisitrassameewong et al. (2012b) Agaricus subtilipes Agaricus suthepensis Zhao et al. (2016) Zhao et al. (2016) Agaricus toluenolens Chen et al. (2015) Agaricus variicystis Zhao et al. (2016) Agaricus xanthosarcus Zhao et al. (2011) Agaricus brunneogracilis Zhou et al. (2016) three currently accepted subgenera, Lactarius subg. Russularia (Fr.) Kauffman is a difficult group to study due to the similarity in macromorphological and latex features 123 amongst species. Species in this subgenus can be recognised in the field by the orange to warm brown to reddish brown fruiting bodies, which are typically dry and fragile, the unchanging latex and the smell of Pentatomidae bugs. In Thailand, although subg. Russularia species are abundant in nature, they have been often overlooked by locals because they are small and fragile and have a poor taste. Subg. Russularia is one of the dominant mushroom groups in terms of the species numbers and numbers of basidiomata distributed in Thai forests. They are associated with several dominant genera of trees, e.g. Dipterocarpus, Shorea, Castanopsis, Lithocarpus, Quercus, Betula and Pinus. Until recently, knowledge of Lactarius subg. Russularia in Thailand was very poor; only L. chichuensis W.F. Chiu, L. gracilis Hongo, and L. subzonarius Hongo were reported (Le 2007). In addition, European names such Fungal Diversity (2018) 93:215–239 219 as L. camphoratus (Bull.: Fr.) Fr., were often applied for Thai species. Thus, the biodiversity of Lactarius subg. Russularia in northern Thailand was explored (Wisitrassameewong et al. 2014a, b, 2015; Liu et al. 2015). More than 100 collections of subg. Russularia were made since 2007, most from the four northern provinces (Chiang Mai, Chiang Rai, Mae Hong Son and Lampang). For species delimitation, we relied on morphology, molecular phylogeny based on ITS and rpb2 regions and to a lesser extent the ecology of host genera. Seventeen novel species (Table 4) were published, along with three new records for Thailand, and another three are new species awaiting description. Thus, in Lactarius subg. Russularia 87% of species collected in northern Thailand are new to science. We predict that with more extensive sampling, more than 30 new cryptic species will be found in coming years. Together with collaboration from Mycology laboratory of Ghent University, we compared our data with data of European species in order to investigate the eventual intercontinental conspecificity of the mycota found in European temperate and Asian tropical regions. There is no case of conspecificity between European and Thai species. Therefore, European or North American names should generally not be used for Thai taxa. Apart from our Table 4 Lactarius subg. Russularia species recorded from Thailand (novel species are in bold) Taxa References Lactarius aquosus Lactarius atrobrunneus Wisitrassameewong et al. (2015) Liu et al. (2015) Lactarius austrorostratus Wisitrassameewong et al. (2015) Lactarius chichuensis Wisitrassameewong et al. (2015) Lactarius crenulatulus Wisitrassameewong et al. (2014a) Lactarius falcatus Verbeken et al. (2014) Lactarius fuscomaculatus Wisitrassameewong et al. (2015) Lactarius grabrigracilis Wisitrassameewong et al. (2014b) Lactarius gracilis Wisitrassameewong et al. (2014b) Lactarius inconspicuus Wisitrassameewong et al. (2015) Lactarius kesiyae Wisitrassameewong et al. (2015) Lactarius laccarioides Wisitrassameewong et al. (2014a) Lactarius pasohensis Wisitrassameewong et al. (2014a) Lactarius perparvus Wisitrassameewong et al. (2014b) Lactarius politus Liu et al. (2015) Lactarius rubrobrunneus Lactarius rubrocorrugatus Wisitrassameewong et al. (2015) Wisitrassameewong et al. (2015) Lactarius sublaccarioides Wisitrassameewong et al. (2014a) Lactarius subzonarius Hongo (1957) Lactarius tangerinus Wisitrassameewong et al. (2015) described species, the sequestrate L. falcatus Verbeken & Van de Putte was also reported from deciduous forest in northern Thailand by Verbeken et al. (2014). Novelty in plant pathogens Diseases caused by plant pathogens may result in considerable losses to food production, as exemplified by black stem rust of wheat (Puccinia spp., Zadoks 1985), late blight of potato (Phytopthora infestans, Fry et al. 2013) and rice blast disease (Magnaporthe oryzae, Ou 1980). The introduction of exotic plant pathogens may also seriously affect farming, forestry and the environment (Jayawardena et al. 2016a; Hyde et al. 2018), as well as global plant trade resulting in huge economic losses to a country (Jayawardena et al. 2016a). Plant pathogens continue to develop resistance against chemicals and host crop defence mechanisms (Crouch 2014) and this has become a challenge in developing control strategies. Most pathogens are microfungi and although the fungus may not be easily seen, the disease symptoms they cause are both highly visual and often occur in epidemic proportions resulting in large yield losses. For this reason, plant pathogens are very well studied and not a group where we would expect to find a high diversity of novel species. In this study, we show that in two prominent plant pathogenic genera collected in northern Thailand, 67% of species in Colletotrichum and 96% in Diaporthe are novel. Colletotrichum Colletotrichum is one of the most important phytopathogenic genera worldwide affecting quality and yield of many economical crops (Hyde et al. 2009; Cannon et al. 2012; Jayawardena et al. 2016b). In a checklist of plant diseases in Thailand (Giatgong 1980), 12 named species of Colletotrichum were listed, while undetermined species were recorded from many different hosts. This host-fungi index was based solely on past literature and taxa were named based on morphological characters. Molecular data are essential to identify Colletotrichum to species level (Shenoy et al. 2007; Cai et al. 2009; Cannon et al. 2012; Hyde et al. 2009, 2014; Jayawardena et al. 2016b; Damm et al. 2019) and therefore these old records must be treated as dubious. For example, in earlier studies carried out in Thailand many species were identified as C. acutatum and C. gloeosporioides. Both of these are now considered as species complexes (Jayawardena et al. 2016b). As there is no herbarium material or cultures we cannot recheck these records. We have been studying Colletotrichum in Thailand since 2007. Colletotrichum gloeosporioides, which was thought to be a common pathogen in tropics, turned out not to be that common and 123 220 may even not be present (Phoulivong et al. 2010). Phoulivong et al. (2010) analyzed DNA sequence data of 25 isolates from eight tropical fruits, which were morphologically identified as C. gloeosporioides in previous studies. Contrary to previous understanding, none of the 25 isolates clustered with the epitype of C. gloeosporioides in the multi-gene phylogenetic analyses. Than et al. (2008a) identified C. acutatum, C. capsici and C. gloeosporioides as the causal agents of anthracnose in chili in Thailand based on morphological characters. However, with the use of ITS and b-tubulin sequence data, Than et al. (2008b) showed that C. acutatum, C. capsici, C. gloeosporioides and C. siamenese are the causal agents of chili anthracnose in Thailand. Morphological characters can be used to differentiate Colletotrichum into species complexes (Hyde et al. 2014; Jayawardena et al. 2016b) but, they cannot be used to separate species within a complex (Phoulivong et al. 2010; Jayawardena et al. 2016b). We have collected more than 200 specimens of Colletotrichum mainly in north Thailand. Of these, eight are new records (hosts/locations), 16 are new species (Table 5) and ten new species that await description. Thus, in Colletotrichum 67% of collected species are new to science (Table 1). The remaining collection representing about 40 species need either more material or additional sequences other than ITS rDNA for a formal description. We predict that with extensive sampling, more cryptic species will be introduced in coming years, with perhaps more than 50 new species. Diaporthe Diaporthe (syn. Phomopsis) species are well known as pathogens, endophytes or saprobes on a range of economical crops, ornamentals and forest trees (Rehner and Uecker 1994; Santos and Phillips 2009; Santos et al. 2011; Udayanga et al. 2011, 2012a, b, 2014; Hyde et al. 2014; Dissanayake et al. 2015, 2017b, c). In the past species of Diaporthe were introduced largely on the basis of host association, which resulted in a proliferation of species names. However, it is now recognised that many of the species are not host-specific and a single species can be found on more than one host (Dissanayake et al. 2017b). Only a few studies related to Diaporthe/Phomopsis pathogens have been conducted in Thailand. Hyde (1991) introduced a novel Phomopsis species: Phomopsis mangrovei, from intertidal prop roots of Rhizophora apiculata in Thailand. Sontirat et al. (1994) listed eight unnamed Diaporthe species and four unnamed Phomopsis species on various host plants in the checklist of Thai pathogens. Based on molecular data, Udayanga et al. (2012a) reported eleven undescribed Diaporthe isolates. Oeurn et al. (2015) found another Phomopsis sp. on dragon fruit stems in Loei 123 Fungal Diversity (2018) 93:215–239 Province, Thailand, but no molecular data were used to support its identity. A survey of leaf spots associated with disease of durian caused by Phomopsis durionis was conducted by Tongsri et al. (2016). Thus, until the incorporation of molecular data, 26 Diaporthe/Phomopsis taxa had been reported to cause diseases on various hosts in Thailand. We have been studying Diaporthe in Thailand since 2012. Twenty-six species were collected mainly in the north, of which none are confirmed existing records, include 14 species formally described (Table 1). Of all Diaporthe species collected in Thailand, 13 are new species and one is new species record that we have described (Table 6) and another twelve are new species waiting to be described. thus, 96% of collected Diaporthe species in northern Thailand are new to science. Novelty in fungi on various hosts Another approach to studying fungal diversity is to target a certain host, and make an inventory of the fungi that are associated with it, either as pathogens, mycorrhizal symbionts, endophytes or epiphytes. Tropical hosts are generally not well studied and fungal novelty might thus be high. We discuss fungi on Pandanaceae and teak (Tectona grandis), and show that 55–74% of species are novel. Pandanaceae The plant family Pandanaceae belongs to monocotyledonous and its species have a worldwide distribution and occur throughout Thailand. Microfungi on Pandanaceae in Thailand have been relatively well studied, although the taxonomic studies lacked molecular data (Manoch et al. 1986; Sivanesan 1987; Tokumasu et al. 1990; Thienhirun 1997; Sivichai et al. 1998; Goh et al. 1999; Pinnoi et al. 2004; Thongkantha et al. 2008; Whitton et al. 2012). The fungi on Pandanaceae in Thailand have been studied by us since 2014. More than 150 specimens were collected, comprising 99 species. Of all the species we collected on Pandanaceae in Thailand, three are confirmed existing records, 21 are new records, 54 are new species (Tables 1, 7) and another 15 are new species waiting to be described. Thus, on Pandanaceae 74% of species are new to science. Teak fungi Teak is one of the most economically valuable hardwood trees globally. The genus Tectona is a member of the family Lamiaceae belonging to order Laminales. Teak is distributed in many countries, and Thailand has a natural distribution of teak forests. Studies on the fungi on teak in Fungal Diversity (2018) 93:215–239 221 Table 5 Colletotrichum species recorded from Thailand (novel species from Thailand are in bold Species Host References Colletotrichum acidae Phyllanthus acidus Samarakoon et al. (2018)a Capsicum annuum Than et al. (2008b) Colletotrichum acutatum Fragaria sp. Fragaria sp. Photita et al. (2004) Capsicum annuum Suwannarat et al. (2017)a Capsicum annuum Diao et al. (2017)a Colletotrichum aeschynomenes Manihot esculenta Sangpueak et al. (2018)a Colletotrichum asianum Coffea arabica Prihastuti et al. (2009) Mangifera indica Colletotrichum boninense Dendrobium sp. Ma et al. (2018)a Colletotrichum brevisporum Manihot esculenta Neoregelia sp., Pandanus pygmaeus Sangpueak et al. (2018)a Noireung et al.(2012)a Colletotrichum cariniferi Dendrobium cariniferum Ma et al. (2018)a Colletotrichum chiangraiense Dendrobium sp. Ma et al. (2018)a Colletotrichum citricola Dendrobium sp. Ma et al. (2018)a Colletotrichum cordylinicola Cordyline fruticosa Phoulivong et al. (2010) Colletotrichum doitungense Dendrobium fimbriatum Ma et al. (2018)a Colletotrichum endophytica Pennisetum purpureum Manamgoda et al. (2013)a Colletotrichum fructicola Coffea arabica Phoulivong et al. (2010)a Capsicum annuum Than et al. (2008a, b), Diao et al. (2017)a Carica papaya Dimocarpus longan Cymbopogon citratus, Manamgoda et al. (2013)a Pennisetum purpureum Dendrobium sp. Ma et al. (2018)a Colletotrichum fusiforme Unknown Ariyawansa et al. (2015)a Colletotrichum gigasporum Alocasia sp., Hibiscus rosa-sinensis Liu et al. (2014)a Capsicum annuum, Fragaria sp., Mangifera indica Than et al. (2008a, b) Magnolia liliifera Promputtha et al. (2004) Stylosanthes fruticosa, Stylosanthes hamata, Stylosanthes humilis, Stylosanthes scabra Masel et al.(1993)N/A Alpinia malaccensis, Draceana sanderiana, Eupatorium thymifolia, Alpinia galanga, Mangifera indica, Musa acuminata, Manihot esculenta Photita et al. (2004), Sangpueak et al. (2018)a Rottboellia cochinchinensis Sherriff et al. (1995)N/A Manihot esculenta Sangpueak et al. (2018)a Colletotrichum musae Musa acuminata, Musa sp. Su et al. (2011)a Colletotrichum orchidearum Hymenocallis sp. Damm et al. (2019)a Colletotrichum orchidophilum Dendrobium sp. Ma et al. (2018)a Colletotrichum pandanicola Pandanus sp. Tibpromma et al. (2018a)a Colletotrichum parallelophorum Dendrobium sp. Ma et al. (2018)a Colletotrichum scovillei Capsicum annuum, Capsicum sp. Damm et al. (2012)a Colletotrichum siamense Coffea arabica Capsicum annuum, Hymenocallis sp. Phoulivong et al. (2010) Than et al. (2008a, b), Yang et al. (2009)a Cymbopogon citratus, Pennisetum purpureum Manamgoda et al. (2013)a Colletotrichum syzygiicola Citrus aurantifolia, Syzygium samarangense Udayanga et al. (2013)a Colletotrichum tropicale Pennisetum purpureum Manamgoda et al. (2013)a Colletotrichum gloeosporioides Colletotrichum graminicola 123 222 Fungal Diversity (2018) 93:215–239 Table 5 (continued) Species Host References Colletotrichum tropicicola Citrus maxima, Paphiopedilum bellatulum Noireung et al. (2012)a Colletotrichum truncatum Capsicum annuum, C. frutescens, Capsicum sp., Manihot esculenta, Solanum melongena, Vigna sesquipedalis, Glycine max, Stylosanthes hamata, Hymenocallis sp., Gossypium sp. Photita et al. (2004), Than et al. (2008a, b), Yang et al. (2009)a, Diao et al. (2017)a, Suwannarat et al. (2017)a, Sangpueak et al. (2018)a Colletotrichum watphraense Dendrobium sp. Ma et al. (2018)a a Phylogenetic studies including other gene regions apart from ITS sequence data Table 6 Diaporthe species recorded from Thailand (novel species from Thailand are in bold) confirmed with molecular data Taxon Host References Diaporthe aseana Unknown dead leaf Hyde et al. (2016) Diaporthe collariana Magnolia champaca Perera et al. (2018) Diaporthe garethjonesii Unknown dead leaf Hyde et al. (2016) Diaporthe neoraonikayaporum Tectona grandis Doilom et al. (2016, 2017a) Diaporthe phaseolorum Hylocereus undatus Udayanga et al. (2012a) Diaporthe pterocarpi Pterocarpus indicus Udayanga et al. (2012b) Diaporthe pterocarpicola Pterocarpus indicus Udayanga et al. (2012b) Diaporthe siamensis Dasymaschalon sp. Udayanga et al. (2012b) Diaporthe tectonae Tectona grandis Doilom et al. (2016, 2017a) Diaporthe tectonendophytica Tectona grandis Doilom et al. (2016, 2017a) Diaporthe tectonigena Tectona grandis Doilom et al. (2016, 2017a) Diaporthe thunbergii Thunbergia laurifolia Udayanga et al. (2012b) Diaporthe thunbergiicola Thunbergia laurifolia Liu et al. (2015) Diaporthe rosae Rosa sp. Wanasinghe et al. (2018) Expected number of new species: [ 100 Thailand are few. Fifteen taxa, mostly without molecular data, have been reported from Thailand, such as Alternaria alternata, Cercospora tectonae, Daldinia eschscholtzii, Hypoxylon haematostroma, Nigrospora sphaerica, Olivea tectonae (: Uredo tectonae), Schizophyllum commune, Xylaria allantoidea and X. feejeensis before we commenced our research (Giatgong 1980; Lorsuwan et al. 1984; Chareprasert et al. 2006; Meeboon et al. 2007; Mekkamol 1998; Okane et al. 2008; To-anun et al. 2011). We have studied teak fungi since 2011. More than 120 specimens were collected mainly in the north. Of all teak fungi we collected in Thailand, none were confirmed as existing records, 24 are new records (two from Meeboon and Takamatsu 2017), 24 are new species (Tables 1, 8) and five are potentially novel species awaiting description. Thus, from teak 55% of collected species are new to science. Novelty in various habitats Specific habitats are generally less well-studied, especially for microfungi in the tropics and therefore we might expect 123 a high novelty if we study such a habitat in detail. In this section we look at the novelty of foliar epiphytes. These are minor plant pathogens and while some are highly visible, most are hard to observe, and as a whole have received little attention. In this study, we show that 88% of foliar epiphytes species are novel. Foliar epiphytes Fungal epiphytes commonly occur on plant surfaces, particularly the leaves (Carroll 1991; Gilbert and Reynolds 2002, 2005; Wu et al. 2011; Hongsanan et al. 2016b). This is a polyphyletic group belonging in the Ascomycota (Schoch et al. 2009; Li et al. 2016; Wu et al. 2011; Hyde et al. 2013; Hongsanan et al. 2016b). Fungal epiphytes, which are obligate parasites, can cause damage to host plants, e.g. resulting in lower yields, chlorosis and plantstunting disease (Ariyawansa et al. 2015; Hongsanan et al. 2014a, 2015a, c, 2016b). The coating of hyphae on the surface of plants in some species may result in marketability problems (Chomnunti et al. 2014). The ecology and taxonomy of fungal epiphytes has been studied Fungal Diversity (2018) 93:215–239 223 Table 7 Fungal species on Pandanaceae reported in Thailand with morphological and molecular data (novel species from Thailand are in bold) Taxa Host Location References Acremoniisimulans thailandensis Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Alternaria burnsiia Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Anthostomelloides krabiensis Pandanus odorifer Krabi Province Tibpromma et al. (2017a) Beltrania krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Beltraniella pandanicola Pandanus sp. Phuket Province Tibpromma et al. (2018b) Beltraniella thailandicus Pandanus sp. Chonburi Province Tibpromma et al. (2018b) Byssosphaeria siamensis Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Canalisporium krabiense Pandanus sp. Krabi Province Tibpromma et al. (2018b) Canalisporium thailandensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Cercospora capsici Pandanus amaryllifolius Chiang Mai Province Tibpromma et al. (2018b) Chaetomium globosum Cladosporium endophyticuma Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Pandanus sp. Krabi Province Tibpromma et al. (2018a) Clonostachys krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Colletotrichum fructicolaa Pandanus sp., Freycinetia sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Colletotrichum pandanicola Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Colletotrichum pandanicolaa Pandanus sp. Chumphon Province Tibpromma et al. (2018a) Curvularia chonburiensis Pandanus sp. Chonburi Province Tibpromma et al. (2018b) Curvularia pandanicola Pandanus sp. Krabi Province Tibpromma et al. (2018b) Curvularia thailandicum Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Deniquelata barringtoniae Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Diaporthe pandanicolaa Pandanus sp. Chumphon Province Tibpromma et al. (2018a) Diaporthe siamensisa Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Dictyochaeta siamensis Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Dictyocheirospora pandanicola Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Dictyosporium appendiculatum Pandanus sp. Nakhon Si Thammarat Province Tibpromma et al. (2018b) Dictyosporium guttulatum Pandanus sp. Krabi Province Tibpromma et al. (2018b) Dictyosporium krabiense Dictyosporium pandanicola Pandanus sp. Pandanus sp. Krabi Province Krabi Province Tibpromma et al. (2018b) Tibpromma et al. (2018b) Distoseptispora thailandica Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Endomelanconiopsis freycinetiaea Freycinetia sp. Ranong Province Tibpromma et al. (2018a) Endopandanicola thailandicaa Pandanus sp., Freycinetia sp. Chumphon Province Tibpromma et al. (2018a) Helicoma freycinetiae Freycinetia javanica Phang Nga Province Tibpromma et al. (2018b) Hermatomyces krabiensis Pandanus odorifer Krabi Province Tibpromma et al. (2016b) Hermatomyces krabiensis (= H. chiangmaiensis) Pandanus sp. Chiang Mai Province Tibpromma et al. (2017b) Hermatomyces pandanicola Pandanus odorifer Phang Nga Province Tibpromma et al. (2016b) Hermatomyces saikhuensis Pandanus odorifer Prachuap Khiri Khan Province Tibpromma et al. (2016b) Lasiodiplodia chonburiensis Pandanus sp. Chonburi Province Tibpromma et al. (2018b) Lasiodiplodia hyalina Pandanus sp. Chiang Mai Province Tibpromma et al. (2018b) Lasiodiplodia pandanicola Pandanus sp. Phatthalung Province Tibpromma et al. (2018b) Lasiodiplodia pseudotheobromae Pandanus sp. Chiang Rai Province Tibpromma et al. (2018b) Lasiodiplodia theobromaea Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Lasionectria krabiense Malaysiasca phaii Pandanus sp. Freycinetia javanica Krabi Province Krabi Province Tibpromma et al. (2018b) Tibpromma et al. (2018b) Massarina pandanicolaa Pandanus sp. Chumphon Province Tibpromma et al. (2018a) Meyerozyma caribbicaa Pandanus sp., Freycinetia sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Montagnula krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Musicillium pandanicola Pandanus sp. Chiang Mai Province Tibpromma et al. (2018b) Mycoleptodiscus endophytica Freycinetia sp. Ranong Province Tibpromma et al. (2018a) 123 224 Fungal Diversity (2018) 93:215–239 Table 7 (continued) Taxa Host Location References Hyde et al. (2018) Neomassarina pandanicola Pandanus sp. Prachuap Khiri Khan Province Neooccultibambusa thailandensis Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Neopestalotiopsis chiangmaiensis Pandanus sp. Chiang Mai Province Tibpromma et al. (2018b) Neopestalotiopsis phangngaensis Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Novomicrothelia pandanicola Pandanus tectorius Chanthaburi Province Zhang et al. (2017a, b) Pandanaceomyces krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Parasarcopodium pandanicola Pandanus sp. Krabi Province Tibpromma et al. (2016a) Parascedosporium putredinis Pestalotiopsis jiangxiensisa Pandanus sp. Pandanus sp. Krabi Province Chumphon, Ranong Province Tibpromma et al. (2018b) Tibpromma et al. (2018a) Pestalotiopsis krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Pestalotiopsis microsporaa Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Phanerochaete chrysosporiuma Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Phyllosticta capitalensisa Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a) Pseudoachroiostachys krabiense Pandanus sp. Krabi Province Tibpromma et al. (2018b) Pseudochaetosphaeronema pandanicola Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Pseudofusicoccum adansoniae Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Pseudohyaloseta pandanicola Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Pseudoornatispora krabiense Pandanus sp. Krabi Province Tibpromma et al. (2018b) Pseudopithomyces pandanicola Pandanus amaryllifolius Chiang Rai Province Tibpromma et al. (2018b) Roussoella solani Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Sirastachys phangngaensis Pandanus sp. Phang Nga Province Tibpromma et al. (2018b) Terriera pandanicola Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) Thozetella pandanicola Torula ficus Pandanus sp. Pandanus sp. Krabi Province Chiang Mai Province Tibpromma et al. (2018b) Tibpromma et al. (2018b) Tubeufia freycinetiae Freycinetia javanica Phang Nga Province Tibpromma et al. (2018b) Tubeufia inaequalis Pandanus sp. Krabi Province Tibpromma et al. (2018b) Tubeufia pandanicola Pandanus sp. Krabi Province Tibpromma et al. (2018b) Tubeufia parvispora Pandanus sp. Krabi Province Tibpromma et al. (2018b) Volutella krabiense Pandanus sp. Krabi Province Tibpromma et al. (2018b) Volutella thailandensis Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b) a Endophytic fungi intensively for many decades, but still there are numerous undiscovered species (Blakeman 1981; Dickinson and Preece 1976; Fokkema and van den Heuvel 1986; Carroll 1991; Gilbert and Reynolds 2002, 2005; Wu et al. 2011; Chomnunti et al. 2014; Hyde et al. 2013, 2016; Hongsanan et al. 2014a, 2015a, b, 2017). This is largely because many taxa will not grow in culture as they are biotrophs and thus it is difficult to obtain sequence data (Hongsanan et al. 2014a, 2017). DNA extraction from fresh material has been used as a core solution to this issue, however, fungal epiphytes often grow intermixed with colonies of other species (Chomnunti et al. 2014), and this may lead to contamination problems and difficulties in obtaining appropriate DNA sequence from targeted organisms. Mostly fungal epiphytes in Thailand have been reported without species-level identification (Athipunyakom and 123 Likhitekaraj 2006). The foliar epiphytes in Thailand have been studied by us since 2008. More than 170 specimens were collected mainly in the north, which included 74 species (Table 1). Of all foliar epiphytes we collected, none are confirmed existing records, nine are new records, 42 are new species (Table 9), and 23 are potentially novel species awaiting description. These 51 described taxa have been shown to belong to Asterinales (8), Capnodiales and Chaetothyriales (26), incertae sedis (3), Meliolales (7), Microthyriaceae and Micropeltidaceae (3), Muyocopronales (1), Zeloasperisporiales (3) (Wu et al. 2011; Hongsanan et al. 2014a, b, 2015a, b, c, 2016a, b, 2017; Liu et al. 2015; Ariyawansa et al. 2015; Hyde et al. 2016; Tibpromma et al. 2017b). Thus, 88% of foliar epiphyte species collected are new to science. Fungal Diversity (2018) 93:215–239 Table 8 Species of fungi on Tectona grandis mostly reported in northern Thailand (novel species from Thailand are in bold) 225 Species References Alternaria tillandsiae Doilom et al. (2017a) Barriopsis tectonae Doilom et al. (2014) Barriopsis thailandica Tibpromma et al.(2017a) Berkleasmium talaumae* Doilom et al. (2017a) Boerlagiomyces macrosporus Doilom et al. (2017a) Ceratocladium purpureogriseum* Doilom et al. (2017a) Chaetomium globosum Maharachchikumbura et al. (2016) Diaporthe neoraonikayaporum Doilom et al. (2017a) Diaporthe tectonae Doilom et al. (2017a) Diaporthe tectonendophytica Doilom et al. (2017a) Diaporthe tectonigena Doilom et al. (2017a) Diatrypella tectonae Shang et al. (2017) Distoseptispora tectonae Hyde et al. (2016) Distoseptispora tectonigena Hyde et al. (2016) Dothiorella tectonae Doilom et al. (2015) Erysiphe mori Meeboon and Takamatsu (2017) Erysiphe tectonae Meeboon and Takamatsu (2017) Helicoma siamense Doilom et al. (2017a) Hermatomyces indicus Doilom et al. (2017a), Koukol et al. (2018) Hermatomyces sphaericus Doilom et al. (2017a), Koukol et al. (2018) Huntiella chinaeucensis Maharachchikumbura et al. (2016) Kirschsteiniothelia tectonae Li et al. (2016) Lasiodiplodia brasiliensis Doilom et al. (2015) Lasiodiplodia pseudotheobromae Doilom et al. (2015) Lasiodiplodia theobromae Doilom et al. (2015, 2017a) Longiostiolum tectonae Li et al. (2016) Macrovalsaria megalospora Doilom et al. (2017a) Manoharachariella tectonae Doilom et al. (2017a) Melanoctona tectonae Tian et al. (2016) Neocosmospora solani (= Fusarium solani) Doilom et al. (2017a) Neooccultibambusa chiangraiensis Doilom et al. (2017a) Paradictyoarthrinium diffractum Liu et al. (2015), Doilom et al. (2017a) Paradictyoarthrinium tectonicola Liu et al. (2015) Phaeoacremonium italicum Doilom et al. (2017a) Phaeoacremonium tectonae Ariyawansa et al. (2015) Phyllosticta capitalensis Wikee et al. (2013a, b) Pseudocoleodictyospora sukhothaiensis Doilom et al. (2017a) Pseudocoleodictyospora tectonae Doilom et al. (2017a) Pseudocoleodictyospora thailandica Doilom et al. (2017a) Pseudofusicoccum adansoniae Doilom et al. (2015) Pseudomonodictys tectonae Ariyawansa et al. (2015) Rhytidhysteron tectonae Doilom et al. (2017a) Sphaeropsis eucalypticola Doilom et al. (2015, 2017a) Stachybotrys levisporus Doilom et al. (2017a) Stachybotrys renisporus Doilom et al. (2017a) Subglobosporium tectonae Doilom et al. (2017a) Thaxteriellopsis lignicola Doilom et al. (2017a) Tubeufia tectonae Doilom et al. (2017a) Expected number of new species: [ 100 a No molecular data available 123 226 What about other countries? One may assume that because Thailand is tropical and was previously poorly studied, we should expect to find a high fungal novelty. However, what is the situation in welldocumented countries? We provided some answers to this question by studying the fungi on Rosaceae and Cornus in Europe. We show that 76% of species on Rosaceae and 32% on Cornus are novel. Fungal Diversity (2018) 93:215–239 The fungi on Cornus species have been studied by us since 2015. More than 100 specimens were collected mainly from Italy and Russia, which included 77 species comprising 52 existing species and 25 novel species waiting to be described. Thus, of all fungi collected on Cornus, 43 are confirmed as existing records from Cornus, and nine are new host records. Therefore, 32% of fungi collected on Cornus species are new to science. We predict that with extensive sampling, over 100 novel species will be introduced in the coming years. Fungi on Rosaceae Novelty of fungi in Europe Rosaceae is one of the largest families of flowering plants including over 3000 species mostly distributed in the northern hemisphere (Wanasinghe et al. 2018). There are more than 4000 records of fungi on Rosaceae species in the U.S. National Fungus Collections Fungus-Host Database (Wanasinghe et al. 2018), but they are poorly investigated in terms of taxonomic relationships with molecular identification. Fungi on Rosaceae species have been reported in recent studies as saprobes (Dissanayake et al. 2017a; Wanasinghe et al. 2017), endophytes (Salgado-Salazar et al. 2008; Rovná et al. 2015), mycorrhizae (Bzdyk et al. 2016; El-Bashiti et al. 2017) or pathogens (Yan et al. 2015; Deng et al. 2017; Santos et al. 2017; Wang et al. 2017). The fungi on Rosa species have been studied by us since 2013. More than 200 specimens were collected mainly from Italy, Russia, Sweden, UK and Uzbekistan and resulted in 59 novel species, with 15 new host records and three confirmed earlier records (Table 10). Thus, 76% of the fungal species on Rosaceae collected were new to science. Fungi on Cornus Cornus (dogwood) is a genus of plants in the family Cornaceae. The genus comprises about 58 species, which are widely distributed in temperate and subtropical (rarely tropical) regions of the northern hemisphere, with a rich diversity in eastern Asia, eastern and western North America (Murrell 1993; Fan 2001; Xiang et al. 2006). In addition, some endemic species of Cornus are reported from South America and tropical Africa (Fan 2001). Members of Cornus are mostly trees and shrubs and rarely perennial herbs with woody rhizomes (Noshiro and Baas 1998; Fan 2001). While approximately 300 fungal species have been reported on Cornus species, only few of them have detailed illustrations and descriptions or are verified by DNA sequence data (Petrak 1921, 1925; Saccardo 1898; Senanayake et al. 2015; Wijayawardene et al. 2016). There is no comprehensive account or checklist of fungi on Cornus. 123 In considering these two examples it seems that the novelty of species in Europe is also surprisingly high. Our other studies on Clematis and several other hosts are also showing a remarkably high novel diversity. Hawksworth and Lücking (2017) estimated that there are 2.2–3.8 million fungal species in the world of which only 120,000 are presently known. Our studies appear to confirm these predicted high numbers. Novel chemistry Aside from the various taxonomic novelties, numerous new and unique biologically active secondary metabolites were also obtained from fungal species of northern Thailand. A selection of their chemical structures is depicted in Fig. 1. The producer organisms were often found to constitute undescribed species. Although not all new chemical structures have yet been named or their activity studied, the novel antibiotics terpene alkaloids named pyristriatins (Richter et al. 2016) were obtained from the novel species, Cyathus pyristriatus (Li et al. 2016) and the rare terpenoid lentinulactam from Panus subfasciatus (Hyde et al. 2016). The cytotoxic polyketides of the gymnopalyne type (Thongbai et al. 2013) and the mildly antibiotic deconins (Surup et al. 2015) were obtained from cultures of Gymnopus and Deconica, respectively, that probably represent new fungal species. The cultivated mushroom Lepista sordida (Thongbai et al. 2017b) yielded nudic acid B, and the nematode trapping species Hohenbuehelia grisea, produced the novel heterocyclic terpenoid pleurothiazol (Sandargo et al. 2018). Mycelial cultures of Agaricus subrufescens (Thongklang et al. 2017) produced the chemotaxonomic marker Blazeisporol A, which was identified as a selective agonist of Liver X Receptor subtype alpha, which offers avenues to the development of the mushroom as a new nutraceutical with chloesterol-lowering activities. Several new molecules were also obtained from species of Xylariales, which are one of the most creative orders of Ascomycota with respect to secondary metabolites (Helaly Fungal Diversity (2018) 93:215–239 Table 9 Foliar epiphytes reported in Thailand (novel species from Thailand are in bold) 227 Species Host References Asterina cynometrae Cynometra sp. Hyde et al. (2016) Asterina phlogacanthi Clinacanthus nutans Hyde et al. (2013) Asterina phoebesicola Phoebes costaricanae Hongsanan et al. (2014a) Capnodium coartatum Psidium guajava Chomnunti et al. (2011) Capnodium coffeicola Coffea sp. Hongsanan et al. (2015a) Ceramothyrium ficus Ficus sp. Hongsanan et al. (2015b) Ceramothyrium longivolcaniforme Unknown Zeng et al. (2016) Ceramothyrium thailandicum Lagerstroemia sp. Chomnunti et al. (2012a) Chaetocapnodium siamense Unknown Liu et al. (2015) Chaetothyrina artocarpi Artocarpus heterophyllus Hyde et al. (2017) Chaetothyrina guttulata Mangifera indica Hongsanan et al. (2016a) Chaetothyrina mangiferae Mangifera indica Singtripop et al. (2016) Chaetothyrina musarum Musa sp. Singtripop et al. (2016) Chaetothyrium bischofiicola Bischofia javanica Chomnunti et al. 2012a Chaetothyriothecium elegans Castanopsis sp. Hongsanan et al. (2014b) Conidiocarpus philippinensis Arecaceae sp. Liu et al. (2015) Conidiocarpus plumeriae Plumeria sp. Hongsanan et al. (2015a) Discopycnothyrium palmae Palm sp. Hongsanan et al. (2017) Irenopsis crotonicola Croton persimilis Zeng et al. (2018a) Irenopsis walsurae Walsura tubulata Hongsanan et al. (2015b) Lembosia albersii Unknown Hongsanan et al. (2014a) Lembosia xyliae Xylia sp. Ariyawansa et al. (2015) Leptoxyphium cacuminum Gossypium herbaceum Chomnunti et al. (2011) Meliola citri-maximae Citrus maxima Hyde et al. (2016) Meliola clerodendri-infortunati Clerodendrum infortunatum Hyde et al. (2017) Meliola clerodendricola Clerodendrum sp. Hyde et al. (2017) Meliola mucunicola Mucuna pruriens Hongsanan et al. (2015c) Meliola tamarindi Tamarindus indica Liu et al. (2015) Meliola thailandicum Dimocarpus longan Hongsanan et al. (2015c) Meliola thailandicum Acacia auriculiformis Hongsanan et al. (2015c) Micropeltis dendrophthoes Dendrophthoe sp. Hongsanan et al. (2015b) Muyocopron lithocarpi Lithocarpus lucidus Mapook et al. (2016) Parameliola acaciae Acacia auriculiformis Li et al. (2016) Parameliola dimocarpi Dimocarpus longan Li et al. (2016) Phaeosaccardinula ficus Ficus sp. Chomnunti et al. (2012a) Phragmocapnias asiaticus Coffea arabica Chomnunti et al. (2011) Phragmocapnias philippinensis Arecaceae sp. Liu et al. (2015) Phragmocapnias siamensis Mangifera indica Chomnunti et al. (2011) Scorias mangiferae Mangifera sp. Hongsanan et al. (2015b) Translucidithyrium thailandicum Syzygium levinei Zeng et al. (2018b) Trichomerium bambusae Poaceae sp. Hyde et al. (2016) Trichomerium deniqulatum Psidium guajava Chomnunti et al. (2012b) Trichomerium foliicola Murraya paniculata Chomnunti et al. (2012b) Trichomerium gloeosporum Ficus sp. Chomnunti et al. (2012b) Trichomerium gloeosporum Gardenia sp. Hongsanan et al. (2016c) Trichomerium siamense Tecoma sp. Liu et al. (2015) Trichopeltina asiatica Strobilanthes sp. Hongsanan et al. (2014c) Tumidispora shoreae Shorea sp. Ariyawansa et al. (2015) Zeloasperisporium ficicola Ficus benjamina Hongsanan et al. (2015d) Zeloasperisporium siamense Unknown Hongsanan et al. (2015d) Zeloasperisporium wrightiae Wrightia religiosa Hongsanan et al. (2015d) Expected number of new species: [ 100 123 228 Fungal Diversity (2018) 93:215–239 Table 10 Species of fungi on Rosaceae reported in the northern hemisphere (novel species are in bold) Species Host References Alternaria doliconidium Rosa canina Wanasinghe et al. (2018) Alternaria hampshirensis Rosa sp. Wanasinghe et al. (2018) Rosa sp. Wanasinghe et al. (2018) Amandinea punctataa Angustimassarina quercicola a Angustimassarina rosarum Rosa canina Wanasinghe et al. (2018) Rosa canina Wanasinghe et al. (2018) Bartalinia rosicola Rosa canina Wanasinghe et al. (2018) Bhatiellae rosae Rosa canina Wanasinghe et al. (2018) Broomella rosae Rosa canina Wanasinghe et al. (2018) Coelodictyosporium rosarum Rosa sp. Wanasinghe et al. (2018) Comoclathris rosae Rosa canina Wanasinghe et al. (2018) Comoclathris rosarum Comoclathris rosigena Rosa canina Rosa canina Wanasinghe et al. (2018) Wanasinghe et al. (2018) Coniochaeta baysunika Rosa sp. Wanasinghe et al. (2018) Coniochaeta rosae Rosa hissarica Wanasinghe et al. (2018) Dematiopleospora rosicola Rosa canina Wanasinghe et al. (2018) Diaporthe eresa Rosa sp. Wanasinghe et al. (2018) Diaporthe foeniculinaa Rosa canina Wanasinghe et al. (2018) Diaporthe rhusicolaa Rosa canina Wanasinghe et al. (2018) Diaporthe rosae Rosa sp. Wanasinghe et al. (2018) Diaporthe rosicola Rosa canina Wanasinghe et al. (2018) Diaporthe rudisa Rosa canina Wanasinghe et al. (2018) Diplodia seriataa Rosa canina Wanasinghe et al. (2018) Endoconidioma rosae-hissaricae Rosa hissarica Wanasinghe et al. (2018) Epicoccum rosae Rosa canina Wanasinghe et al. (2018) Keissleriella rosacearum Rosa canina Wanasinghe et al. (2018) Keissleriella rosae Rosa canina Wanasinghe et al. (2018) Keissleriella rosarum Lasiodiplodia theobromaea Rosa canina Rosa canina Wanasinghe et al. (2018) Wanasinghe et al. (2018) Lecidella elaeochromaa Rosa canina Wanasinghe et al. (2018) Lophiostoma rosae Rosa sp. Wanasinghe et al. (2018) Marjia tianschanica Rosa canina Wanasinghe et al. (2018) Marjia uzbekistanica Cerasus tianschanica Wanasinghe et al. (2018) Melanodiplodia tianschanica Rosa ecae Wanasinghe et al. (2018) Monoseptella rosae Rosa sp. Wanasinghe et al. (2018) Rosa canina Wanasinghe et al. (2018) Muriformistrickeria rosae Muriformistrickeria rubi a Murilentithecium rosae Rosa canina Wanasinghe et al. (2018) Rosa canina Wanasinghe et al. (2018) Neoascochyta rosicola Rosa canina Wanasinghe et al. (2018) Neoconiothyrium rosae Rosa canina Wanasinghe et al. (2018) Neofusicoccum australea Rosa sp. Wanasinghe et al. (2018) Neopaucispora rosaecae Rosa ecae Wanasinghe et al. (2018) Neosetophoma rosarum Neosetophoma rosigena Rosa canina Rosa canina Wanasinghe et al. (2018) Wanasinghe et al. (2018) Wanasinghe et al. (2018) Paraconiothyrium rosae Rosa canina Paraphaeosphaeria michotiia Rosa canina Wanasinghe et al. (2018) Paraphaeosphaeria rosae Rosa canina Wanasinghe et al. (2018) Paraphaeosphaeria rosicola Rosa canina Wanasinghe et al. (2018) Pararoussoella rosarum Rosa sp. Wanasinghe et al. (2018) 123 Fungal Diversity (2018) 93:215–239 229 Table 10 (continued) Species Host References Parathyridaria rosae Rosa sp. Wanasinghe et al. (2018) Paraxylaria rosacearum Rosa sp. Wanasinghe et al. (2018) Phragmocamarosporium rosae Rosa canina Wanasinghe et al. (2018) Pleospora rosae Rosa canina Wanasinghe et al. (2018) Pleospora rosae-caninae Rosa canina Wanasinghe et al. (2018) Pleurophoma pleurosporaa Rosa sp. Wanasinghe et al. (2018) Poaceicola rosae Rosa canina Wanasinghe et al. (2018) Populocrescentia rosae Pseudocercospora rosae Rosa hissarica Rosa canina Wanasinghe et al. (2018) Wanasinghe et al. (2018) Pseudopithomyces rosae Rosa canina Wanasinghe et al. (2018) Pseudostrickeria rosae Rosa canina Wanasinghe et al. (2018) Sclerostagonospora rosae Rosa sp. Wanasinghe et al. (2018) Sclerostagonospora rosicola Rosa sp. Wanasinghe et al. (2018) Seimatosporium rosicola Rosa canina Wanasinghe et al. (2018) Seimatosporium rosigenum Rosa canina Wanasinghe et al. (2018) Seiridium rosarum Rosa canina Wanasinghe et al. (2018) Sigarispora cauliuma Rosa canina Wanasinghe et al. (2018) Sigarispora rosicola Rosa sp. Wanasinghe et al. (2018) Sporormurispora pruni Prunus erythrocarpa Wanasinghe et al. (2018) Suttonomyces rosae Rosa canina Wanasinghe et al. (2018) Teichospora rubriostiolataa Rosa multibracteata Wanasinghe et al. (2018) Uzbekistanica rosae-hissaricae Rosa hissarica Wanasinghe et al. (2018) Uzbekistanica yakutkhanika Wojnowicia rosicola Rosa hissarica Rosa canina Wanasinghe et al. (2018) Wanasinghe et al. (2018) Xenomassariosphaeria rosae Rosa canina Wanasinghe et al. (2018) Expected number of new species: [ 100 a New host records et al. 2018). As an example, the lenormandins (Kuhnert et al. 2015) constitute highly specific pigments of the stromata of the Hypoxylon lenormandii complex. Work on the secondary metabolism of fungi from northern Thailand is ongoing, and papers on additional new and interesting molecules are in preparation. For instance, even other groups of Ascomycota including the Diaporthales that are treated above, are also well-known to be extremely creative secondary metabolite producers (Chepkirui and Stadler 2017). Potential future avenues For decades, mycologists have estimated fungal species numbers using various criteria. Such estimates have ranged from 500,000 to almost 10 million species, with mycologists generally agreeing on 1.5–5 million (Hawksworth 1991; Hawksworth and Lücking 2017). In the most recent estimates, Hawksworth and Lücking (2017) suggested between 2.2 and 3.8 million fungal species and that only 120,000 (8%) have been described. Hyde (2001) suggested that the ‘missing fungi’ might be found in poorly studied countries and hosts, or poorly studied habitats or niches. Tedersoo et al. (2014) used DNA metabarcoding data from hundreds of globally distributed soil samples, and demonstrated that climatic factors, followed by edaphic and spatial variables constituted the best predictors of fungal richness and community composition at the global scale. Tedersoo et al. (2017) provided phylogenetic placement and principal niche analysis for [ 40 previously unrecognized fungal groups from global soil samples at the order and class level based on combined 18S (nSSU) and 28S (nLSU) rRNA gene sequences, and showed that within the fungal kingdom, tropical and non-tropical habitats were equally likely to harbor novel groups. In this paper, we have provided an insight to show where missing fungi could be found. From data accumulated to date, and with randomized sampling only in northern Thailand, the fungal diversity with new species recovered far exceeds our expectations. We should revisit 123 230 Fungal Diversity (2018) 93:215–239 HO N O H O O Nudic Acid B OH O NH Cl OH H HO Gymnopalyne B Lentinulactam O N O O O O O O H S O O HO H O OH OH Blazeispirol A Deconin C Pleurothiazol O O O H O OH H H O Pyristriatin B O O O HO N OH Lenormandin B Fig. 1 Chemical structures of selected biologically active secondary metabolites from Thai fungi our current sampling strategies to target more novel species and similar studies could be extended to other parts of Thailand and surrounding countries. For some groups, the proportion of new species from samples collected is well above 50%, but this number might still be an underestimate because we assume we have still understudied the areas in which we have collected the material. Our studies on speciose genera, such as Colletotrichum and Pestalotiopsis have revealed many new species and this means that despite the high number of already described species, there is still much to be discovered. Tibpromma et al. (2018) recovered novel cultured endophytic species from Pandanaceae, but certainly there are unculturable orphan fungal species, that could represent a reservoir of novel species with a panoply of unexploited bioactive compounds. How will this impact on our anticipated number of fungi? Possibly we can argue that previously simple blind sampling strategies and inadequate DNA sequence analyses, limit new species discovery. Our studies in northern Thailand have revealed more new species as the taxonomic assessment methods used became more reliable and substrates/environments sampled were strategic. There are obviously other major fungal groups that warrant investigation (e.g. aquatic fungi, entomopathogens, dung fungi, 123 edible mushrooms, mycorrizhae, and wood decay fungi) that are globally distributed, but poorly sampled in Thailand. There is a need to further sample unexplored habitats (e.g. extreme environments) and substrata. One research area that we have yet to incorporate is sequence-based fungal community analyses. This will undoubtedly reveal and unravel an astonishing diversity of novel species, but as OTUs (Hongsanan et al. 2018). The major research challenges, however, to decipher fungal diversity in largely unexplored regions require more personnel to undertake multifaceted approaches to recover, identify and conserve the potentially new species waiting to be discovered. Conclusion Huge advances have been made in the understanding of the fungi in northern Thailand using polyphasic approaches and considerable advances in arranging the classification of fungi at the higher levels (cf. Tedersoo et al. 2018) have been concluded. Many novel fungi have been inventoried for the region, but much work remains. For example, many more species in the edible genus Agaricus await description. In these relatively well known mushroom genera we Fungal Diversity (2018) 93:215–239 are finding that more than 93% of species collected are new to science. In the microfungi which appear to be relatively poorly studied, the percentage does not appear to be as high. The studied regions mainly includes three provinces in northern Thailand. The southern, eastern and central provinces of Thailand and surrounding countries of Cambodia, Myanmar, Laos and Vietnam have barely been studied for fungi and thus we predict that there are huge numbers of new species waiting to discovered in this region. At the same time, we have been finding ways to exploit these fungi. Our work has resulted in the discovery of at least ten new species which are being developed as novel industrial mushrooms. We have also isolated at least ten novel medicinal compounds from Thai fungi and are also looking at ways to exploit them in biocontrol. All of the fungi mentioned above are known to produce various therapeutic metabolites with high biological activities. It is therefore very important to properly characterize not only these compounds, but to carefully resolve the species names, so that researchers can better identify and screen potential taxa for future biotechnological applications. Fungi have been poorly exploited and yet have a huge potential in biocontrol, bioremediation, novel compound discovery as well as basic industrial organisms as edible mushrooms, fertilizers and cosmetics. With such high novelty, there is a need for extensive research to exploit the biotechnological potential of these fungi. Acknowledgements K.D. Hyde would like to thank the Thailand Research Fund for the grant ‘‘Domestication and bioactive evaluation of Thai Hymenopellis, Oudemansiella, Xerula and Volvariella species (basidiomycetes)’’ Grant No. as : DBG6180033 for funding this work. Mae Fah Luang University for the grant ‘‘Taxonomy diversity, phylogeny and evolution of fungi in Capnodiales’’ (Grant No: 666713), for supporting this study. The authors extend their appreciation to the German Academic Exchange Service (DAAD) for a joint TRF-DAAD [PPP 2017–2018] academic exchange grant to K.D. Hyde and M. Stadler. Mingkwan Doilom thanks grant for postdoctoral researchers funded by Yunnan Human Resources and Social Security Department, and The 64th Grant from China Postdoctoral Science Foundation. References Abiala MA, Popoola OO, Olawuyi OJ, Oyelude JO, Akanmu AO, Killani AS, Osonubi O, Odebode AC (2013) Harnessing the potentials of vesicular arbuscular mycorrhizal (VAM) fungi to plant growth—a review. 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Fungal Divers 51:279–296 Zhao RL, Desjardin DE, Callac P, Parra LA, Guinberteau J, Soytong K, Karunarathna S, Zhang Y, Hyde KD (2012a) Two species of Agaricus sect. Xanthodermatei from Thailand. Mycotaxon 122:187–195 Zhao RL, Hyde KD, Karunarathna SC, Desjardin DE, Raspé O, Soytong K, Guinberteau J, Callac P (2012b) Agaricus flocculosipes sp. nov., a new potentially cultivatable species from the palaeotropics. Mycoscience 53:300–311 Zhao RL, Zhou JL, Chen J, Margaritescu S, Sánchez-Ramı́rez S, Hyde KD, Callac P, Parra LA, Li GJ, Moncalvo JM (2016) Towards standardizing taxonomic ranks using divergence times—a case study for reconstruction of the Agaricus taxonomic system. Fungal Divers 78:239–292 Zhou JL, Su SY, Su HY, Wang B, Callac P, Guinberteau J, Hyde KD, Zhao RL (2016) A description of eleven new species of Agaricus sections Xanthodermatei and Hondenses collected from Tibet and the surrounding areas. Phytotaxa 257:99–121 Affiliations Kevin D. Hyde1,2 • Chada Norphanphoun2 • Jie Chen3 • Asha J. Dissanayake2 • Mingkwan Doilom1,4,5 • Sinang Hongsanan6,7 • Ruvishika S. Jayawardena2 • Rajesh Jeewon8 • Rekhani H. Perera2 • Benjarong Thongbai10 Dhanushka N. Wanasinghe1,4 • Komsit Wisitrassameewong9 • Saowaluck Tibpromma1,2,4 • Marc Stadler10 1 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Science, Kunming 650201, Yunnan, People’s Republic of China 123 2 Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand • Fungal Diversity (2018) 93:215–239 3 Instituto de Ecologı́a, A.C., CP 91070 Xalapa, Veracruz, Mexico 4 World Agro Forestry Centre, East and Central Asia, 132 Lanhei Road, Kunming 650201, Yunnan, People’s Republic of China 239 7 Shenzhen Key Laboratory of Laser Engineering, College of Optoelectronic Engineering, Shenzhen University, Shenzhen, People’s Republic of China 8 Department of Health Sciences, Faculty of Science, University of Mauritius, Reduit 80837, Mauritius 5 Department of Biology, Faculty of Science, Chiang Mai University, Chiang Rai 50200, Thailand 9 National Science Technology and Innovation Policy Office, Bangkok 10330, Thailand 6 Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Sciences and Oceanography, Shenzhen University, Shenzhen, People’s Republic of China 10 Department of Microbial Drugs and German Centre for Infection Research (DZIF), partner site HannoverBraunschweig, Helmholtz Centre for Infection Research, Inhoffenstrasse 7, 38124 Brunswick, Germany 123