R EVIEW ARTICLE ScienceAsia 46S (2020): 1–11 doi: 10.2306/scienceasia1513-1874.2020.S001 The rise of in Asia

Kevin D. Hydea,b, K.W.T. Chethanaa, Ruvishika S. Jayawardenaa, Thatsanee Luangharna,c, a,d e,f g,h i, Mark S. Calabon , E.B.G. Jones , Sinang Hongsanani , Saisamorn Lumyong ∗ a Center of Excellence in Fungal Research and School of Science, Mae Fah Luang University, Chiang Rai 57100 Thailand b Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Guangzhou 510225 China c Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201 China d Research Foundation, Mae Taeng, Chiang Mai 50150 Thailand e Department of Botany and , College of Science, King Saud University, 11451 Saudi Arabia f Nantgaredig, 33B St Edwards Road, Southsea, Hants., PO5 3DH, UK g Shenzhen Key Laboratory of Laser Engineering, College of Physics and Optoelectronic Engineering, Shenzhen University, Shenzhen 518060 China h Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Sciences and Oceanography and Shenzhen University, Shenzhen 518060 China i Center of Excellence in Microbial Diversity and Sustainable Utilization, Faculty of Science, Chiang Mai University, Chiang Mai 50200 Thailand

∗Corresponding author, e-mail: [email protected] Received 13 Mar 2020 Accepted 26 Mar 2020

ABSTRACT: Mycology was a well-studied discipline in Australia and New Zealand, Europe, South Africa and the USA. In Asia (with the exception of Japan) and South America, the fungi were generally poorly known and studied, except for the result of forays from some American and European mycologists. However, in the last 20 years, the situation has changed. With the development of Asian economies, the funding for science research and development has greatly increased. Mycological research has also diversified in many fields. Many studies have focused on applied aspects and new journals and websites have been established as a platform for Asian mycologists to publish their research. This paper will briefly review the history of the study of fungi in Asia and then discuss how it advanced during the last two decades. It will examine the current situation using case studies in plant pathogens, terrestrial saprobes, aquatic fungi, evolution studies, genomics and applied mycology and biotechnology. Finally, it will suggest research that is needed in the future.

KEYWORDS: Asian mycology, fungal diversity and evolution, genomics and applied mycology, mycological websites and journals

INTRODUCTION soil, mosses, man-made materials and or associated with other organisms [3]. According to the state of The diversity of fungi is immense and yet less than world fungi report in 2018, the estimated number 10% have been described [1,2 ]. Fungi have unique of fungal species on earth is between 2.2–3.8 mil- attributes and are essential for the well-being lion [4]. Due to the increased use of molecular- of as decomposers, mutualists, and based techniques, specifically DNA-based, the rate pathogens. They also have huge biotechnological of description of new fungal species has increased to potential in agriculture (biocontrol agents, biofertil- more than 2000 species per year. These discoveries izers, growth-promoting hormones and pathogens), represent taxa mainly from Ascomycota, followed and as sources for many other industries such as by Basidiomycota. In 2017, the majority of new biofuel, beverages, cosmeceuticals, food, mycore- fungal species were recorded from Asia, accounting mediation, and pharmaceuticals [3]. Fungi being for 35%, followed by Europe, Oceania, South Amer- cosmopolitan, grow almost everywhere; mostly in ica, North America, and Africa [4]. These numbers terrestrial and aquatic habitats, and even in extreme reflect the areas where most fungal taxonomists environments, in deep sea vents, on rocks, plants,

www.scienceasia.org 2 ScienceAsia 46S (2020) work. For example, among these, the majority of the (e.g., Thailand). The development of mycology in records were from China, Australia ranks second, Thailand and Malaysia has been documented by followed by Thailand [4]. Asian mycologists are Jones et al [6,8 ]. In Thailand, the enthusiasm taking the initiative to actively participate in the on- of M. Tantichareon in establishing workshops and going quest to explore fungi and their applications. annual conferences and many international confer- Most Asian regions have tropical and subtropical ences, did a great deal to stimulate investigations climates, where fungi thrive due to the favourable of tropical fungi. Tzean et al [9] published five temperatures and humidity. Therefore, this region volumes on the Fungi of Taiwan with descriptions has a rich fungal diversity. In recent years, many and excellent illustrations thus contributing to our young mycologists and fungal taxonomists have knowledge of this region, and establishing an active been trained in Asia and they are providing sig- Mycological Society. nificant contributions to improve mycology in their During the latter part of the 1900s, Asia estab- respective countries. lished societies to promote mycology. For example, This paper briefly reviews the history of mycol- the Indian Mycological Society was established in ogy in Asia before the 20th century and addresses 1954. Among the 49% of the new fungal species the current status of mycology. We provide case (including fossils) introduced from 1981–1990 by studies in plant pathology, aquatic fungi, terrestrial the tropical countries, 9.7% were contributed by saprobes, evolution studies, genomics and applied India and was ranked second to the USA, which mycology and biotechnology. We will suggest future contributed 10.1% of new species [10]. China, areas of research needed in mycology in Asia. being the second largest country in Asia, carries a significant history of mycology which runs back to Mycology before the 20th century in Asia 1245 [4, 11] when the first monograph of Chinese Half of the worlds’ population lives in Asia, but fungi was published with 15 fungal species. During many countries were economically disadvantaged the 19th century, several projects such as the Com- and lacked resources for establishing laboratories prehensive Scientific Expeditions in the Qinghai- for fungal research. Taxonomic studies of Asian Tibet Plateau (1973–1983), the Bioresources Sur- fungi were limited because of reliance on overseas vey and Evaluation (1988–1990) and many more expertise, lack of local centers, language and a poor individual investigations advanced the knowledge understanding of fungi by the public. In most of in the region. However, until the beginning of the Asia (except Japan), fungal studies were carried 21st century, fungal species identification in China out by visiting scientists or expatriate experts in as well as in most parts of Asia was based mainly the 18th and most of the 19th centuries. For ex- on morphological characters and sometimes on the ample, an Irish mycologist, E.J. Butler contributed geographical distribution [12]. The development significantly to establish the foundation for mycol- of mycology in the region was therefore limited. ogy and plant pathology in India [5]. Visiting Furthermore, Thailand, one of the more active coun- mycologists M.J. Berkeley, C.E. Broome and T. Petch tries for fungal research, mainly focused on ap- contributed significantly to mycology in Sri Lanka. plied mycology [13]. Much research was related to Their contributions increased the recorded number agriculture [13], while others focused on industrial of fungi to over 2000 described species belonging to applications, such as mushroom cultivation, citric 640 genera. The eminent botanist and mycologist, acid soy sauce, alcohol fermentation, sweetened rice E.J.H. Corner made remarkable contributions to the production, pigment production and biodeteriora- South-East Asian macrofungi, mainly in Malaysia tion [13]. Due to an increased interest and impor- and Singapore [6]. Most mycological laborato- tance of mycology in the region, the International ries employed conventional techniques, resulting in Mycological Association established a committee for lower numbers of novel discoveries and publica- Asia in 1977, later known as the Asian Mycological tions. These reasons challenged the establishment Association to promote mycology. of a successful mycological community in most of With the rapid development of DNA based tech- Asia. Due to isolation from the world, niques, molecular phylogenetic studies have ad- Japanese mycologists and phytopathologists, how- vanced rapidly in Asian laboratories and used to ever, published accounts of Japanese fungi during study species diversity and systematics of many this period [7]. With the help of visiting scholars, fungal groups [14–17]. Incorporation of molecular however, several laboratories were set up in some data into fungal research has allowed the extension parts of Asia in the latter part of the 19th century of studies into diverse areas of life science, including www.scienceasia.org ScienceAsia 46S (2020) 3 ecology, traditional taxonomy, phylogenetics, evo- marinefungi.org), all providing illustrations and lution, microbial community and chemotaxonomy, plates of fungal species and genera. The websites and mushroom cultivation [15, 18]. With the devel- provide up-to-date outlines and accounts on orders, opment of next-generation sequencing in the Asian families, genera, and species levels, keeping abreast region, specifically in China, many mycologists have of the current literature. Other websites are: www. adopted fungal genomics and transcriptomics ap- onestopshopfungi.org which provides a stable plat- proaches into their studies. This has opened new form for the taxonomy of plant pathogenic fungi opportunities for mycologists to study the evolution and related organisms. It combines all molecular or co-evolution of species, natural products discov- data (phylogenetic analyses, DNA sequence acces- ery, biological control, medical mycology (disease sion numbers), recommendations on correct names, mechanisms and virulence factors), host-fungal in- data on type material, geographical and ecological teractions, bioremediation, bio-fuel production, and observations, ecological and economic significance, genetic engineering of fungal strains for industrial industrial relevance, and literature into a compre- purposes [19]. As a result, many publications in the hensive and uniform treatise, so that fungal taxa can above-mentioned fields came from the Asian region, be identified. The website www.facesoffungi.org especially from China [20, 21]. provides descriptions, photoplates, keys for identi- China, being the largest country in the region, fication, phylogenetic analyses, sequence accession has made significant advances in mycology. A large data and ecological and economic significance of number of studies have been conducted in China in registered taxa. The website www.outlineoffungi. the 20th century, resulting in more than 6700 fungal org provides an up-to-date classification for all fungi species [4, 22]. China is the world’s largest edible and fungi-like organisms, while www.fungalgenera. mushroom producer with an estimated annual yield org provides a short account of all accepted fungal of 38.42 million tons, accounting for around 75% of genera of Ascomycota, Basidiomycota, and Lower the global demand [4]. With the increased demand, Fungi with links to data. research on the mushroom industry has accelerated in China. This has influenced other countries to Mycological journals in the Asian region focus on the mushroom related research [4]. With the increased interest in mycology, there are The increased interest in mycology in the region about 34 mainstream journals which publish myco- has also led to an increase in journals published out logical research in 2011. Besides these, several re- of Asia (Table S1). The number of publications in gional journals, plant pathology journals and many taxonomic mycology from Asian in 1999 was 117, bulletins of national mycological societies publish with an estimated 32 active taxonomic mycologists. research articles. Many new mycological journals These numbers have rapidly increased until the have been introduced from the Asian region and present day. China, Japan and Thailand have taken some have SCI recognition. The mainstream peer- the lead in publishing in the Asian region with 427, reviewed journals devoted to mycology are given in 125 and 94 SCI articles published in 2019. India, Table S1. Taiwan and South Korea have also contributed to a lesser extent. Fungal research has increased greatly Major advances by Asian mycologists in Thailand, with Mae Fah Luang University ranked There has been a great increase in mycological first in the world for Mycology by CWUR World research and publications from Asian institutes and University Rankings 2019 (https://cwur.org/2017/ universities, while formerly strong research groups subjects.php#Mycology). were found in Japan and Taiwan. Prominent examples are Beijing Forestry University (1955– Website development present), BIOTEC, Bangkok (1995–present), Chiang Online databases serve as a repository for pub- Mai University (1996–present), The University of lished data and several are being developed in Asia. Hong Kong (1994–2008), Mae Fah Luang Univer- These websites are dedicated to updating the tax- sity, Chiang Rai (2009–present), Institute of My- onomy, classification or distribution of fungi. Cur- cology & , CAS, Beijing (1988–present), rently, four webpages deal with different taxonomic Kunming Institute of Botany (1989–present) and groups (www.basidio.org, www.coelomycetes.org, National Taiwan Ocean University (2008–present). www.dothideomycetes.org, www.sordariomycetes. Dai et al [23] showed that many more fun- org), while two webpages are dedicated for ecology gal taxa were being described from China, while and taxonomy (www.freshwaterfungi.org, www. Hyde et al [24] showed that the fungal diversity in

www.scienceasia.org 4 ScienceAsia 46S (2020) northern Thailand was remarkably high, with up to morphology alone. Later, it was established that 93% of some genera (e.g, ) being new to this concept can be used for all the complex gen- science. era, leading to a new process to understand these Asia has been leading the way in many areas of complex fungi. Following the same concept, Cai and mycology and in particular in understanding fungal colleagues advanced the understanding on other classification. Some of the most significant pa- plant pathogenic genera and recommended genes pers are Families of Dothideomycetes [25], Taxonomy to provide a better resolution. Maharachchikum- and phylogeny of hyaline-spored Coelomycetes [26], bura et al [38] took the initiative to identify the Notes for genera: Ascomycota [27], Outline of As- morphology, phylogeny and biochemistry of an im- comycota: 2017 [28], Notes, outline and diver- portant plant pathogenic genus, Pestalotiopsis. They gence times of Basidiomycota [29], Refined fam- tested ten gene regions to establish the best gene ilies of Sordariomycetes [30], Notes for genera: combination (ITS-Tub2-TEF1) for a better species basal clades of fungi (including Aphelidiomycota, Ba- resolution. Later, they segregated two novel gen- sidiobolomycota, Blastocladiomycota, Calcarisporiel- era from Pestalotiopsis, namely Neopestalotiopsis and lomycota, Caulochytriomycota, Chytridiomycota, En- Pseudopestalotiopsis based on molecular and mor- tomophthoromycota, Glomeromycota, Kickxellomy- phological approaches [41]. Udayanga et al [40] cota, Monoblepharomycota, Mortierellomycota) [31] assessed eight gene regions to establish the best and Outline of the fungi and fungi-like taxa [32]. gene combination for resolving Diaporthe species. These texts are incredibly important to students and In their study, the Genealogical Concordance Phylo- young mycologists, providing classifications of most genetic Species Recognition (GCPSR) concept was groups of fungi. The various series, Fungal Diversity applied to resolve species boundaries based on notes and Mycosphere notes with 50 or more taxa, individual and combined analyses of seven genes many new to science, treated in each paper and also (except ITS), resolving the D. eres species com- mostly by Asian mycologists. plex. Manamgoda et al [39] revised the boundaries The fungal diversity of China, Malaysia and between Bipolaris and Curvularia. In their study, Thailand, where the main research has taken place, they accepted 47 species in Bipolaris and clarified are still poorly known. In Malaysia, Chip [33] list the taxonomy, host associations, geographic distri- 861 fungi, while Lee et al [34] list circa 3000. butions and species’ synonymies. Another major Thailand has more than 6000 known species of contribution of Asian mycologists to the field of fungi and between 2009–2020 Hyde and cowork- plant pathology is the starting of a paper series one ers have described 700 new species mostly from stop shop. Hyde et al [42] initiated this concept to northern Thailand. Yet collections have been from provide all the information, such as history of the many new habitats, hosts or areas, with numerous disease, symptoms, life cycle, on plan pathogenic new species documented, including those in new fungal genera in one place which has provided easy genera, families and orders [4, 24]. The bottleneck access to plant pathologists to all the relevant infor- in describing these new species is the lack of stu- mation on a specific plant pathogenic genus. OSS2 dents for the necessary detailed morphological and and OSS3 detailing 50 pathogenic genera have since phylogenetic studies and ability to obtain molecular been published [43]. data. Case studies in plant pathogens, terrestrial Asian mycologists have contributed enormously saprobes, aquatic fungi, evolution studies, genomics in the establishment of a stable taxonomy of the and applied mycology and biotechnology, where pathogenic genus Colletotrichum. Hyde et al [36] Asians have advanced mycology, are provided. provided the first ever most comprehensive overview of the genus with 66 common names Plant pathogens in use and 19 doubtful names. Another article Asian mycologists have contributed a great deal published by the same author highlighted the to the understanding of plant pathogenic genera importance of revising the genus by using molecular during the past two decades. Many of these mycolo- methods. These studies are the foundation of the gists worked on complex pathogenic genera, such as still ongoing revision of the genus based on multi- Bipolaris-Curvularia, Colletotrichum, Diaporthe and locus sequence data. Cai et al [35] introduced the pestaloid taxa, establishing guidelines for species use of polyphasic approaches in identifying this delimitation [35–40]. Cai et al [35] proposed that in group of fungi. Mycological groups led by K.D. order to correctly identify Colletotrichum to species Hyde and L. Cai have introduced many species level, a polyphasic approach is better than using of Colletotrichum based on not only concatenated www.scienceasia.org ScienceAsia 46S (2020) 5 loci, but also with the application of GCPSR. Table 1 Fungal taxonomic novelties from Asian aquatic Jayawardena et al [44] provided an account environment. of Colletotrichum accepting 190 species, with a Orders/Families Genera Species backbone phylogenetic tree and notes for each Orders Halodiatrype Bacusphaeria nypae species. Distoseptisporales Striatiguttula Fusicolla bharatavarshae Dyfrolomycetales Bacusphaeria Halodiatrype avicenniae Diaporthe is another plant pathogenic genus Jahnulales Thyridariella Striatiguttula nypae that the Asian mycologists have contributed im- Families Halotestudina Thyridariella mangrovei Striatiguttulaceae Gesasha mensely to clarify the taxonomy. Through his Distoseptisporaceae studies, Udayanga et al [40] contributed in under- Ceratosphaeriaceae Triadelphiaceae standing species delimitation in this genus. Many Dyfrolomycetaceae Asian mycologists also contributed in identifying the species on economically important plants and forest plants 45 . Gao and collaborators showed that this [ ] paper 55 which reported 2344 strains belonging genus is paraphyletic. Dissanayake et al 45 pro- [ ] [ ] to 610 species in 253 genera and introduced 33 vided accounts for 171 accepted species with a back- new species. Mapook et al 56 described the fungi bone tree for the genus. Manawasinghe et al 46 [ ] [ ] on Siam weed, a species from Central America. contributed to the understanding of this genus Remarkably, one family, 12 genera and 47 species by studying the high genetic diversity and species were new to science and must have jumped to this complexity of Diaporthe associated with grapevine host from surrounding hosts. Preliminary studies on dieback. the fungi on seeds and fruits resulted in eight new Botryosphaeriaceae is a family of plant genera and 65 new species 57, 58 . In fact, the pathogenic fungi and many Asian mycologists [ ] fungi on most terrestrial hosts in Asia have barely have contributed to the understanding of the been studied and we predict that tens of thousands genera. Liu et al 47 used a phylogenetic approach [ ] of species await discovery. together with a study of herbarium specimens and provided detailed accounts of the accepted genera Aquatic mycology in this family. Dissanayake et al [48] provided a backbone phylogenetic tree for the family and notes The majority of the research on aquatic fungi in on accepted genera and species. recent years has been from Asia and studies have incorporated molecular techniques to achieve a bet- Terrestrial saprobes ter classification of taxa. As a result, over 500 new Dead plant material is mainly decomposed by fun- species and some new genera, families and orders gal saprobes including ascomycetes, basidiomycetes (Table 1) have been introduced [15, 27, 28, 30, 54]. and basal fungi. They are essential for nutrient The Greater Mekong Subregion has been relatively cycling and are the major decomposers of lignin [3]. well-explored for freshwater fungi [16, 59]. Most Asia has provided major advances in describing of the studies follow the direct observation of fruit- many of its fungi. Hyde et al [24] showed that 96% ing bodies on different substrates and use culture- of Amanita and 93% of Agaricus species collected dependent techniques [16]. There are few reports in mainly northern Thailand were new to science. on the unculturable fungi in freshwater habitats in Other important contributions were monographs Asia [61], hence, culture-independent techniques of Boletales [49] and Amanita [50] and studies need to be followed to understand the true diver- on Polyporaceae [51] and advances in the genus sity among freshwater fungi in Asia. For example, Agaricus. However, perhaps the most important Kagami et al [61] analyzed the planktonic fungi advance was the notes and an outline to all Basid- in Lake Inba, Japan using denaturing gradient gel iomycota [29]. electrophoresis and revealed Chytridiomycota as the The study of microfungi is, however, in its in- dominant group, followed by Basidiomycota and fancy. Research on the microfungi on bamboo [52], Ascomycota. However, it is not clear whether the teak [53] and Pandanaceae [17] provides prelim- resulting operational taxonomic units (OTUs) have inary knowledge, but certainly did not describe any role or are present. A world monograph all taxa from these hosts. Zhang et al [54] re- of all freshwater Sordariomycetes was published ported 246 species in 116 genera from two unnamed by Luo et al [16] and will be followed by one on Karst cave in Guizhou, China, including 20 new Dothideomycetes. species. This first study is followed by a second The study of in Asia has been

www.scienceasia.org 6 ScienceAsia 46S (2020) prolific and has predominantly been led by EBG. port from evolutionary evidence. Despite the recent Jones who worked in BIOTEC, Thailand, Univer- molecular clock dating of the Fungi [75] studies sity of Malaya, Malaysia and other Asian regions. on co-evolution and evolution of fungal characters, Marine environments in Asia have been a genesis further studies are needed. of many novel fungal taxa [62, 63], with molecular data being included in most studies. Countries with reports of new species of marine fungi over the past decade (2010–2019) are India [64], Japan [65, Genomics 66], Malaysia [67], Saudi Arabia [68, 69] and Thai- land [63]. As a highpoint to his life’s work, EBG. The genome provides a glimpse into the molecular Jones has developed a website (www.marinefungi. basis of fungal biological diversity. Genome se- org) on marine fungi with his Asian colleagues [70]. quencing of fungi represents a significant milestone Besides taxonomic studies, marine fungi have also in mycology. Accelerated progress in sequencing been studied as a source of biologically active com- technologies and bioinformatics enable access to pounds [71, 72]. genomes at rapidly reducing costs. Therefore, initia- tives have been launched to sequence a wide range Evolution studies of fungal genomes [76]. Developments in the field Knowledge of fungal evolution helps to provide an of genomics have paved way for comparative ge- understanding of the diversity of life as similarities nomics; this, in turn, benefits many fields including between species/higher-level taxa occur as heritable agriculture and biotechnology. In Asia, fungal ge- changes over time and in populations. Fungi in nomics has received much attention from scientists Asia are clearly highly diverse, especially in tropical in China, Japan, Korea, India, Iran and Thailand, Asia [24]. Although the numerous fungi discovered where considerable investments in fungal genomics in Asia are morphologically and phylogenetically research have been made. China has taken the well-studied, there have been relatively few evolu- lead in Asia with many laboratories initiating fun- tionary studies. Mycologists have applied evolution- gal genomic projects [19], which have accelerated ary study to understand fungi in different research the pace of mycological research in China. Asia, disciplines. For example, the study of ancestral in particular China, has focused on economically characters not only explains the evolution of mor- important fungi in many genome projects. The phology in fungal groups, but can also estimate landscape of Asian genomic and functional genomic the probability of ancestral habitat or nutritional projects have concentrated on areas, such as plant modes. Co-evolution of plant- relationships and human pathogenic fungi [21], fungal-host in- visualize how fungal species are ecologically linked, teractions [77, 78], fungi as biocontrol agents [79], and how they co-evolved to survive in a specific fungi as food [80] and [20, 81]. time period [73]. The study of fungi which are Asian large scale sequencing bodies, such as Bei- resistant to fungicides through evolution is ongoing jing Genomics Institute (China), Macrogen (Korea), and may lead to methods to resolve this issue [74]. Chun Lab (Korea), Softbank (Japan), Temasek (Sin- These studies are needed because of the extensive gapore), Prenetics (Hong Kong), Xcode (India) and use of fungicides in Asia. According to evolutionary Genomico (Iran), have also been collaborating with changes over time, which gives rise to new species, large scale fungal genomic projects. The rapid fungal evolution with divergence time estimation development of bioinformatics infrastructures has has been used for classification and ranking of fungi facilitated a boom in Asian fungal genomics in the at higher levels [14, 75]. In these analyses, it is region [19, 82]. In addition, several mycological necessary to include a starting point for any molec- breakthroughs have been published by Asian scien- ular dating, paleoecological re-construction, and tists, such as massive decoding of non-model fun- morphological character evolutionary study [18]. gal genomes [19, 20] and successful application of Hence, reliable fossils information of fungi is needed Agrobacterium-mediated fungal transformation and to define the boundary of estimation [18], and more CRISPR/CAS9 systems [83]. While China leads sequence data of novel species are requested to the way, many governments and private sectors in accurate analyses. Asia are investing billions of dollars into genetics Fungal evolution has become clearer with var- research. As genomics takes a firm hold in Asia, ious Asian studies [15, 56, 62]. This is being im- regulators will need to establish standards on the proved by the introduction of higher taxa with sup- genetic data management. www.scienceasia.org ScienceAsia 46S (2020) 7

Applied mycology and biotechnology cesses such as mycoremediation, mycofumigation, Applied mycology is a discipline where we use the and fungal degradation of plastics and polycyclic full potential of fungi for industrial exploitation. aromatic hydrocarbons [3]. Studies to assess the Initially, applied mycology mainly focused on agri- genetic manipulation of these processes are being culture or where fungi caused damage. However, conducted. Asian industrial production of fungal- after several decades significant developments were mediated biofuels, organic acids, cosmeceuticals, made globally towards using fungi for new pro- enzymes, and textile dyes are commonplace, but studies are needed to improve these processes 3 . cesses, and environmental-friendly products [3]. [ ] The trend for fungal genetic research has moved Knowing the gene and protein function of ben- from genes to genomics, to functional genomics. In eficiary fungal traits has boosted the fungal biotech- Asia, research on functional genomics of economi- nology research arena. Bioengineering has dra- cally important fungal taxa has accelerated and this matically improved yield, quality, resistance and resulted in the accumulation of genetic informa- the nutritional value of foods and also facilitates sustainable agriculture. Asia scientists have stud- tion [77, 78]. The data are currently being used to enhance industrial processes and obtain value- ied the heterogeneous expression of fungal sec- added products, such as bulk manufacturing of or- ondary metabolite gene clusters [90], enabling the ganic acids, proteins, enzymes, secondary metabo- biosynthesis of antibiotics and natural products [3]. Applied mycology will continue to be the direct lites and active pharmaceutical ingredients [3]. A common application of mycology in Asia is for beneficiary of future advances in fungal genomics agricultural purposes. These applications can vary and fungal biotechnology. The progress achieved from using fungi as biocontrol agents, biofertilizers, by Asia scientists in transferring basic mycology mitigating abiotic stresses in plants and the pro- knowledge to applied mycology, and biotechnolog- duction of biofungicides, insecticides, nematicides, ical applications during the last two decades has been impressive. herbicides and growth-promoting hormones [3, 84, 85]. Furthermore, fungal genes have been genet- The future ically engineered into plants to produce resistant Where does the future lead us? With tens of varieties [86]. Owing to their antibiotic, antimy- cotic, antiviral, anticancer, antidiabetics and im- thousands of new species awaiting collection and munosuppressive properties, there have been many description, there is so much research needed. How- studies in Asia to assess the potential of fungi in the ever, the future challenge for the mycologists is not only to discover novel fungal taxa and understand pharmaceutical industry [3, 87]. In addition, fungi have been used in Traditional Chinese Medicine and their evolutionary relationships, but also discover this practice is being used in Japan and other parts potential diversities of dark taxa hidden in fungal sequences and place them into the tree of life. The of Asia [3]. Domestication and industrial production of knowledge gathered from these studies provides occurs across Asia. China has the great- exciting opportunities for the molecular mycologists est global demand for food including mushrooms. to predict fungal properties that can be exploited Mushroom production in Asia has changed from further. Such research may be basic, but it will manual to automated and also, novel hybridized provide us with the resources and tools for numer- strains are being developed mainly in China and ous applications. For example, new edible species of mushrooms can be commercialized and increase Thailand [4]. Studies are also being conducted to genetically engineer or manipulate mushrooms the variety of foods in the market. Medicinal mush- rooms with anti-cancer and anti-diabetic properties for higher yields and longer shelf-lives [88, 89]. Apart from these applications, fungi are vastly used can be commercialized and sold dried or fresh or in Asian food and beverage industries as food packaged in teas or powdered capsules. The novel colourants, bio-flavours, probiotics and in wine, microfungi are likely to span across the fungi and thus provide a powerhouse of metabolic beer and spirit production [3]. Furthermore, fungi have been used as a sustainable solution for many pathways that can be utilized in the industry as biofertilizers, biofuels, enzymes, organic acids, pro- global problems [3]. For example, in many parts of Asia, agricultural waste has been recycled as teins, secondary metabolites, textile dyes, and active substrates for mushroom production. In addition, pharmaceutical and cosmeceutical ingredients. The fungi or their products have been utilized in pro- aim of future fungal research should be to convert the results of these basic sciences to practical appli-

www.scienceasia.org 8 ScienceAsia 46S (2020) cations. ways we can exploit fungi industrially. Fungal Divers The accelerated pace of fungal genomics re- 97, 1–136. search and its development in the region has pro- 4. Willis KJ (2018) State of the World’s Fungi 2018, vided genomic resources for more than 1000 fungal Royal Botanic Gardens, Kew. species and resulted in a huge amount of data. 5. Subramanian CV (1986) The progress and status of Even though fungal genomics evolved rapidly, very mycology in India. Proc Plant Sci 96, 379–392. 6. Jones EBG, Hyde KD, Vikineswary S (2007) few biotechnological developments resulted from Malaysian Fungal Diversity, Mushroom Research omics sciences. The omics technologies such as Centre, University of Malaya and Ministry of Natural functional genomics, transcriptomics, proteomics, Resources and Environment, Malaysia. metabolomics and interactomes are currently un- 7. Hosoya T (2006) History and the current status of dergoing a development phase. Future studies us- fungal inventory and databasing in Japan. Nat Sci ing omics technologies should reveal information Mus Monogr 34, 51–61. related to fungal adaptations, evolution, industri- 8. Jones EBG, Tantichareon M, Hyde KD (2004) Thai alizations and fungal interactions. The future of Fungal Diversity, BIOTEC, Thailand. Asian mycology depends on innovative studies fo- 9. Tzean SS, Hsieh WH, Chang TT, Wu SH (2005) cused on enhancing fungal metabolic processes or Fungal Flora of Taiwan, National Science Council, engineering industrially important fungi for maxi- Taiwan. mum production. For mycology to thrive in this 10. Hawksworth DL, Rossman AY (1997) Where are all region, collaborations among Asian mycologists and the undescribed fungi? Phytopathology 87, 888–891. 11. Zang M, Li X-J (2008) Studies on fungi and improvements for bioinformatics infrastructures are bryophytes from Yunnan in the last one hundred necessary. years–review and prospect. Acta Bot Yunnan 30, Appendix A. Supplementary data 382–326. 12. Dai YC (1999) Phellinus sensu lato (Aphyllophorales, Supplementary data associated with this arti- Hymenochaetales) in East Asia. Acta Bot Fenn 166, cle can be found at http://dx.doi.org/10.2306/ 1–115. scienceasia1513-1874.2020.S001. 13. Suwanarit P (1986) Mycological research in Thai- land. Proc Plant Sci 96, 343–355. Acknowledgements: We thank the Thailand Research 14. Hyde KD, Norphanphoun C, Abreu VP, Bazzicalupo Fund for the grant RDG6130001 entitled Impact of cli- A, Thilini Chethana KW, Clericuzio M, Dayarathne mate change on fungal diversity and biogeography in the MC, Dissanayake AJ, et al (2017) Fungal diversity Greater Mekong Subregion. K.D. Hyde thanks Kunming notes 603–708: taxonomic and phylogenetic notes Institute of Botany, Chinese Academy of Sciences (project on genera and species. Fungal Divers 87, 1–235. 2013T2S0030), for the Visiting Professorship for Senior 15. Hongsanan S, Maharachchikumbura SSN, Hyde KD, International Scientists. K.D. Hyde also thanks Chiang Samarakoon MC, Jeewon R, Zhao Q, Al-Sadi AM, Mai University for the position as Visiting Professor. Bahkali AH (2017) An updated phylogeny of Sordari- Gareth Jones is supported under the Distinguished Scien- omycetes based on phylogenetic and molecular clock tist Fellowship Program (DSFP), King Saud University. MS evidence. Fungal Divers 84, 25–41. Calabon is grateful to the Mushroom Research Foundation 16. Luo ZL, Hyde KD, Liu JK, Maharachchikumbura SSN, Jeewon R, Bao DF, Bhat DJ, Lin CG, et al and Department of Science and Technology – Science (2019) Freshwater Sordariomycetes. Fungal Divers Education Institute. 99, 451–660. REFERENCES 17. Tibpromma S, Hyde KD, McKenzie EHC, Bhat DJ, Phillips AJL, Wanasinghe DN, Samarakoon MC, 1. James TY, Berbee ML (2012) No jacket required – Jayawardena RS, et al (2018) Fungal diversity notes new fungal lineage defies dress code. BioEssays 34, 840-928: micro-fungi associated with Pandanaceae. 94–102. Fungal Divers 93, 1–160. 2. Hawksworth DL, Lücking R (2017) Fungal diversity 18. Samarakoon MC, Hyde KD, Hongsanan S, McKenzie revisited: 2.2 to 3.8 million species. In: Heitman EHC, Ariyawansa HA, Promputtha I, Zeng XY, Tian J, Howlett BJ, Crous PW, Stukenbrock EH, JamesTY, Q, et al (2019) Divergence time calibrations for Gow NAR (eds) The Fungal Kingdom, Wiley Online ancient lineages of Ascomycota classification based Library, pp 79–95. on a modern review of estimations. Fungal Divers 96, 3. Hyde KD, Xu J, Rapior S, Jeewon R, Lumyong S, 285–346. Niego AGT, Abeywickrama PD, Aluthmuhandiram 19. An Z, Wang C, Liu X, Bennett JW (2010) China’s JVS, et al (2019) The amazing potential of fungi: 50 fungal genomics initiative: a whitepaper. Mycology 1, 1–8. www.scienceasia.org ScienceAsia 46S (2020) 9

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Appendix A. Supplementary data

Table S1 Mainstream Asian mycological journals.

Journal Publisher Reference

a,b Asian Journal of Mycology CEFR, Mae Fah Luang Univ, https://asianjournalofmycology.org Thailand Current Research in Environmental Beijing Academy of Agriculture & http://www.creamjournal.org and Applied Mycologya,b,c Forestry Sciences, China b,c Fungal Diversity Kunming Institute of Botany, https://www.springer.com/journal/13225 (IF: 15.596) CAS, China b Fungal Science Mycological Society of Taiwan http://mycology.sinica.edu.tw/ TaiwanMycoSoc/Default.aspx a Kavaka Mycological Society of India http://www.fungiindia.co.in/index.php a,b Medical Mycology Journal Japanese Society for Medical https://www.jstage.jst.go.jp/browse/ (IF: 2.851) Mycology mmj/-char/en a,c Mycobiology Korean Society of Mycology https://www.tandfonline.com/toc/ (IF: 1.369) tmyb20/current a,b,c Mycology Chinese Mycological Society https://www.tandfonline.com/toc/ (IF: 0.56) tmyc20/current b,c Mycoscience Mycological Society of Japan https://www.journals.elsevier.com/ (IF: 1.229) mycoscience a,b,c Mycosphere Guizhou Academy of Agricultural http://mycosphere.org (IF: 2.015 ) Sciences, China b Mycosystema Mycological Society of China and http://manu40.magtech.com.cn/Jwxb/ Institute of Microbiology, CAS CN/1672-6472/home.shtml a,b Plant Pathology and Quarantine Agriculture College, Guizhou http://www.plantpathologyquarantine.org Univ, China a,b Studies in Fungi Institute of Animal Science, http://www.studiesinfungi.org CAAS, China a,c The Korean Journal of Mycology Korean Society of Mycology http://www.kjmycology.or.kr a Dielectric data obtained at room temperature and a frequency of 1 kHz b No publication costs c Scopus indexed journals

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