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
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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.
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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
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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
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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
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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
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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.
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Chukeatirote E, Alias SA, McKenzie EH, Hyde KD (2013a)
Phyllosticta capitalensis, a widespread endophyte of plants.
Fungal Divers 60:91–105
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Wikee S, Lombard L, Nakashima C, Motohashi K, Chukeatirote E,
Cheewangkoon R, McKenzie EHC, Hyde KD, Crous PW
(2013b) A phylogenetic re–evaluation of Phyllosticta
(Botryosphaeriales). Stud Mycol 76:1–29
Wisitrassameewong K, Karunarathna SC, Thongklang N, Zhao R,
Callac P, Moukha S, Ferandon C, Chukeatirote E, Hyde KD
(2012a) Agaricus subrufescens: a review. Saudi J Biol Sci
19:131–146
Wisitrassameewong K, Karunarathna SC, Thongklang N, Zhao RL,
Callac P, Chukeatirote E, Bahkali AH, Hyde KD (2012b)
Agaricus subrufescens: new records to Thailand. Chiang Mai J
Sci 39:281–291
Wisitrassameewong K, Nuytinck J, Hyde KD, Verbeken A (2014a)
Lactarius subgenus Russularia (Russulaceae) in Southeast Asia:
1. Species with very distant gills. Phytotaxa 158:023–042
Wisitrassameewong K, Nuytinck J, Hampe F, Hyde KD, Verbeken A
(2014b) Lactarius subgenus Russularia (Russulaceae) in Southeast Asia: 2. Species with remarkably small basidiocarps.
Phytotaxa 188:181–197
Wisitrassameewong K, Nuytinck J, Le Thanh H, De Crop E, Hampe
F, Hyde KD, Verbeken A (2015) Lactarius subgenus Russularia
(Russulaceae) in Southeast Asia: 3. New diversity in Thailand
and Vietnam. Phytotaxa 207:215–241
Wolfe BE, Tulloss RE, Pringle A (2012) The irreversible loss of a
decomposition pathway marks the single origin of an ectomycorrhizal symbiosis. PLoS ONE 7:e39597
Wu HX, Schoch CL, Boonmee S, Bahkali AH, Chomnunti P, Hyde
KD (2011) A reappraisal of Microthyriaceae. Fungal Divers
51:189–248
Wurzbacher C, Nilsson RH, Rautio M, Peura S (2017) Poorly known
microbial taxa dominate the microbiome of permafrost thaw
ponds. ISME J 11:1938–1941
Xiang QY, Thomas DT, Zhang W, Manchester SR, Murrell Z (2006)
Species level phylogeny of the genus Cornus (Cornaceae) based
on molecular and morphological evidence—implications for
taxonomy and Tertiary intercontinental migration. Taxon
55:9–30
Xiao YP, Wen TC, Hongsanan S, Sun JZ, Hyde KD (2017)
Introducing Ophiocordyceps thanathonensis, a new species of
entomogenous fungi on ants, and a reference specimen for O.
pseudolloydii. Phytotaxa 328:115
Xiao YP, Wen TC, Hongsanan S, Jeewon R, Luangsa-ard JJ, Brooks
S, Nadeeshan D, Long FY, Hyde KD (2018) Multigene
phylogenetics of Polycephalomyces (Ophiocordycipitaceae,
Hypocreales), with two new species from Thailand. Sci Rep
(accepted)
Yan YC, Zhang ZX, Song YJ, Deng DF, Liu ZY (2015) First report of
Prunus serrulata stem canker caused by Botryosphaeria dothidea in China. Plant Dis 100:858
Yang ZL (1997) Die Amanita-Arten von Südwestchina. Bibl Mycol
170:1–240
Yang YL, Liu ZY, Cai L, Hyde KD, Yu ZN, McKenzie EHC (2009)
Colletotrichum anthracnose of Amaryllidaceae. Fungal Divers
39:123–146
Zadoks JC (1985) On the conceptual basis of crop loss assessment:
the threshold theory. Annu Rev Phytopathol 23:455–473
Zeng XY, Wen TC, Chomnunti P, Liu JK, Boonmee S, Hyde KD
(2016) Ceramothyrium longivolcaniforme sp. nov., a new
species of Chaetothyriaceae from northern Thailand. Phytotaxa
267:51–60
Zeng XY, Jeewon R, Wen TC, Hongsanan S, Boonmee S, Hyde KD
(2018a) Simplified and efficient DNA extraction protocol for
Meliolaceae specimens. Mycol Prog 17:403–415
Zeng XY, Hongsanan S, Hyde KD, Putarak C, Wen TC (2018b)
Translucidithyrium thailandicum gen. et sp. nov.: a new genus in
Phaeothecoidiellaceae. Mycol Prog 17:1087–1096
Zhang LF, Yang JB, Yang ZL (2004) Molecular phylogeny of eastern
Asian species of Amanita (Agaricales, Basidiomycota): taxonomic and biogeographic implications. Fungal Divers
17:219–238
Zhang SN, Hyde KD, Jones EG, Cheewangkoon R, Boonmee S,
Doilom M, Mapook A, Liu JK (2017a) Novomicrothelia
pandanicola sp. nov., a non-lichenized, Trypetheliaceae species
from Pandanus. Phytotaxa 321:254–264
Zhang MZ, Li GJ, Dai RC, Xi YL, Wei SL, Zhao RL (2017b) The
edible wide mushrooms of Agaricus section Bivelares from
Western China. Mycosphere 8:1640–1652
Zhao RL (2008) Systematics of Agaricus, Cyathus and Micropsalliota
in Northern Thailand. PhD Thesis, KMITL, Bangkok, Thailand
Zhao R, Karunarathna S, Raspé O, Parra LA, Guinberteau J, Moinard
M, De Kesel A, Barroso G, Courtecuisse R, Hyde KD, Guelly
AK (2011) Major clades in tropical Agaricus. 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
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