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Thailand’s amazing diversity: up to 96% of fungi in northern Thailand
may be novel
Kevin D. Hyde
1,2
•Chada Norphanphoun
2
•Jie Chen
3
•Asha J. Dissanayake
2
•Mingkwan Doilom
1,4,5
•
Sinang Hongsanan
6,7
•Ruvishika S. Jayawardena
2
•Rajesh Jeewon
8
•Rekhani H. Perera
2
•
Benjarong Thongbai
10
•Dhanushka N. Wanasinghe
1,4
•Komsit Wisitrassameewong
9
•Saowaluck Tibpromma
1,2,4
•
Marc Stadler
10
Received: 8 October 2018 / Accepted: 14 November 2018
ÓSchool 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 organ-
isms. 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
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
&Mingkwan Doilom
j_hammochi@hotmail.com
Extended author information available on the last page of the article
123
Fungal Diversity
https://doi.org/10.1007/s13225-018-0415-7(0123456789().,-volV)(0123456789().,-volV)
and promote plant growth, including crops in many envi-
ronments (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 (Gostinc
ˇar
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 micro-
fungi 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 Dia-
porthe, the less well-studied hosts Pandanaceae and Tec-
tona 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 well-
studied 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.
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 (Dar-
anagama 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 sap-
robes 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; Kar-
unarathna 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 addi-
tion, the cosmopolitan species A. subrufescens,A. endox-
anthus (‘‘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 col-
lected 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
123
tasty, then numerous new species could be introduced to
international cuisine (Thawthong et al. 2014). Agaricus
subrufescens is an important medicinal species (Wisi-
trassameewong 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 Thai-
land 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 sec-
tion 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 col-
lected more than 250 specimens of Amanita, mainly in the
north (Table 3). Of all Amanita species collected in Thai-
land, 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
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
Number of
described
new species
Number
remaining
undescribed
New records
of known
species to
Thailand
Percentage
novelty
(%)
Expected
new species
in future
studies
Agaricus 17 0 75 38 32 5 93 40?
Amanita 27 3 69 17
a
40 9 83 25?
Lactarius
subg.
Russularia
0 0 23 17
b
3 3 87 30?
Colletotrichum 12 5 39 16 10 8 67 50?
Diaporthe 26 0 26 13 12 1 96 100?
Pandanaceae 87 3 93 54 15 21 74 100?
Tectona
grandis
15 0 53 24 5 24
c
55 100
Foliar
epiphytes
6 0 74 42 23 9 88 100
a
Three species recorded by Li et al. (2016);
b
One species recorded by Verbeken et al. (2014);
c
Two species recorded by Meeboon and Takamatsu
(2017)
Fungal Diversity
123
three currently accepted subgenera, Lactarius subg. Rus-
sularia (Fr.) Kauffman is a difficult group to study due to
the similarity in macromorphological and latex features
amongst species. Species in this subgenus can be recog-
nised 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 abun-
dant 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 basid-
iomata 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. Rus-
sularia 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
Table 2 Agaricus species recorded from Thailand (novel species are
in bold)
Taxon References
Agaricus albosquamosus Zhao et al. (2016)
Agaricus angusticystidiatus He et al. (2018a)
Agaricus atrodiscus Ariyawansa et al. (2015)
Agaricus badioniveus Chen et al. (2017)
Agaricus bisporiticus Thongklang et al. (2014)
Agaricus brunneolutosus Chen et al. (2017)
Agaricus brunneosquamulosus Chen et al. (2015)
Agaricus chiangmaiensis Karunarathna et al. (2014)
Agaricus endoxanthus Zhao et al. (2012a)
Agaricus erectosquamosus Zhao et al. (2016)
Agaricus exilissimus Ariyawansa et al. (2015)
Agaricus fimbrimarginatus Chen et al. (2017)
Agaricus flammicolor Chen et al. (2017)
Agaricus flavicentrus Liu et al. (2015)
Agaricus flocculosipes Zhao et al. (2012b)
Agaricus fuscopunctatus Thongklang et al. (2014)
Agaricus haematinus Ariyawansa et al. (2015)
Agaricus hanthanaensis Liu et al. (2015)
Agaricus heterocystis Zhao et al. (2011)
Agaricus inthanonensis Zhao et al. (2016)
Agaricus leucocarpus Chen et al. (2017)
Agaricus leucolepidotus Zhao et al. (2016)
Agaricus luteofibrillosus Li et al. (2016)
Agaricus luteopallidus Chen et al. (2017)
Agaricus megacystidiatus Karunarathna et al. (2014)
Agaricus megalosporus Chen et al. (2012)
Agaricus microvolvatulus Thongklang et al. (2014)
Agaricus murinocephalus Zhao et al. (2012a)
Agaricus niveogranulatus Chen et al. (2015)
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 Zhao et al. (2016)
Agaricus suthepensis 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)
Table 3 Amanita species recorded from Thailand (novel species are
in bold)
Taxon References
Amanita atrobrunnea Li et al. (2016)
Amanita ballerina Thongbai et al. (2017a)
Amanita brunneitoxicaria Thongbai et al. (2017a)
Amanita brunneoprocera Thongbai et al. (2018)
Amanita brunneosquamata Thongbai et al. (2018)
Amanita brunneoumbonata Thongbai et al. (2018)
Amanita castanea Thongbai et al. (2016)
Amanita cinnamomea Thongbai et al. (2018)
Amanita concentrica Thongbai et al. (2016)
Amanita digitosa Li et al. (2016)
Amanita esculenta Thongbai et al. (2018)
Amanita flavidocerea Thongbai et al. (2018)
Amanita flavidogrisea Thongbai et al. (2018)
Amanita fuligineoides Thongbai et al. (2017a)
Amanita gleocystidiosa Li et al. (2016)
Amanita hemibapha sensu lato Sanmee et al. (2008)
Amanita luteoparva Thongbai et al. (2018)
Amanita macrocarpa Thongbai et al. (2017a)
Amanita cf. oberwinklerana Thongbai et al. (2017a)
Amanita pyriformis Li et al. (2016)
Amanita pseudoporphyria Sanmee et al. (2008)
Amanita rimosa Thongbai et al. (2016)
Amanita rubromarginata Thongbai et al. (2016)
Amanita rubrovolvata Sanmee et al. (2008)
Amanita cf. spissacea Thongbai et al. (2017a)
Amanita strobilipes Li et al. (2016)
Amanita suborientifulva Thongbai et al. (2018)
Amanita subovalispora Thongbai et al. (2018)
Amanita zangii Thongbai et al. (2016)
Fungal Diversity
123
as L. camphoratus (Bull.: Fr.) Fr., were often applied for
Thai species. Thus, the biodiversity of Lactarius subg.
Russularia in northern Thailand was explored (Wisi-
trassameewong 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
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 consid-
erable 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,Ou1980). The intro-
duction 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 (Jayawar-
dena et al. 2016a). Plant pathogens continue to develop
resistance against chemicals and host crop defence mech-
anisms (Crouch 2014) and this has become a challenge in
developing control strategies. Most pathogens are micro-
fungi 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 phy-
topathogenic 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 identi-
fied 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
Table 4 Lactarius subg. Russularia species recorded from Thailand
(novel species are in bold)
Taxa References
Lactarius aquosus Wisitrassameewong et al. (2015)
Lactarius atrobrunneus 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 Wisitrassameewong et al. (2015)
Lactarius rubrocorrugatus Wisitrassameewong et al. (2015)
Lactarius sublaccarioides Wisitrassameewong et al. (2014a)
Lactarius subzonarius Hongo (1957)
Lactarius tangerinus Wisitrassameewong et al. (2015)
Fungal Diversity
123
may even not be present (Phoulivong et al. 2010). Phou-
livong et al. (2010) analyzed DNA sequence data of 25
isolates from eight tropical fruits, which were morpho-
logically 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 dif-
ferentiate 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 Col-
letotrichum 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 Col-
letotrichum 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 eco-
nomical 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 man-
grovei, 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
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 con-
ducted by Tongsri et al. (2016). Thus, until the incorpo-
ration 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 spe-
cies 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 sym-
bionts, endophytes or epiphytes. Tropical hosts are gener-
ally 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 monocotyle-
donous 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 stud-
ied by us since 2014. More than 150 specimens were col-
lected, 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
123
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
Colletotrichum acutatum Capsicum annuum
Fragaria sp.
Than et al. (2008b)
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
Mangifera indica
Prihastuti et al. (2009)
Colletotrichum boninense Dendrobium sp. Ma et al. (2018)
a
Manihot esculenta Sangpueak et al. (2018)
a
Colletotrichum brevisporum Neoregelia sp., Pandanus pygmaeus 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
Carica papaya
Dimocarpus longan
Than et al. (2008a,b), Diao et al. (2017)
a
Cymbopogon citratus,
Pennisetum purpureum
Manamgoda et al. (2013)
a
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
Colletotrichum gloeosporioides 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
Colletotrichum graminicola 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 Phoulivong et al. (2010)
Capsicum annuum, Hymenocallis sp. 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
Fungal Diversity
123
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 com-
menced 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
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, par-
ticularly 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 plant-
stunting 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 mar-
ketability problems (Chomnunti et al. 2014). The ecology
and taxonomy of fungal epiphytes has been studied
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
Fungal Diversity
123
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 burnsii
a
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 Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b)
Cladosporium endophyticum
a
Pandanus sp. Krabi Province Tibpromma et al. (2018a)
Clonostachys krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Colletotrichum fructicola
a
Pandanus sp., Freycinetia sp. Chumphon, Ranong Province Tibpromma et al. (2018a)
Colletotrichum pandanicola Pandanus sp. Phang Nga Province Tibpromma et al. (2018b)
Colletotrichum pandanicola
a
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 pandanicola
a
Pandanus sp. Chumphon Province Tibpromma et al. (2018a)
Diaporthe siamensis
a
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 Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Dictyosporium pandanicola Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Distoseptispora thailandica Pandanus sp. Prachuap Khiri Khan Province Tibpromma et al. (2018b)
Endomelanconiopsis freycinetiae
a
Freycinetia sp. Ranong Province Tibpromma et al. (2018a)
Endopandanicola thailandica
a
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 theobromae
a
Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a)
Lasionectria krabiense Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Malaysiasca phaii Freycinetia javanica Krabi Province Tibpromma et al. (2018b)
Massarina pandanicola
a
Pandanus sp. Chumphon Province Tibpromma et al. (2018a)
Meyerozyma caribbica
a
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 endophytic
a
Freycinetia sp. Ranong Province Tibpromma et al. (2018a)
Fungal Diversity
123
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 epi-
phytes often grow intermixed with colonies of other spe-
cies (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
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), Muyoco-
pronales (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.
Table 7 (continued)
Taxa Host Location References
Neomassarina pandanicola Pandanus sp. Prachuap Khiri Khan Province Hyde et al. (2018)
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 Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Pestalotiopsis jiangxiensis
a
Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a)
Pestalotiopsis krabiensis Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Pestalotiopsis microspora
a
Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a)
Phanerochaete chrysosporium
a
Pandanus sp. Chumphon, Ranong Province Tibpromma et al. (2018a)
Phyllosticta capitalensis
a
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 Pandanus sp. Krabi Province Tibpromma et al. (2018b)
Torula ficus Pandanus sp. Chiang Mai Province 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
Fungal Diversity
123
Table 8 Species of fungi on
Tectona grandis mostly
reported in northern Thailand
(novel species from Thailand
are in bold)
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
Fungal Diversity
123
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 well-
documented 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.
Fungi on Rosaceae
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 iden-
tification. 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; Rovna
´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 Cor-
naceae. 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.
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.
Novelty of fungi in Europe
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 Lu
¨cking (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 rep-
resent new fungal species. The cultivated mushroom
Lepista sordida (Thongbai et al. 2017b) yielded nudic acid
B, and the nematode trapping species Hohenbuehelia gri-
sea, produced the novel heterocyclic terpenoid pleuroth-
iazol (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 sub-
type alpha, which offers avenues to the development of the
mushroom as a new nutraceutical with chloesterol-lower-
ing 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
123
Table 9 Foliar epiphytes
reported in Thailand (novel
species from Thailand are in
bold)
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
Fungal Diversity
123
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)
Amandinea punctata
a
Rosa sp. Wanasinghe et al. (2018)
Angustimassarina quercicola
a
Rosa canina Wanasinghe et al. (2018)
Angustimassarina rosarum 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 Rosa canina Wanasinghe et al. (2018)
Comoclathris rosigena Rosa canina 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 eres
a
Rosa sp. Wanasinghe et al. (2018)
Diaporthe foeniculina
a
Rosa canina Wanasinghe et al. (2018)
Diaporthe rhusicola
a
Rosa canina Wanasinghe et al. (2018)
Diaporthe rosae Rosa sp. Wanasinghe et al. (2018)
Diaporthe rosicola Rosa canina Wanasinghe et al. (2018)
Diaporthe rudis
a
Rosa canina Wanasinghe et al. (2018)
Diplodia seriata
a
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 Rosa canina Wanasinghe et al. (2018)
Lasiodiplodia theobromae
a
Rosa canina Wanasinghe et al. (2018)
Lecidella elaeochroma
a
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)
Muriformistrickeria rosae Rosa canina Wanasinghe et al. (2018)
Muriformistrickeria rubi
a
Rosa canina Wanasinghe et al. (2018)
Murilentithecium rosae Rosa canina Wanasinghe et al. (2018)
Neoascochyta rosicola Rosa canina Wanasinghe et al. (2018)
Neoconiothyrium rosae Rosa canina Wanasinghe et al. (2018)
Neofusicoccum australe
a
Rosa sp. Wanasinghe et al. (2018)
Neopaucispora rosaecae Rosa ecae Wanasinghe et al. (2018)
Neosetophoma rosarum Rosa canina Wanasinghe et al. (2018)
Neosetophoma rosigena Rosa canina Wanasinghe et al. (2018)
Paraconiothyrium rosae Rosa canina Wanasinghe et al. (2018)
Paraphaeosphaeria michotii
a
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)
Fungal Diversity
123
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 mycolo-
gists generally agreeing on 1.5–5 million (Hawksworth
1991; Hawksworth and Lu
¨cking 2017). In the most recent
estimates, Hawksworth and Lu
¨cking (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 demon-
strated 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 unrecog-
nized 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 accumu-
lated to date, and with randomized sampling only in
northern Thailand, the fungal diversity with new species
recovered far exceeds our expectations. We should revisit
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 pleurospora
a
Rosa sp. Wanasinghe et al. (2018)
Poaceicola rosae Rosa canina Wanasinghe et al. (2018)
Populocrescentia rosae Rosa hissarica Wanasinghe et al. (2018)
Pseudocercospora rosae Rosa canina 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 caulium
a
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 rubriostiolata
a
Rosa multibracteata Wanasinghe et al. (2018)
Uzbekistanica rosae-hissaricae Rosa hissarica Wanasinghe et al. (2018)
Uzbekistanica yakutkhanika Rosa hissarica Wanasinghe et al. (2018)
Wojnowicia rosicola Rosa canina Wanasinghe et al. (2018)
Xenomassariosphaeria rosae Rosa canina Wanasinghe et al. (2018)
Expected number of new species: [100
a
New host records
Fungal Diversity
123
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 Pan-
danaceae, but certainly there are unculturable orphan fun-
gal species, that could represent a reservoir of novel
species with a panoply of unexploited bioactive com-
pounds. How will this impact on our anticipated number of
fungi? Possibly we can argue that previously simple blind
sampling strategies and inadequate DNA sequence analy-
ses, limit new species discovery. Our studies in northern
Thailand have revealed more new species as the taxonomic
assessment methods used became more reliable and sub-
strates/environments sampled were strategic. There are
obviously other major fungal groups that warrant investi-
gation (e.g. aquatic fungi, entomopathogens, dung fungi,
edible mushrooms, mycorrizhae, and wood decay fungi)
that are globally distributed, but poorly sampled in Thai-
land. 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 descrip-
tion. In these relatively well known mushroom genera we
N O
OH
O
N
HO
O
O
OH
H
HO
NH
OH
O
H
H
OOH
O
O
OO
H
H
Lenormandin B
Pyristriatin B
Lentinulactam
Nudic Acid B
OO
O
O
N
S
H
H
Pleurothiazol
O
O
Cl
Gymnopalyne B
O
O
HO OH
O
OH
Deconin C
O
O
HO
O
Blazeispirol A
Fig. 1 Chemical structures of selected biologically active secondary metabolites from Thai fungi
Fungal Diversity
123
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 Cam-
bodia, 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 appre-
ciation 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 postdoc-
toral researchers funded by Yunnan Human Resources and Social
Security Department, and The 64th Grant from China Postdoctoral
Science Foundation.
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(2018a) Simplified and efficient DNA extraction protocol for
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Translucidithyrium thailandicum gen. et sp. nov.: a new genus in
Phaeothecoidiellaceae. Mycol Prog 17:1087–1096
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Asian species of Amanita (Agaricales, Basidiomycota): taxo-
nomic and biogeographic implications. Fungal Divers
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Doilom M, Mapook A, Liu JK (2017a) Novomicrothelia
pandanicola sp. nov., a non-lichenized, Trypetheliaceae species
from Pandanus. Phytotaxa 321:254–264
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in Northern Thailand. PhD Thesis, KMITL, Bangkok, Thailand
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Affiliations
Kevin D. Hyde
1,2
•Chada Norphanphoun
2
•Jie Chen
3
•Asha J. Dissanayake
2
•Mingkwan Doilom
1,4,5
•
Sinang Hongsanan
6,7
•Ruvishika S. Jayawardena
2
•Rajesh Jeewon
8
•Rekhani H. Perera
2
•Benjarong Thongbai
10
•
Dhanushka N. Wanasinghe
1,4
•Komsit Wisitrassameewong
9
•Saowaluck Tibpromma
1,2,4
•Marc Stadler
10
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
2
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
Fungal Diversity
123
3
Instituto de Ecologı
´a, 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
5
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Rai 50200, Thailand
6
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Sciences and Oceanography, Shenzhen
University, Shenzhen, People’s Republic of China
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
9
National Science Technology and Innovation Policy Office,
Bangkok 10330, Thailand
10
Department of Microbial Drugs and German Centre for
Infection Research (DZIF), partner site Hannover-
Braunschweig, Helmholtz Centre for Infection Research,
Inhoffenstrasse 7, 38124 Brunswick, Germany
Fungal Diversity
123
