Fungal Diversity
https://doi.org/10.1007/s13225-022-00498-w
REVIEW
Predicting global numbers of teleomorphic ascomycetes
Indunil C. Senanayake1,2 · Dhandevi Pem3,4 · Achala R. Rathnayaka3,4 · Subodini N. Wijesinghe3,4
Saowaluck Tibpromma5,7 · Dhanushka N. Wanasinghe5,6,7 · Rungtiwa Phookamsak5,6,7 ·
Nuwan D. Kularathnage2,3,4 · Deecksha Gomdola3,4 · Dulanjalee Harishchandra3,4,8 ·
Lakmali S. Dissanayake9 · Mei‑mei Xiang2 · Anusha H. Ekanayaka3,4,5 · Eric H. C. McKenzie10 ·
Kevin D. Hyde2,3,4,5 · Hao‑xing Zhang1 · Ning Xie1
·
Received: 25 September 2021 / Accepted: 20 January 2022
© The Author(s) 2022
Abstract
Sexual reproduction is the basic way to form high genetic diversity and it is beneficial in evolution and speciation of fungi.
The global diversity of teleomorphic species in Ascomycota has not been estimated. This paper estimates the species number
for sexual ascomycetes based on five different estimation approaches, viz. by numbers of described fungi, by fungus:substrate
ratio, by ecological distribution, by meta-DNA barcoding or culture-independent studies and by previous estimates of species
in Ascomycota. The assumptions were made with the currently most accepted, “2.2–3.8 million” species estimate and results
of previous studies concluding that 90% of the described ascomycetes reproduce sexually. The Catalogue of Life, Species
Fungorum and published research were used for data procurement. The average value of teleomorphic species in Ascomycota
from all methods is 1.86 million, ranging from 1.37 to 2.56 million. However, only around 83,000 teleomorphic species have
been described in Ascomycota and deposited in data repositories. The ratio between described teleomorphic ascomycetes
to predicted teleomorphic ascomycetes is 1:22. Therefore, where are the undiscovered teleomorphic ascomycetes? The
undescribed species are no doubt to be found in biodiversity hot spots, poorly-studied areas and species complexes. Other
poorly studied niches include extremophiles, lichenicolous fungi, human pathogens, marine fungi, and fungicolous fungi.
Undescribed species are present in unexamined collections in specimen repositories or incompletely described earlier species. Nomenclatural issues, such as the use of separate names for teleomorph and anamorphs, synonyms, conspecific names,
illegitimate and invalid names also affect the number of described species. Interspecies introgression results in new species,
while species numbers are reduced by extinctions.
Keywords Ascomycota · Estimates · Habitat diversity · Molecular techniques · Species concepts
Introduction
Ascomycota Caval-Sm. is the largest fungal phylum comprising around 93,000 extant species and are generally
known as “sac fungi” (Bennett and Turgeon 2017; Clark
et al. 2018; Catalog of Life 2021). Members of Ascomycota are ubiquitously spread in various terrestrial and fresh
or marine ecosystems (Naranjo-Ortiz and Gabaldón 2019).
Most ascomycetes are saprobes while some are soil or dung
inhabitants (Richardson 2019). Some are animal, human and
Handling Editor: Antonio Roberto Gomes de Farias
* Ning Xie
Shainin@msn.cn
plant pathogens or parasites such as epiphytes or fungicolous fungi (Wu et al. 2011), while others are symbionts as
endophytes, lichenicolous and mycorrhizae (Lawrey and
Diederich 2003; Chomnunti et al. 2014; Kim et al. 2017;
Sun et al. 2019; Hyde et al. 2020b). The sexual reproduction in ascomycetes often occurs as a response to adverse
environmental conditions (Nieuwenhuis and James 2016)
and it results high genetic diversity between species (Lee
et al. 2010). Sexual reproduction helps to purge deleterious mutations and also selects beneficial mutations to adapt
to a fluctuating environment (Otto and Lenormand 2002).
Ascospores are more resistant to environmental stress and
more widely dispersed than the conidia (Kirschner 2019).
Sexual reproduction in ascomycetes comprising the matingtype loci (MAT) which encodes key transcription factor genes
Extended author information available on the last page of the article
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that govern speciation (Paoletti et al. 2005). Two compatible
partners fuse their genetic materials by recombination or crossing-over and also meiosis and mitosis to produce genetically
diversified offspring (O'Gorman et al. 2009). Sexual reproduction occurs in the same mycelium (homothallic/self-fertile) or
two different mycelia (heterothallic) and they produce spores
in a sac-like structure called an ascus (Fig. 1). The sexual
structures such as asci and ascospores are contained in fruiting
bodies. Released ascospores from fruiting bodies germinate
when contact with a suitable substrate and produce the mating
Fig. 1 Sexual reproduction of filamentous ascomycetes. a Reproduction cycle. b Ascus formation within fertilized ascogonium (drawn from
Peraza-Reyes and Berteaux-Lecellier 2013)
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Fungal Diversity
type male (+) and female (−) mycelia (Taylor et al. 2006). The
male mycelium produces an antheridium and the female produces an ascogonium, which are different reproductive organs.
Sexual reproduction in Ascomycota includes three stages
as plasmogamy, karyogamy and meiosis (Wallen and Michae
2018). During the first step (plasmogamy), two haploid cells are
fused and lead to a dikaryotic stage where two haploid nuclei
coexist in a single cell. During the second step (karyogamy), the
haploid nuclei of + and − mycelia fuse to form a diploid zygote
nucleus. The diploid zygote nucleus produces unique haploid
nuclei by meiosis and these haploid nuclei form haploid spores
with mitosis and cell division within the sac-like structures called
ascus. These ascospores are then released and germinate to form
new hyphae in new environments. The production of these highly
specialized sexual tissues is initiated and controlled by gene,
protein, and secondary metabolite networks and those proteins
regulate the expression of primary MAT genes (MAT1-1-1 and
MAT1-2-1) (Dyer 2007). Both MAT genes are typically essential
for successful fertilization and ascomatal development (Ferreira
et al. 1998). Additionally, the MAT1-1-1 gene is also critical
for ascospore production in some species (Debuchy et al. 2010)
and alteration of both MAT genes results in failure to form even
immature sexual structures (Lee et al. 2003).
Some anamorphic ascomycetes shuffle their genetic material
by parasexuality. Parasexual reproduction results in recombination of genes from different individuals but does not involve
meiosis and formation of a zygote by fertilization as in sexual
reproduction. Parasexuality generates both genotypic and phenotypic diversity in species (Hirakawa et al. 2017). The variations in genotypes create by shuffling of different chromosome
homologs, recombination between homologs and the generation
of cells in various ploidy states with one or more supernumerary chromosomes (Forche et al. 2008). The mutations in genetic
materials and haploidization occur inside the heterokaryotic
hyphae prior to conidial formation (Becker and de Castro-Prado
2006). However, the mixing-up genetic materials without forming sexual structures does not discuss here and this study only
estimates species formed by true sexual reproduction.
Teleomorphic ascomycetes are morphologically diversified (Fig. 2) and ubiquitous taxa that can survive in various
ecological habitats in both terrestrial and aquatic ecosystems
(Gould 2008; Schoch et al. 2009). Teleomorphic species are
reported from only 17 classes in Ascomycota including all
classes in subphylum Pezizomycotina and Neolectomycetes
in subphylum Taphrinomycotina.
Why should we estimate the global species
richness of teleomorphic ascomycetes?
In mycology, “species” is simply defined as a diagnosable
cluster of individuals within a parental pattern of lineage
displays a pattern of phylogenetic ancestry and descent
among units and hence, it is valuable to give it a species
name (Brown 2002; Aldhebiani 2018; Maharachchikumbura et al. 2021). Finalizing the global fungal inventory
is a challenge due to their morphological, ecological and
physiological diversity (Purvis and Hector 2000). Estimates of the total number of teleomorphic ascomycetes
have major inferences for systematics, resource management and classification (Hawksworth 1991) as they play
key roles in ecosystems as decomposers, mutualists and
pathogens individually and with the interactions of each
other (Schmit and Mueller 2007) and some of them in
the plant rhizosphere protect root systems from pathogens (Mendes et al. 2013). Mutualistic ascomycetes are
associated with their host without causing harm. It is a
beneficial relationship for both fungi and the host (Volk
2013). Ascomycetes associate with algae or cyanobacteria to form lichens (Weber and Büdel 2011) and Arthoniomycetes, Dothideomycetes, Eurotiomycetes, Lecanoromycetes, Lichinomycetes and Sordariomycetes comprise
the lichenized species (Grube and Winka 2002; Andersen
and Ekman 2005). More than 40% of lichenized fungi are
species in Ascomycota (Brodo et al. 2001; Schoch et al.
2009). Mycorrhizae are symbiotically associated with
plant roots while endophytes are associated with living
plants (Volk 2013).
There are numerous of plant pathogenic teleomorphic
ascomycetes causing various diseases of economic crops
and forest trees (Lu et al. 2003). The increasing number
of virulent fungal infectious diseases is regarded as a
worldwide threat to food security (Hyde et al. 2018). An
unprecedented number of diseases caused by fungi including teleomorphic ascomycetes have resulted in some of the
most severe diebacks in economic crops and wild species
(Fisher et al. 2012; Hyde et al. 2019). Many species can
be harmless endophytes in some plants, however cause
severe damages in others (Hardoim et al. 2015; Terhonen
et al. 2019; Song et al. 2021). Therefore, description and
cataloging of teleomorphic ascomycetes helps to identify
fungal pathogens and prevent future disasters.
The nutritional sources of the teleomorphic ascomycetes vary from dead organic matter to synthesized
compounds by other organisms and they decompose litter, maintain the nutrient cycles and improve soil quality
(Gams 2007; Gould 2008; Frąc et al. 2018; Senanayake
et al. 2020a). However, the mycota involved in decomposition incompletely known and there may be many species
interactions (Frey-Klett et al. 2011; Volk 2013). Hyaloscyphaceae, Melanommataceae, Mytilinidiaceae and Savoryellaceae are some ascomycetous families which have
many saprobic teleomorphic species (Hernández-Restrepo
et al. 2017). Identifying and describing the teleomorphic
ascomycetes involved in litter degradation is important in
organic farming and fertilizer production (Peyvast et al.
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Fungal Diversity
Fig. 2 Diversity of fruitting bodies in teleomorphic Ascomycota. a, j, k, p, t, w, x, z Leotiomycetes, d Neolectomycetes, b, h, m, y Dothideomycetes, n, r Eurotiomycetes, e Geoglossomycetes, o Orbiliomycetes, q, v Pezizomycetes, c, f, g, i, l, s, u Sordariomycetes
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Fungal Diversity
2008). Isolation of those fungi and application to soil as
a microbial assortment to enhance plant growth and yield
is required (Khalid et al. 2017). Additionally, antibiotics,
anticancer, anti-inflammatory and some medically important chemicals are extracted from teleomorphic ascomycete cultures (Rajamanikyam et al. 2018; Al-Fakih and
Almaqtri 2019; Wu et al. 2019b). Many organic acids and
enzymes such as citric acid, gluconic acid, amylases and
proteases are produced by teleomorphic ascomycetes.
Morels, truffles, Hypomyces lactifluorum are edible ascomycetes (Acton and Sandler 2008; Splivallo et al. 2010).
Therefore, revealing the undescribed teleomorphic ascomycetes, estimating the species number and exploring
their chemical and biological properties are important
(Yang et al. 2018).
How many teleomorphic species
in Ascomycota based on different estimation
methods?
Estimated number of teleomorphic ascomycetes
based on numbers of described fungi
Traditionally, taxonomic studies of teleomorphic ascomycetes were based mostly on morphological characters,
subcellular arrangement, bio-chemical, physiological and
ecological studies (Yang 2011; Maharachchikumbura et al.
2021). During 1960–2000, phenotypic taxonomic studies
were improved by microscopy and in vitro culturing (Klopfstein 2016). Many groups of teleomorphic ascomycetes have
been intensively studied (Todd et al. 2014) and simultaneously significant taxonomic monographs were published
(Barr 1978, 1987; Kohlmeyer and Kohlmeyer 1979; Schmit
and Lodge 2005; Senanayake et al. 2017, 2018). In the previous 20 years, molecular methods have modernized and
studies are based on biogeography, phylogeny, population
genetics, systematics and taxonomy (Yang 2011).
Since 1982, there has been a periodic update in the classification of taxa in Ascomycota especially in the Journal
Systema Ascomycetum. The Species Fungorum database
(http://www.speciesfungorum.org) has recorded the number of new species described each year. The taxonomy of the
phylum Ascomycota has been updated at a fast pace over the
last few years (Hyde et al. 2013, 2020a, b, c; Jaklitsch et al.
2016; Maharachchikumbura et al. 2016; Liu et al. 2017;
Ekanayaka et al. 2018). Accepted families with descriptions and list of genera in the Ascomycota were provided by
Jaklitsch et al. (2016). Currently there are 22 classes in the
phylum Ascomycota as Archaeorhizomycetes, Arthoniomycetes, Candelariomycetes, Coniocybomycetes, Dothideomycetes, Eurotiomycetes, Geoglossomycetes, Laboulbeniomycetes, Lecanoromycetes, Leotiomycetes, Lichinomycetes,
Neolectomycetes, Orbiliomycetes, Pezizomycetes, Pneumocystidomycetes, Saccharomycetes, Sareomycetes, Schizosaccharomycetes, Sordariomycetes, Taphrinomycetes, Xylobotryomycetes and Xylonomycetes (Lumbsch and Huhndorf
2010; Voglmayr et al. 2019; Beimforde et al. 2020).
The “2.2–3.8 million” species estimate (Hawksworth and
Lücking 2017) is considered as the most rational estimate by
many mycologysts (Hyde et al. 2020b). There are approximately 150,000 extant fungal species (Roskov et al. 2019;
Species Fungorum 2021), however this is only 15–26% of
the estimated species (Hyde et al. 2020b). Early mycologists
believed that only half of ascomycetes are meiosporic fungi
which obligatory sexually reproduced and do not produce
asexual spores (Reynolds and Taylor 1993). However, the
rest are probably obligatory mitosporic or facultative mitosporic fungi with undetected teleomorphs (Nieuwenhuis and
James 2016). Reynolds and Taylor (1993) showed that about
5% of obligatory anamorphic ascomycetes are known to be
pleomorphic and thus discretionary sexually reproduce.
However, the most accepted value is that may be the 90% of
the described ascomycetes sexually reproduce (Judson and
Normark 1996; Normark et al. 2003).
There are 92,725 described species in Ascomycota
(Catalog of Life 2021; Species Fungorum 2021). Wijayawardene et al. (2017) listed 8897 species in Ascomycota
with undetermined teleomorphs. There are 523 anamorphic ascomycetes with undetermined teleomorph have been
introduced from 2018 to 2020 (31 December 2020) (Species
Fungorum 2021). Hence, 9420 ascomycetous species are
obligatory anamorphic species and therefore around 83,305
teleomorphic ascomycetes have been described. The reproduction arrangements appear to be similar across the phylum
Ascomycota even though the life cycles among the major
groups are different (Nieuwenhuis and James 2016). Hence,
there should be 1.25–2.17 million teleomorphic ascomycetes based on described number of species considering the
2.2–3.8 million species estimate.
Estimated number of teleomorphic ascomycetes
based on fungus:substrate or host ratio
The ratio of fungal species to each plant species was one
of the key elements in estimating global species richness
as 1.5 million (Hawksworth 1991) and this estimate was
assumed from independent data sets which shown that the
number of fungi in all environments was six times higher
than the vascular plants present, inferred on a global scale.
The “1.5 million estimate” was considered too low because,
the number of plant species and the fungus: plant ratios were
too conservative and many were collected from other substrata such as insects (Hawksworth 2001). The number of
plant species has increased from 250,000 to 390,000 (Pimm
and Joppa 2015) signifying that the estimated number of
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Fungal Diversity
fungal species must rise to 2.4–3 million. Some authors also
suggested that the ratio of fungi: plants is about 10:1, those
found in soil, insects or lichen were excluded (O’Brien et al.
2005; Blackwell 2011). However, there are several complications in the fungus: plant ratio concept. The total inventory
of species described in a particular area increases gradually, while the number of plant species remains more or less
unchanged (Hawksworth and Lücking 2017). Therefore, the
fungus: plant ratio decreases gradually with the description
of new species.
Some studies based on meta-DNA sequencing of decaying litter samples showed that the fungus: plant ratio is 13:1
(Hawksworth and Lücking 2017). Therefore, the actual ratio
in a particular area may be significantly higher than the ratio
indicated by traditional inventory techniques. Further, the
whole planet has not been screened evenly and known species number in some places is higher than in others. For
example, fungal diversity in North America, Europe and
Japan is well-studied compared to South Asia and Africa
(Hawksworth 2001; Větrovský et al. 2019). O’Brien et al.
(2005) noted that the fungus: plant ratios of two forests in
North Carolina gave 19:1 and 13:1 suggesting that there
may be 3.5 to 5.1 million species. Further, this ratio changes
according to the substrata and Taylor et al. (2010) showed
that fungus: plant ratio is 7.5:1 in forest soils in Alaska.
Tedersoo et al. (2014) analyzed soil samples using meta-barcoding molecular methods and concluded that the number
of species had been overestimated by 1.5 to 2.5 times from
data based on plant: fungus species ratios.
Therefore, the estimates based on studies of fungus: plant/
insect/lichen/plant OTUs in soil ratios in a site, obtained
by field survey and molecular approaches, have generated
lower ranges from 0.42 to 3.5 million to (O’Brien et al.
2005; Tedersoo et al. 2014) an upper range from 0.6 to
5.1 million (O’Brien et al. 2005; Piepenbring et al. 2012)
(Table 1). Considering the average of the upper and lower
range of previous estimates, we estimated species number
is 1.96–2.85 million based on fungus: host ratio. There are
Table 1 Species estimates based
on fungus: host ratio
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around 150,000 described species and 92,725 are ascomycetes, which is around 63.4%. Therefore, there should be
about 1.11–1.62 million estimated teleomorphic ascomycetes (Table 1), excluding 10% obligatory anamorphic species (Normark et al. 2003).
The patterns of introducing new species are biased with
more described from economically important plants (Cannon and Hawksworth 1995). Most early described species
were collected from temperate floral communities and host
specificity in tropical plants are not well-reported and new
host records are not published (Tedersoo et al. 2010; Piepenbring et al. 2011). Therefore, fungus: plant/insect/substrate
ratio is not an ideal method to estimate species numbers
because of uneven exploration of global species in habitats.
Estimated number of teleomorphic ascomycetes
based on ecological distribution
The traditional quantification approaches of teleomorphic
ascomycetes are established on the opportunistic collections
of specimens based on host, substrate, area and transects
(Schmit and Lodge 2005). Opportunistic collecting requires
highly trained collectors who can recognize taxa in the
field without a bias. Some collectors only perceive favored
particular groups of teleomorphic ascomycetes. Further,
conspicuous species and more common species are often
overlooked (Lodge et al. 2004). Teleomorphic ascomycetes
produce fruit bodies in different types of substrata (Lodge
1996; Huhndorf and Lodge 1997; Schmit and Lodge 2005;
Sainz et al. 2018). Some ascomycetes sexually reproduce
rather dependably while others do so only occasionally, and
therefore require long periods to be recorded from a particular area (Straatsma et al. 2001). Further, fruiting patterns,
abundance and dispersion of ascomycetes differs among
substrata (Lodge et al. 2004).
Some teleomorphic ascomycetes show a wide range of
host and substrate variation and also different modes of life.
Daldinia eschscholtzii, one of the common endophytes in
Ratio
Estimated species
in millions
Host/substrate
References
N/A
8:1
13–19:1
7.5:1
6:1
1.8:1
17:1
1.8:1
50–53:1
9.8:1
~ 1.5
~ 1.5
3.5–5.1
1.9–2.8
0.8–5.1
0.45–0.6
1.5–6
0.42–2.72
0.9–0.95
2.2–3.8
Fungi:insect
Fungi:plants
Fungi:plants
Fungi:plant OTUs in soil
Fungi:plants
Fungi:plants
Fungi:plant OTUs in soil
Fungi:plant OTUs in soil
Fungi:lichens
Fungi:plants
Hywel-Jones (1993)
Hawksworth (2001)
O’Brien et al. (2005)
Taylor et al. (2010)
Blackwell (2011)
Piepenbring et al. (2012)
Taylor et al. 2014)
Tedersoo et al. (2014)
Zhang et al. (2015), Wang et al. (2016)
Hawksworth and Lücking (2017)
Fungal Diversity
plants (Stadler et al. 2014; Helaly et al. 2018) and marine
algae (Tarman et al. 2012), has been reported as an endosymbiont of a mantis gut (Zhang et al. 2011), and a human
pathogen (Chan et al. 2015). Further, Diaporthe sojae, a
known pathogen of soybean, was also isolated from infected
skin of an immunocompromised patient after kidney transplantation (Garcia-Reyne et al. 2011). Diaporthe toxica is
a plant endophyte and occasionally a plant pathogen (Williamson et al. 1991) and produces secondary metabolites that
result in toxicoses of animals such as liver disease known as
lupinosis of sheep (Gardiner 1975; Allen and Wood 1979;
Williamson et al. 1994). Therefore, it is necessary to understand the ecology and life strategies of teleomorphic ascomycetes before estimating the species number. Further, many
endophytes do not sporulate in culture (Sun and Guo 2012)
and some ascomycetes change colony morphology while
growing and sub-culturing on different media (Senanayake
et al. 2017). Some ascomycetes do not sexually reproduce
or need specific conditions for sexual reproduction (Sun and
Heitman 2011). Direct morphological examination of fruiting structures on substrata or media only is therefore biased
in estimating number of teleomorphic ascomycetes (Guo
et al. 2001; Promputtha et al. 2004).
Case studies from marine ascomycetes
Marine ascomycetes are recovered repeatedly from marine
habitats, able to grow and/or sporulate on substrata in
marine environments, form symbiotic relationships with
other marine organisms, adapt and evolve at the genetic
level or be metabolically active in marine environments
(Pang et al. 2016). They are observed in a range of marine
substrates, including mangrove plant wood and leaves, driftwood, saltmarsh plants, algae, dead coral, and sand grains
on beaches (Gonçalves et al. 2021; Walker and Robicheau
2021), along with severe marine ecosystems such as deepsea trenches, hydrothermal vents, deep-sea subsurfaces, cold
methane seeps and hypersaline, anoxic, and suboxic waters
(Raghukumar and Ravindram 2012; Xu et al. 2018). Marine
ascomycetes colonize a variety of substrata based on their
ability to degrade complex substrata such as lignocellulose,
keratin, chitin and calcareous structures and ascomycetes are
the major decomposers in marine ecosystems (Kohlmeyer
and Volkmann-Kohlmeyer 2001; Walker and Campbell
2010). Marine ascomycetes are also known as symbionts
and pathogens of marine algae and marine fauna (Hyde et al.
1998).
The accessibility and the nature of substrate for colonization, competition, pH, temperature, and saltiness of water
affect the diversity of marine ascomycetes (Jones 2000,
2011). Most marine fungi are recognized to have a cosmopolitan distribution (Pugh and Jones 1986). However, basic
biogeographic diversity data are lacking for marine ascomycetes in most parts of the world (Walker and Robicheau
2021). Some marine fungi such as Aniptodera chesapeakensis Shearer & M.A. Mill., Ceriosporopsis Halima Linder,
Corollospora maritima Werderm., Lignincola laevis Höhnk,
Savoryella lignicola E.B.G. Jones & R.A. Eaton, and Torpedospora radiata Meyers have diverse geographic dispersion
which is classified as tropical to subtropical while Lulwoana
uniseptata (Nakagiri) Kohlm. et al. is reported from temperate habitats only (Torta et al. 2015; Tibell et al. 2020).
Mora et al. (2011) presented an approach to estimate species numbers on earth and ocean and predicted that 0.005
million species are marine. However, 91% of species in the
ocean await description and increasing the sampling intensity is required to characterize the underexplored species
of marine biodiversity (Walker and Robicheau 2021). It
is estimated that more than 10,000 marine fungal species
exist globally (Jones 2011; Walker et al. 2017) and only
around 1000 have been described (Jones et al. 2015; Pang
et al. 2016). Jones et al. (2019) listed 1257 marine species
belonging to 539 genera and 943 of them are ascomycetes
(Jones et al. 2009, 2015; Abdel-Wahab et al. 2010; Pang
et al. 2010; Abdel-Wahab and Nagahama 2011; Dayarathne
et al. 2016, 2019).
The number of species is estimated as 2.2–3.8 million
(Hawksworth and Lücking 2017) while only around 150,000
species have been described (Species Fungorum 2021). If
there are 943 described marine ascomycetous species, then it
is predicted that 13,831–23,889 marine ascomycetes should
be in oceans. However, considering only 90% of described
ascomycetes are sexually reproduced (Judson and Normark
1996; Normark et al. 2003), then there are 12,448–21,500
marine, teleomorphic ascomycetes.
Case studies from freshwater ascomycetes
Freshwater ascomycetes are an ecological assortment rather
than a taxonomic group and they reproduce sexually or asexually residing on sunken or partially submerged woody substrata in freshwater environments (Tsui et al. 2016; Calabon
et al. 2021). In spite of their importance as decomposers and
food sources in freshwater food webs, there has been little
research on their global distribution, community structure
and species diversity (Shearer et al. 2015). Freshwater ascomycetes occur on submerged or partially submerged substrata in lotic and lentic aquatic habitats. The teleomorphic
ascomycetes are more dominant on submerged wood, while
the anamorphic ascomycetes occur on submerged leaf litter
(El-Elimat et al. 2021).
Phylogenetically, freshwater ascomycetes are grouped
mostly throughout the class Dothideomycetes, Leotiomycetes and Sordariomycetes in Ascomycota (Shearer et al.
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Fungal Diversity
2009, 2014). They have soft fruiting bodies during teleomorphic stage with appendage baring ascospores (Hyde
et al. 1998). Asci developed in ascomata possess pathways
for efficient spore discharge and ascospores are frequently
appendaged or have sheaths. The appendages facilitate spore
dissemination and bonding to the substrata (Hyde and Goh
2003).
Freshwater ascomycetes are observed across the both lentic and lotic ecosystems, and they are commonly associated
as endophytes and parasites on algae and aquatic macrophytes along with the saprobes on the dead plant matter (Lu
et al. 2018). Many freshwater ascomycetes are believed to
have evolved from terrestrial ancestors through a wide range
of evolutionary pathways (Vijaykrishna et al. 2006; Grossart
et al. 2019).
The number of aquatic species has been estimated as
0.5–10 million based on molecular data (Bass and Richards 2011; Blackwell 2011; Mora et al. 2011). A significant
number of freshwater species are teleomorphic ascomycetes
(Shearer et al. 2007; Hu et al. 2013; Shearer and Raja 2021).
About 738 species of freshwater ascomycetes are known
from their teleomorph, belonging to approximately 170 genera (El-Elimat et al. 2021; Shearer and Raja 2021). There are
around 83,305 described teleomorphic ascomycetes (Species
Fungorum 2021) and if there are 738 described freshwater teleomorphic ascomycetes, it is estimated that there are
19,490–33,664 aquatic teleomorphic ascomycetes based on
“2.2–3.8 figure”.
suspected that these yeasts also might provide nutritional
supplements.
Insect associated fungi were estimated to be 1.5 million (Hywel-Jones 1993) and Stork (2018) updated this to
5.5 million, while 1–2% of them may be cryptic species.
Therefore, including the cryptic species, the consensus
estimate of insect associated species ranges from 5.505 to
5.511 million. However, this is more deviated from currently
estimated species numbers. The diversity of insect associated ascomycetes has been extensively studied (Aung et al.
2008; Mora et al. 2011; Hyde et al. 2018) and they are taxonomically distributed in Clavicipitaceae, Cordycipitaceae,
and Ophiocordycipitaceae in Hypocreales, ambrosia fungi
(e.g., Ceratocystis, Ophiostoma) in the Ophiostomatales, all
families in Laboulbeniomycetes and some species in Saccharomycetes (Sung et al. 2007; Vega et al. 2012; Araújo
and Hughes 2016; Maharachchikumbura et al. 2016; Wijayawardene et al. 2018). Mueller and Schmit (2007) estimated
around 50,000 insect associated species, when there are 750
described species. However, currently, there are more than
4000 insect-associated species described (Species Fungorum 2021). Hence, there should be 52,800–91,200 insect
associated teleomorphic ascomycetes according to “2.2–3.8
species estimate” and considering generally 90% of ascomycetes reproduce sexually (Judson and Normark 1996;
Normark et al. 2003).
Case studies from coprophilous ascomycetes
Case studies from insect‑associated
ascomycetes
Insects are an extremely diversified group of organisms in
all ecosystems (Stork 1988) and include dragonflies, mayflies, grasshoppers, cockroaches, termites, stoneflies, true
bugs, flies, beetles, butterflies, moths, ants, bees, and wasps
(Stork et al. 2015, 2018). Insects and fungi share a long
history of relationship in the similar habitats and (Bourtzis
and Miller 2003) those interactions can be mutualistic or
harmful. Insects involved in associations with fungi include
members of the Coleoptera, Diptera, Homoptera, Hymenoptera, and Isoptera.
Fungal biotrophic parasites of insects are rare, except
for the very successful associations of Laboulbeniomycetes
(Blackwell et al. 2020; Haelewaters et al. 2021). Vega and
Dowd (2005) highlighted the role of yeast-insect endosymbionts in supporting the digestion and detoxification of plant
materials ingested by insects and discovered an enormous
number of species of Saccharomycetes. Some fungi also
interact with insects by providing nutritional supplements
(Vega and Blackwell 2005). Suh et al. (2001) described
around 200 new yeast species from the gut of beetles. It is
13
Coprophilous fungi grow, sporulate and germinate on herbivore dung (Tretter et al. 2014; Lazarus et al. 2017) and
they are specialized to survive in the harsh environment of
the gastrointestinal tract of animals (Richardson 2001b; Bell
2005; Kirschner et al. 2015; De Souza et al. 2017; Lavrinienko et al. 2021). Coprophilous fungi recycle the nutrients
in animal dung and release nutrients to the soil (Basumatary
and McDonald 2017; Florenzano 2019).
Species richness and composition of coprophilous ascomycetes differ with abiotic and biotic factors. Intra- and
inter-specific interactions in a dung pile affect fungal succession and species composition (Maynard et al. 2018; Lavrinienko et al. 2021). Many coprophilous ascomycetes are most
common on only one or a few dung types (Lundqvist 1972)
and dung from animals that live together generally show a
similar species composition (Richardson 2001a).
However, the fungal community varied more between
animal dung types than between the various grassland habitats (Angel and Wicklow 1983). Coprophilous ascomycetes
can be found more frequently on dung of herbivores than
carnivores (Lundqvist 1972; Richardson 2001a). In addition, they have seldom been reported on reptile or amphibian
dung, indicating that coprophily in fungi developed among
Fungal Diversity
the warm-blooded animals (Webster 1970). Some ascomycetes are strictly coprophilous and they have a distinct lifecycle restricted to dung pile, plant surface and animal gut
(Wicklow 1992). However, some spores disperse in soil. The
spores of coprophilous ascomycetes are highly pigmented,
with thick walls and are protected against the harmful ultraviolet sunlight (Ingold and Hudson 1993). Therefore, spores
remain in soil alive and those species have been reported as
soil fungi.
Kruys (2005) reported that many coprophilous ascomycetes belong to order Pleosporales in Dothideomycetes and
three of the families are solely or mostly coprophilous, viz.
Delitschiaceae, Phaeotrichaceae and Sporormiaceae. Calaça
et al. (2015) listed 143 coprophilous ascomycetous species
recorded from Brazil. Melo et al. (2020) studied the diversity
and species richness of coprophilous fungi in Brazil. A total
of 271 species are reported from dung substrata and among
them, 70% of recorded species are ascomycetes. Most species are included in Sordariales, Hypocreales and Microascales and 9% of recorded species are anamorphic ascomycetes (Saumell et al. 1999, 2000; Saumell and Padilha 2000).
Calaça et al. (2020) listed the coprophilous species
recorded in Brazil during 1900–2013 and 117 from 210
coprophilous species are ascomycetes. They were collected
from 12 states of Brazil and total area of these 12 states
is 303146 km2. Therefore, one coprophilous ascomycetous
species was collected from each 2591 km2. If this value
inferred for land area of earth, there should be 196,874
coprophilous ascomycetes. If 9% of recorded species are
anamorphic species (Saumell et al. 1999, 2000; Saumell and
Padilha 2000), then there should be 177,187 teleomorphic
coprophilous ascomycetes. However, this value is 443,061
according to Melo et al. (2020). Therefore, it is assumed
that there should be around 177,000–443,000 teleomorphic
coprophilous ascomycetes.
Case studies from soil ascomycetes
Fungi occur in the soil or soil-associated environments at
least for some stage in their life-cycle known as soil fungi
(Bridge and Spooner 2001). They are active, freely growing fungi closely associated with other organisms or inactive dormant propagules (Rämä and Quandt 2021). The
role of soil fungi are an extremely complex and are fundamental to the soil ecosystem (Hawksworth et al. 1995).
Soil fungi carry out many different functions in soils such
as the degradation of dead organic matter, binding soil particles to improve the aeration, water penetration, destroy
soil pathogens, and improve soil health by formation of
propagules (Zin and Badaluddin 2020; Jayaraman et al.
2021).
Soil fungi can only be consistently identified if they produce fruiting bodies (Hibbett et al. 2016) and conventional
techniques are unable to reliably identify the species that are
assumed to be present in any given soil sample due to the
fastidious nature of the great majority of species (Wardle and
Lindahl 2014). Fungal communities in soil can be extremely
species rich and patchy at small spatial scales (Taylor and
Sinsabaugh 2015). High throughput sequencing of soil fungi
in boreal forest sites revealed around 300 taxa in 0.25 g soil
and the dominant taxa in the sites were quite distinct from
each other (Taylor and Sinsabaugh 2015).
Around 80% of all soil-inhabiting taxa cannot be identified to species and 20% cannot be reliably assigned to known
orders (Hawksworth 2001; Vartoukian et al. 2010; Tedersoo
et al. 2014, 2017). The number of soil fungal species is considerably greater than the described amount and studies with
the integration of molecular, genetic and ecological factors
may reveal more species. The number of species identified
by traditional culture dependent methods doubles when
the same soil samples are analysed by culture-independent
methods (Lord et al. 2002; Arenz et al. 2006; Malosso et al.
2006; Smith and Jaffee 2009; Zachow et al. 2009; Hirsch
et al. 2013; Rodolfi et al. 2016).
There are no significant estimates of the number of species made for soil fungi. Gilman (1957) included around 700
fungal species which grew on only non-selective media by
soil dilution method and many species were later included
(Barron 1968; Domsch et al. 1993). Watanabe (1994) suggested that at least 1200 species have been isolated from soil.
Pugh (1969) showed that only 17% of soil fungi can be readily grown in culture media. Therefore, Hawksworth (1991)
estimated that around 7000 species could be considered as
soil fungi based on Watanabe (1994). However, there are
more than 80,000 fungal species so far named and described,
and they are likely to occur in the soil environment at some
stage in their life-cycle (Bridge and Spooner 2001). These
species are mainly distributed in the subphylum Taphrinomycotina as the fission yeasts, animal and plant pathogens,
the root-associated, sporocarp-forming, filamentous fungi
(Schoch et al. 2009), while the Saccharomycotina includes
the budding yeasts. Pezizomycotina contains lichen-forming
fungi, mycorrhizal fungi, dark-septate endophytes, pathogens and saprotrophs. Further, Bridge and Spooner (2001)
proposed that at least 10% of the described fungal species
are obligatory soil fungi and around 75% of them are ascomycetes (Taylor and Sinsabaugh 2015). If this applies to
2.2–3.8 estimate considering 90% of them are teleomorphs
(Normark et al. 2003), there should be 148,500–256,500
teleomorphic ascomycetes in soil.
13
Fungal Diversity
Case studies from lichenized ascomycetes
Case study from ligninolytic ascomycetes
Lichens are stable self-supporting associations of a mycobiont and a photobiont (Maria et al. 2021). They produce many
secondary metabolites such as phenolic compounds, dibenzofurans, depsides, depsidones, depsones, lactones, quinones and pulvinic acid derivatives which are accumulated
externally on the hyphae rather within the cells (Tehler and
Irestedt 2007). These compounds are unique to each species
and can be used as food, fodder, dyes, and pharmaceuticals.
The lichens are the best bio-indicators of air pollution (Garty
2001). The mycobiont is usually an ascomycete but in a few
cases it is a basidiomycete. The photosynthetic partners are
generally green algae or cyanobacteria (Richardson 2002).
The relationship between fungi and lichens can be endolichenic and lichenicolous (Tripathi and Joshi 2019).
Most of the endolichenic fungi and other accessory fungi
reported from inside the lichen thalli are phylogenetically
distinct from lichenicolous fungi (Miadlikowska et al. 2004)
and more closely related to endophytic ascomycetes in vascular plants (Miadlikowska and Lutzoni 2004). Generally,
most of the lichenized fungi belong to Ascomycota or rarely
to Basidiomycota. Further, the ascolichens mainly belong
to Sordariomycetes, Lecanoromycetes and Eurotiomycetes.
The Lecanoromycetes is almost an entirely lichenized class
comprising the remarkable population of lichen-forming
species (Nash 2008).
A study of the diversity and distribution of the fungal
communities that were associated with seven lichens in the
Ny-Ålesund Region (Svalbard, High Arctic) using Roche
454 pyrosequencing method reported 370 OTUs of which
294 belonged to Ascomycota (Zhang et al. 2015). Among
these, Leotiomycetes, Dothideomycetes, and Eurotiomycetes
were the major classes, with Helotiales, Capnodiales, and
Chaetothyriales as the dominant orders. Further, Wang et al.
(2016) studied fungal diversity associated with a common
lichen Hypogymnia hypotrypa in China and 28 were ascomycetes from 50 species. It is assumed roughly that lichen:
fungi ratio as 1:45 (Fernández-Mendoza et al. 2011; Muggia
and Grube 2018).
There are around 20,000 described lichen species
(Çobanoğlu et al. 2010) and about 98% have an ascomycetous mycobiont (François et al. 2001). Therefore, there
should be 19,600 endolichenic ascomycetous species. Further, about 40% of species in the Ascomycota are lichenized
or lichenicolous fungi (Kirk et al. 2008). Currently, there
are around 93,000 described species in Ascomycota and
40% is 37,200. However, excluding 10% obligatory anamorphic species, there are around 17,640–33,480 teleomorphic
lichenized or lichenicolous species in Ascomycota.
Fungi play a vital role in plant litter decomposition in ecosystems (Boddy et al. 2008; Watkinson et al. 2015; Baldrian
2017). They can degrade different types of organic compounds in the litter (Baldrian and Lindahl 2011), which
other organisms are unable to degrade (de Boer et al. 2005).
Ligninolytic fungi produce several kinds of extracellular
enzymes that help to degrade cellulose and other organic
compounds in litter and helps nutrient turnover (Sinsabaugh et al. 2002; Romaní et al. 2006). Most common
extracellular enzymes produced by ligninolytic fungi are
α-glucosidase, β-glucosidase, cellobiosidase, xylosidase,
polyphenol oxidase, N-acetyl-polyphenol oxidase, N-acetylβ-glucosaminidase and acid phosphatase (Marx et al. 2001;
De-Forest 2009).
The majority of ligninolytic fungi are ascomycetes (Seena
et al. 2019), that colonize during the early stages of decomposition (Aneja et al. 2006; Voříšková and Baldrian 2013;
Prakash et al. 2015). It was proposed that ascomycetes dominate during the initial stages of litter decay presumably due
to a superior ability to degrade cellulose (Weber et al. 2011)
and decreases during the process of degradation as they are
gradually replaced by other non-ascomycetous saprobes
(Frankland 1998; Osono 2007). The classes Dothideomycetes, Eurotiomycetes, Saccharomycetes and Taphrinomycetes are the most ligninolytic species abundant classes in
Ascomycota (Zhang et al. 2018). A few studies have denoted
that endophytes living in plants shift their lifestyle to saprotrophs when the substrates die and they play a key role in
early stage decomposition (Purahong and Hyde 2011; Fesel
and Zuccaro 2015; Purahong et al. 2016; Szink et al. 2016).
However, NGS data may partly reflect the fungal succession from ascomycetes to other fungi in early to later stages
of litter decomposition and it does not clearly provide an
idea about the species richness and abundance (Amend et al.
2010; Peršoh 2015).
Many studies have been revealed the species composition in decaying litter. However, the number of ligninolytic
fungal species has not been estimated. Haňáčková et al.
(2015) analyzed fungal species involved in decomposition
of pine needle litter through culture dependent and culture
independent methods. This study proved that the ratio of
species recognition of culture dependent method to culture
independent method is 1:2. Purahong et al. (2016) sampled
leaf litter 473 times to study decomposing fungi and showed
that the percentage of detection frequency of ascomycetes
was 66–82%. Zhang et al. (2018) studied the ligninolytic
fungal diversity in China and revealed 2621 fungal OTUs
which mainly belong to Ascomycota, Basidiomycota and
13
Fungal Diversity
Zygomycota. Further, 75% are ascomycetes. Meanwhile,
Seena et al. (2019) studied the ligninolytic fungal diversity
in 19 globally distributed streams and the total number of
fungal OTUs revealed in this study was 1311 with 79.7%
being ascomycetes. Voříšková and Baldrian (2013) did a
similar study and revealed that 71% of species are ascomycetes. Osono (2019) obtained 127 fungal species from 1133
leaf litter isolates and 95 are ascomycetes.
Based on the above studies, it is assumed that around
70–80% of ligninolytic fungi are ascomycetes and therefore, around 1045–1966 ligninolytic ascomycetes have
been recorded in culture dependent and culture independent studies (Zhang et al. 2018; Seena et al. 2019). Further,
Dashtban et al. (2010) reported that more than 14,000 fungal
species produce ligninolytic enzymes and all litter degrading fungi must produce ligninolytic enzymes (Kumar and
Chandra 2020). Assuming that 70–80% of above ligninolytic enzyme producing fungi are ligninolytic ascomycetes,
there are around 10,500 described ligninolytic ascomycetes.
If this applies to 2.2–3.8 estimate, considering 90% of them
are teleomorphs (Normark et al. 2003), it is assumed that
there are around 138,600–239,400 teleomorphic species of
ligninolytic ascomycetes.
Case studies from endophytic ascomycetes
Endophytes are mutualists that colonize asymptomatically
inside of any tissues of living plants at least in any phase in
their life cycle (Singh and Dubey 2015). Bills (1996) proposed that some type of mycorrhizae such as ericoid mycorrhizae and pseudomycorrhizae can be endophytes. Endophytic colonization generally does not cause any damage to
its host and does not produce any structures emerging from
the external plant (Azevedo and Araújo 2007). Some endophytes can grow invitro in culture media.
Endophytes are ubiquitous and occur within a broad
range of host plants, such as mosses, ferns, grasses, shrubs,
deciduous and coniferous trees and lichens (Guo et al. 2008;
Albrectsen et al. 2010; Mohamed et al. 2010; Su et al. 2010;
Sun et al. 2011). Endophytes are an important component
in natural ecosystems and they produce various bioactive
chemicals, promote host growth, improve resistance to environmental stress and decompose litter (Aly et al. 2010; Saikkonen et al. 2010; Xu et al. 2010; Purahong and Hyde 2011;
Tejesvi et al. 2011; Gouda et al. 2016).
Endophytic fungi have not been seriously considered in
the estimation of fungal numbers (Hawksworth 1991). However, there could be more than 1 million endophytic fungal
species based on ratios of vascular plants to fungal species
of 1:4 (Petrini 1991). Dreyfuss and Chapela (1994) proposed
that there should be 1.3 million endophytic fungal species.
Most culturable plant endophytes are ascomycetes belonging
to orders Amphisphaeriales, Capnodiales, Diaporthales,
Hypocreales, Pleosporales, Sordariales, Trichosphaeriales
and Xylariales (Guo et al. 2001; Crozier et al. 2006; He et al.
2012; Koukol et al. 2012).
Hamzah et al. (2018) revealed that the ratio of endophytic
Ascomycota: Basidiomycota is around 25:1 and there are
around 30,000 described, endophytic basidiomycetes species
(Anke 1989; Anke and Steglich 1988). Hence, there should
be 750,000 endophytic ascomycetes. Further, any vascular
plant species can host somewhere 4–5 different endophytic
fungal species (Sun and Guo 2012). There are 372,383 species of vascular plants and therefore, there could be 1.49
million endophytic fungal species. Hence, it is estimated
that there are 675,000–1,341,000 endophytic teleomorphic
ascomycetes excluding 10% obligatory anamorphs (Normark
et al. 2003).
Case studies from epiphytic ascomycetes
Epiphytic fungi reside either permanently or casually on the
surface of plants (Langvad 1980). They can multiply and
grow on the surface of healthy leaves without any adverse
effect to the host, while casual epiphytes land on the healthy
leaf surface in the form of spores or mycelia but cannot grow
like residents (Kharwar et al. 2010. The coexistence of epiphytic and endophytic microorganisms may play an important role for plant health and plant protection (Andrews and
Harris 2000) as well as contributing to microbial biodiversity (Hawksworth and Rossman 1997).
Epiphytic fungi are dominant in Ascomycota and Basidiomycota with very few in other phyla and Sordariomycetes,
Dothideomycetes and Eurotiomycetes are the most frequent
among the all classes in Ascomycota (Dong et al. 2021).
Among epiphytic fungi in the phyllosphere, 70–98% is ascomycetes while the rhizosphere comprises 73% of epiphytic
species in Ascomycota (Oliveira et al. 2017). A comparative study of endophytic and epiphytic fungal association
in leaves of Eucalyptus citriodora Hook. revealed 279 epiphytes out of 478 fungal isolates. This means number of
epiphytic fungi is 1.4 times higher than endophytes. Further, Dong et al. (2021) analyzed the epiphytic and endophytic fungal communities of tomato plants and revealed
161 epiphytic fungal OTUs and 119 endophytic fungal
OTUs. This suggested that the number of epiphytic fungi
is around 1.4 higher than the endophytic fungi (Kharwar
et al. 2010). In this study, we concluded that there should
be 335,000–675,000 teleomorphic, endophytic ascomycetes and therefore, we suggest that there should be
469,000–945,000 teleomorphic epiphytic ascomycetes.
Based on the above case studies, it is estimated that there
are around 1,710,000–3,405,000 teleomorphic ascomycetes
in different ecological habitats and the average is around
13
Fungal Diversity
2,558,000 species. However, all the predictions are based
on the available data and some ecological groups are wellstudied while others are poorly examined.
Estimated number of teleomorphic ascomycetes
based on meta‑DNA and culture‑independent
studies
The identification of some teleomorphic ascomycetes such
as fungal symbionts, endophytes, marine species associated
with plants and green algae, and parasites is challenging
due to their unculturable nature (Blackwell 2011). However,
advanced molecular techniques facilitate the discovery of
undescribed species from unculturable samples (Zhang
et al. 2010; Blackwell 2011). The fungal diversity estimate
increases with the advent of more uncultured fungi and
fungi from environmental samples. Environmental DNA
(meta-DNA) can be genetic material acquired directly from
environmental samples, such as soil, sediment, water and
others devoid of any clear signs of biological material is
an effective, safe and quick standardized sampling method
(Prosser and Hedgpeth 2018). The development of advanced
molecular techniques such as high-throughput sequencing
has greatly contributed in identification of undescribed species (Barnes and Turner 2015). More fungal species were
identified by culture-independent approaches than by culture-dependent methods (Zhang et al. 2010) and the fungal species detected by one method is really different from
other method, even for the dominant fungal species (Wu
et al. 2019a).
Environmental DNA is a powerful tool to explore the
hidden teleomorphic ascomycetes and it challenges understanding of global biodiversity (Venter et al. 2004). It was
estimated that the number of fungal species on earth ranged
between 3.5 and 5.1 million when considering the species
recorded from environmental samples (Blackwell 2011). The
class Archaeorhizomycetes in the sub-phylum Taphrinomycotina was introduced based on only environmental DNA,
even its precise ecological niches and life cycle is unknown
(Rosling 2011). Further, an unknown, basal clade of phylum
Ascomycota which is characterized by unicellular zoospores
with a single, non-chitin or non-cellulose-walled microtubular flagellum was described as Cryptomycota based on
meta-DNA sequences (Jones et al. 2011).
Even though there is an argument as to use environmental
DNA for nomenclature, Hawksworth and Rossman (1997)
proposed to use this technique to explore the fungi existing
in un-examined niches as well as known habitats. Therefore,
the fungal species number could be much higher than the
current reliable estimates of 2.2–3.8 million.
In a study based on fungal DNA assemblages and their
spatial structure in river water using environmental DNA
metabarcoding targeting of ITS locus revealed 985 fungal
13
OTUs with 97% sequence similarity (Matsuoka et al. 2019).
Totally, 770 OTUs were assigned as Ascomycota and it
is 78.2% of total fungal OTUs. However, when there are
150,000 described fungal species, only 92,725 are ascomycetes and it is 61.8% in total. Therefore, environmental DNA
metabarcoding method provides 16.4% additional amount of
species and hence number of species in Ascomycota should
be 117,325. If this applies to 2.2–3.8 estimate, considering
90% of them are teleomorphs (Normark et al. 2003), it is
assumed that there are 1,548,690–2,675,010 species.
A study based on high-throughput sequencing of fungus: plant ratios revealed that the number of fungal species may be around 3.5–5.1 million species (O’Brien et al.
2005). Around 62% of described fungi are ascomycetes
(Species Fungorum 2021) and if this applies to 3.5–5.1
estimate (O’Brien et al. 2005), considering 90% of them
are teleomorphs (Normark et al. 2003), there should be
1,947,225–2,836,620 species. Here, we estimate around
1,747,958–2,755,815 teleomorphic species based on
O’Brien et al. (2005), Hawksworth and Lücking (2017) and
Matsuoka et al. (2019). Wu et al. (2019a) suggested that the
range of species numbers based on environmental DNA is
8.8 times higher than the traditional culture dependent methods and this gives 11–19 million species for our estimate
1.25–2.17 million species based on described species in the
data bases. This is quite large value and it is significantly
different from other estimates in this study.
Estimated number of teleomorphic ascomycetes
based on previous estimates of Ascomycota
De Meeûs and Renaud (2002) studied the phylogenetic
relationship between the parasites and the eukaryotes.
This study estimated that there should be 60,000 species in
Ascomycota. Further, Aptroot (2001) studied fungal diversity of Elaeocarpus sp. and estimated that there should be
40,000–70,000 species of ascomycetes. However, these estimates were done two decades ago and more species have
been introduced in last two decades. About 1900 fungal
species were described per year over the past two decades
(Hawksworth and Lücking 2017) and there should be around
38,000 more described species. Therefore, the updated estimate in Aptroot (2001) is 0.078–0.108 million. If 90% of
described ascomycetes are teleomorphic species (Normark
et al. 2003), it ranges for teleomorphic ascomycetes from
0.070 to 0.097 million with 0.084 million average.
Mueller and Schmit (2007) studied several groups of
ascomycetes and estimated the species number. This study
was based on Rossman (1994), Hawksworth et al. (1995)
and data of the Dictionary of Fungi. They estimated species number for several groups in Ascomycota including
Endomycetales, Helotiales, Hypocreales, insect-associated
fungi, macrolichens, non-dematiaceous hyphomycetes and
Fungal Diversity
coelomycetes, other perithecioid ascomycetes, Pezizales and
Xylariales. There were 40,706 described species in above
groups and it was predicted as 694,000 species (Mueller and
Schmit 2007) with the ratio of described species: estimated
species as 1:17. If this ratio applys for the average of updated
estimate in Aptroot (2001) (0.084 million), there should be
1,190,000–1,649,000 teleomorphic ascomycetous species.
Updated estimation of teleomorphic species
number in Ascomycota
We evaluated species number for teleomorphic ascomycetes
based on five approaches; number of described species in
databases, fungus:substrate ratio, ecological distribution
(Table 2), meta-DNA and culture-independent methods, and
previous estimates by other authors (Table 3). The average
of each method was used to propose the updated value for
species number of teleomorphic ascomycetes and it is 1.86
million. The species number of teleomorphic ascomycetes
ranges 1.37–2.56 million and the ratio between described
teleomorphic ascomycetes to predicted teleomorphic ascomycetes is 1:22. Further, there should be 3.3 million fungi
when 150,000 species has been described.
The estimates based on ecological distributions and
environmental DNA analysis provide large numbers. It is
suggested that one species can occur in several different
Table 2 Estimated species number for teleomorphic ascomycetes
based on different ecological studies
Ecological group
Range of species number
Average
of species
number
Marine species
Freshwater species
Insect associated species
Coprophilous species
Soil inhabiting species
Lichenized species
Ligninolytic species
Endophytic species
Epiphytic species
Total fungal species
12,448–21,500
19,490–33,664
52,800–91,200
177,000–443,000
148,500–256,500
17,640–33,480
138,600–239,400
675,000–1,341,000
469,000–945,000
1,710,478–3,404,744
16,974
26,577
72,000
310,000
202,500
25,560
189,000
1,008,000
707,000
2,557,611
Table 3 Updated estimates
for number of teleomorphic
species in Ascomycota based on
different approaches
habitats or in different life modes and counted several times
as the different species. Additionally, environmental DNA
reveal the hidden diversity in habitats and most unculturable species or species rarely produce teleomorph can be
trace. Therefore, the species number in these two methods
are higher than others (Fig. 3).
Limitations in estimation methods:
Why estimating the species number
of teleomorphic ascomycetes is challenging
The number of existing teleomorphic species in Ascomycota is not well-predicted and this estimation depends on
the known number of species. So, what are the difficulties
in knowing or describing teleomorphic ascomycetes? The
diversity of teleomorphic ascomycetes is much higher due
to their easily adaptable capability to different ecological
conditions and therefore, in any given community or ecosystem, teleomorphic ascomycetes can be abundantly discovered (Berbee and Taylor 1992). A high level of reproductive
plasticity and different life cycles with exciting teleomorphic
and anamorphic reproduction mechanisms can be observed
in ascomycetes due to their diversity (Wilson et al. 2019).
These variations and diversity have led to a high level of
species richness across different ecological niches.
There are three distinct phases in species introduction:
an ascending phase in 1750s to 1860s, a steep phase in the
1870s to 1880s and a relatively constant phase from the
1890s to the present day based on publications (Hawksworth
and Lücking 2017). However, the number of described species may be greater than this in ascending and steep phases.
The internet was not previously available and most mycologists worked independently. Conferences, scientific meetings
and societies were limited, and funding to attend was often
unavailable. Most mycologists were unable to directly share
their knowledge and experiences with others (Agerer et al.
2000). Further, scientific research was not done for commercial purposes. Most described species could not be published
or published papers were destroyed during world wars. Additionally, studies in mycology and description of new fungal species decreased during the Second World War period
(Hawksworth and Lücking 2017). Further, many books,
notes, experimental observations and fungal specimens were
Method
Range
Average
Based on numbers of described fungi
Based on fungus:substrate ratio
Based on ecological distribution
Based on meta-DNA/culture-independent studies
Based on previous estimates of species in Ascomycota
1.25–2.17
1.11–1.62
1.71–3.40
1.74–2.80
1.19–1.65
1.71
1.37
2.56
2.25
1.42
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Fungal Diversity
Fig. 3 Estimated numbers of teleomorphic species in Ascomycota based on five different approaches. The range of each approach
and their mean values are marked in black dots and blue diamonds,
respectively. The red dashed-line shows the average value for estimated species number of the five approaches
destroyed and most mycologists had to move to other places
or retired.
Recent intensive studies based on comprehensive inventories of ascomycetous genera and families have neglected
morphology and are mostly based on molecular data (Senanayake et al. 2018). Over 90% of the collected specimens
may constitute undescribed species (Hyde et al. 2018). The
young mycologists and students are willing to describe new
collections as novel taxa rather than assign them to existing species (Hawksworth and Lücking 2017). Further, it is
required to combine the data on biogeographic distributions,
levels of endemism and host specificity into the described
species list when estimating the number of teleomorphic
ascomycetes (Mueller and Schmit 2007). However, the number of described species has increased due to application of
molecular techniques for species delimitation. There are up
and downs in the number of described species after 2010.
This may reflect the resolve of cryptic species, synonymize
and link teleomorphs and anamorphs, rather than introducing new collections as new species.
The grid map-based method for predicting species richness introduced in Lücking et al. (2014) has been used to
predict species richness in the lichenized family Graphidaceae in Ascomycota (Aptroot and Cáceres 2016; Cáceres
et al. 2017; Mendonça et al. 2020). This method uses
known occurrence records to provide a prediction and a
precise estimate of species richness. Thus, to predict the
species richness of a large group of teleomorphic species in
Ascomycota, the accuracy of the used records is important.
Errors during species introduction, incorrect nomenclature,
misidentification of cryptic species or species complexes,
fungi from understudied fungal habitats and hosts can be
problematic. The correct estimates of teleomorphic species
richness in Ascomycota can be a difficult task (Hyde et al.
2020b). There are several limitations of the species estimates
as below.
13
One name for one fungus (1N1F)
for pleomorphic species
The “One name = One fungus” system used nowadays has
been effective in establishing standards for naming fungi
in the scientific community. Before the 1N1F system came
into effect, teleomorphs and anamorphs were given separate names depending on the circumstances from which they
were discovered. Some ascomycetes produce an anamorph
in their life cycle (Seifert and Samuels 2000) and the anamorph sometimes becomes the prominent, commonly available
morph in nature (Li et al. 2020).
With the use of DNA sequence data for species identification, the accuracy in identification and linking teleomorphs
and anamorphs of a species are important. Limitations in
available DNA sequence data could however lead to erroneous identification. Recent studies have been resolved errors
made when selecting one name for pleomorphic fungal
genera (Hawksworth 2012, 2015; Réblová et al. 2016; May
2017; Taylor et al. 2016). There are plenty of teleomorphic
species epithets in the species catalogs without linking to
anamorphic species due to a lack of molecular data from
ex-type and other authentic cultures or poor morphological descriptions (Seifert and Samuels 2000; Hawksworth
Fungal Diversity
et al. 2013). Most of the early introduced species do not have
molecular data and species introduction was based on morphology (Vellinga et al. 2015; Koukol and Delgado 2021).
Further, cultural studies not performed often and morphology of anamorph or teleomorph derived from pure cultures
was not recorded. Therefore, recollection of earlier species
or use fungarium materials to obtain DNA is required (Seifert and Rossman 2010; Aime et al. 2021).
Even after the implementation of 1N1F species system,
certain research areas still predominantly use the older
names in studies where the main focus is not related to
taxonomy and nomenclature. The teleomorph of fungi are
rarely encountered in plant pathology, thus plant pathologists tend to name the pathogenic species related to their
anamorph, but the link between the two states was rarely
established (Wingfield et al. 2012). These types of basic
errors in species naming would hinder the possibility of
accurate species estimates. Article F.8 for pleomorphic
fungi in Shenzhen code states names proposed simultaneously for separate morphs (anamorph and teleomorph) of a
taxon of non-lichen-forming Ascomycota and Basidiomycota
are necessarily heterotypic and are not therefore alternative
names (Turland et al. 2018). This code facilitates both teleomorphic and anamorphic names in the legitimate state and
those legitimate names are treated equally when establishing
priority to conserve the accepted name regardless of the lifehistory or stage of the type.
Rossman et al. (2015a, b) and Réblová et al. (2016) have
provided recommendations for conservation or use of pleomorphic generic names in Dothideomycetes and Sordariomycetes. However, linking teleomorph and anamorph of
species is challenging. Pure cultures obtained from single
germinating ascospores often sporulate and anamorphs are
formed (Senanayake et al. 2020a). Sometime, strains of pleomorphic species obtained from different specimens cluster
together with strong support in phylogenetic trees (Karunarathna et al. 2017; Wanasinghe et al. 2018). The colony characters and nucleotide identity of molecular sequences should
be checked even if cultures do not sporulate. Further, some
unculturable ascomycetes that are only known from their
teleomorph cannot be linked to the anamorph.
However, some other classes such as Orbiliomycetes, the
generic names are much more complicated because of the
names based on single morphological differences without
molecular data (Baral et al. 2018). In Orbiliomycetes, a
narrow concept has used for the demarcating the generic
boundaries of the anamorphs, while a broad concept relies
on the teleomorphs. Therefore, more generic names have
been established for the anamorphs. Hence, it should be
avoided to adopt a certain generic concept prematurely, as
this may imply a lot of unnecessary name changes (Baral
et al. 2018). Therefore, linking teleomorph and anamorph of
species is an important practice in nomenclature and it can
affect species number.
Phenotypic plasticity
Phenotypic plasticity in fungi denotes that changes in morphology, behavior and physiology in response to the environmental variation (Price et al. 2003). Phenotypic plasticity allows teleomorphic ascomycetes to respond to climatic
changes within their lifetime (Williams et al. 2008). This is
important for species to survive as evolutionary responses
for climatic changes by natural selection takes time to make
any adaptation.
More than 40% of ascomycetes live in symbiosis as
lichens (Kirk et al. 2008). Lichens show high phenotypic
plasticity together with geographical distributions (Divakar
et al. 2013; Muggia et al. 2014). Parmeliaceae is a hyperlichenized fungal family mainly distributed in the tropics
(Kraichak et al. 2015). The type genus Parmelia includes
several distinct species by phenotypic plasticity (Valladares
et al. 2007). Parmelia discordans and P. omphalodes were
described based on morphological differences. However,
molecular data showed that these two species are conspecific
and phenotypic variations are made according to environmental changes (Divakar and Upreti 2005). Nipponoparmelia pseudolaevior and N. laevior show phenotypic plasticity
in this family (Molina-Montenegro et al. 2016).
Phenotypic stasis
Phenotypic stasis is explained by natural selection and
genetic drift, or by constraints imposed by mutation and
recombination of standing genetic variation (Mallard et al.
2019). This is a basic method in speciation and genetic variations (Chethana et al. 2020). Gene variations formed by
phenotypic stasis can completely disappear to reduce the
genetic variation or initially rare alleles become much more
frequent to dominant the gene variations (Mallard et al.
2019). However, the morphological variations formed by
phenotypic stasis are retained in a population if only individuals survive and reproduce. Phenotypic stasis forms species
morphologically similar, but genetically different. Therefore,
species estimates must include these species.
Homoplasy
Homoplasy is a trait that has been gained or lost independently in separate lineages with evolution (Torres-Montúfar
et al. 2018) and it can arise by selection pressures or genetic
13
Fungal Diversity
drift (Stearns and Hoekstra 2005; Hall and Colegrave 2008).
Homoplasy mostly appears in similarity of morphological
characters, but also in molecular sequences (Reece et al.
2011), life cycle (Silberfeld et al. 2010) and behavior (de
Queiroz and Wimberger 1993).
Jiang et al. (2020) showed in a phylogenetic study of foliicolous lichens that a new lineage sister with Strigulaceae
(Dothideomycetes) was formed, however morphologically
similar to Porina (Lecanoromycetes). This new clade represents a monogeneric family Tenuitholiascaceae which is typified by Tenuitholiascus with a single species T. porinoides.
This species is morphologically similar to the genus Porina
in external morphology, ascospore type, the thin-walled asci
and unbranched paraphyses. Further, Schmitt (2011) showed
that homoplasy affects the evolution of fruiting body type
and ascus at the class level within the phylum Ascomycota.
This may increase the species number described if only
based on phenotypic characters.
Synonyms and conspecific species
Synonym is a scientific name currently applies to a taxon
that goes by a different scientific nameand synonyms form
strong, monophyletic clades with currently applied taxon in
phylogenetic trees. Therefore, these synonyms are known as
conspecific species (Rossman et al. 2015a). Hence, there is
often more than one scientific names for a single species and
the morphs had been described in different genera (McNeill
et al. 2006). Two or more names for different morphs of
the same species are not accepted according to the Melbourne Code (McNeill et al. 2012). Hence, Wijayawardene
et al. (2012), Rossman et al. (2015a,b), Réblová et al. (2016)
have proposed recommendations to determine which name
to conserve. Proposals were based on excluding synonymy,
giving priority to basionyms, commonly used names or the
commonly occurring morph in nature.
Conspecific species being identified as distinct species
through morphological data, but with molecular data providing evidence for them being identical, has also led to
incorrect species identification. Some sexually compatible
conspecific fungal species can also produce new pathogens
via interspecific hybridization and reproductive interference
(Giordano et al. 2019). The genus Diatrype is typified by
D. disciformis (Fries 1849). Libertella betulina, the type
species of Libertella, is the anamorph of Diatrype stigma
(Grove 1937; Kutorga et al. 2006), while L. disciformis is
the anamorph of D. disciformis. Diatrype disciformis and
D. stigma are conspecific (Trouillas et al. 2010). Further,
Libertella is the older name as it was erected in 1830 while
Diatrype was only erected in 1842, thus Diatrype and Libertella are synonyms. However, Diatrype has a great number
13
of species including important plant pathogens. Hence,
Diatrype was recommended for protection over Libertella
(Réblová et al. 2016).
There are many recommendations proposed for taxa
of Xylariaceae (Réblová et al. 2016). The genus Daldinia
is typified by D. concentrica (Stadler et al. 2014). The
monotypic genera Annellosporium which is typified by A.
nemorosa and Versiomyces typified by V. cahuchucosus
Whalley & Watling, have been synonymized under Daldinia
as D. nemorosa based on the phylogeny and D. cahuchucosa based on morphology and chemotaxonomic evidences
(Stadler et al. 2014). Daldinia is common with many species
and has been recommended for use. Therefore, recognizing excluded synonyms is essential to estimate the actual
number of fungi.
Illegitimate and invalid names
Published taxonomic names may be illegitimate and invalid.
This means the species exist, but are nomenclaturally incorrect due to contravening some of the articles laid down by
the nomenclature codes. If a published species name is not
accepted as a proven valid species, then it can be superfluous as a synonym of a known species, non-compliant with
nomenclature codes thus considered a “bad” name and
doubtful name with insufficient study (Wang et al. 2019).
The number of accepted names, synonyms, invalid or illegitimate names, and unstudied names has been compared
by Wang et al. (2019). They found that accepted names
increased markedly over time and increased significantly
after the 1900s. The number of synonyms, invalid or illegitimate names increased slowly and it is evident that the
quality of fungal taxonomic work has improved with the
application of molecular techniques. The International Code
of Nomenclature for Algae, Fungi, and Plants is updated
every four years and proposes and regulates all the articles
related to nomenclature (Turland et al. 2018).
Introgression and natural hybridization
Introgression means transfer and incorporates alleles
from one species into the gene pool of another species by
hybridization and backcrossing (Schardl and Craven 2003;
Stukenbrock 2013, 2016; Restrepo et al. 2014). There are
many occasions when genetic information can be transfered
between closely related species and thus gene flow between
cryptic species has frequently been found (Hawksworth
2001; Bickford et al. 2007; Hawksworth and Lücking 2017).
Therefore, species boundaries in morphologically indistinct
species and species complexes may be doubtful (Barton
and Hewitt 1985; Barton and Gale 1993). If a portion of
Fungal Diversity
the introgressed gene pool of each of the hybridizing taxa
remains constant and uncontaminated then different distinct
gene pools can be recognized as new species.
Beneficial alleles tend to introgress easily for habitat
adaptation or reproduction (Barton 1979). Thus, patterns
of differential introgression across hybrid zones in genes
or genome regions are important for habitat adaptation and
speciation (Payseur 2010; Shaw and Mullen 2011; Nachman and Payseur 2012). Sometimes, the gene flow between
species is limited or prevented in nature by a set of basic
barriers. These limits control transfer of the genetic material which affects phenotypic variations between species and
determines if species reproduce individually (Bouck et al.
2005; Lemmon et al. 2007; Roe and Sperling 2007; Wagner
et al. 2013). However, the question is how many species
evolves presently by introgression and hybridization? The
answer is unpredictable. Therefore, this should be considered when estimate the teleomorphic species in Ascomycota.
Lumping and splitting of Xylaria species in phylogenetic
trees forming unresolved lineages has occurred over time.
Most phylogenetic analyses have shown that Xylaria species
do not form a monophyletic clade and are scattered within
Xylariaceae (Hsieh et al. 2010; Senanayake et al. 2015;
Maharachchikumbura et al. 2016; Wendt et al. 2018). Few
Xylaria species are clustered with Amphirosellinia, Astrocystis, and Collodiscula without strong statistical support
(Wendt et al. 2018; Konta et al. 2020).
Generic polyphyly does not markedly change the number
of existing species, but species polyphyly changes the number of species. The “special status” of a species comprises
the unique, observable morphological characters (Queiroz
and Donoghue 1988). It is implicitly assumed that species
are monophyletic or at least paraphyletic. However, hybrid
speciation arguably leads to polyphyletic species (Hörandl
and Stuessy 2010). Hybrid species are a common phenomenon that allows for rapid speciation (Linder and Risenberg
2004) and polyphyletic species develop into different species later.
“Man‑made” or domesticated species
Adaptive hybridization is used to obtain industrially important species additionally to natural hybridization (Burgarella
et al. 2019). Genetic materials are changed during domestication of wild species and new species or varities may form
(Shibata et al. 2015). Industrial cultivation of some teleomorphic ascomycetes such as Cordyceps, morels, truffles
requires hybrid varities to obtain high yield. However, the
obtained number of hybrid species in a particular period
is undetermined. Additionally, the behavior of hybrid species with the wild species gene pool is not well studied.
Therefore, an idea of the number of domesticated species
is needed to estimate the number of species in Ascomycota.
Polyphyletic genera and species
The polyphyletic nature of fungal genera derived from more
than one common evolutionary ancestor or ancestral group
cannot taxonomically be in the same genus. Sometime morphologically similar species cluster in different sub-clades
in phylogenetic trees representing several distinct genera
(Phookamsak et al. 2015, 2017; Konta et al. 2020). This
affects the number of described species. Some studies have
used slightly different morphological characters of taxa
along with the phylogenetic analyses to introduce new genera (Hyde et al. 2020a).
Xylariaceae comprises several polyphyletic genera which
are phylogenetically distantly related to each other (Peršoh
et al. 2009; Senanayake et al. 2015). Xylaria is typified by
X. hypoxylon (Schrank 1789; Greville 1824) and species in
Xylaria are saprobes or endophytes (Thomas et al. 2016).
Extinct or endangered species
The evolution of fungi begun around 1.5 billion years ago
(Wang et al. 1999; Brundrett 2002). There is evidence
that fungal communities in Ascomycota were present in
the Devonian period, 416–359 million years ago (StrulluDerrien et al. 2018). Ascomycetes diversified rapidly in terrestrial environments and therefore, they occupied numerous ecological niches. However, teleomorphic ascomycetes
are recently threatened by habitat loss, loss of symbiotic
hosts, pollution, over exploitation of the animal and plants,
destruction of ecosystems and climate change and they are
also becoming extinct (Wang et al. 1999). However, the
vast majority of teleomorphic ascomycetes have not been
assessed. The IUCN has listed 280 threaten fungal species
under several catogaries, such as critically endangered (CR,
15 species), endangered (EN, 59 species), vulnerable (VU,
90 species), near threatened (NT, 40 species), least concern
(LC, 54 species) and data deficient (DD, 22 species). The
IUCN Red List contains 46 threaten teleomorphic ascomycetes (Table 4) (IUCN 2021). The objective of the global
IUCN red list of threatened fungal species is to determine
conservation issues to the public and policy makers and help
the international community decrease species decline and
extinction (Lughadha et al. 2020). The IUCN Red List is the
most comprehensive, objective global approach for evaluating the conservation status of fungal species. The largest
number of threatened species is from Europe (IUCN 2021).
IUCN organized three workshops in Chile, and the UK in
2020 and this effort will assess the conservation status of
13
Fungal Diversity
Table 4 IUCN red-list category of teleomorphic species in Ascomycota issued by International Union for Conservation of Nature (IUCN)
Red-list category
Species name
Population trend
Classification
CR
CR
CR
CR
CR
CR
CR
CR
CR
EN
EN
EN
EN
EN
EN
EN
EN
EN
EN
EN
VU
VU
VU
VU
VU
VU
VU
VU
VU
VU
NT
NT
NT
NT
NT
NT
LC
LC
LC
LC
LC
DD
DD
DD
DD
DD
Acanthothecis leucoxanthoides
Acanthothecis paucispora
Buellia asterella
Erioderma pedicellatum
Hypocreopsis amplectens
Loxospora assateaguensis
Ramalina portosantana
Rinodina chrysomelaena
Sulcaria isidiifera
Arthonia kermesina
Cetreliopsis papuae
Cladonia perforata
Gymnoderma insulare
Ramalina confertula
Ramalina erosa
Ramalina timdaliana
Rinodina brodoana
Santessoniella crossophylla
Sticta alpinotropica
Sulcaria badia
Antrelloides atroceracea
Anzia centrifuga
Berggrenia aurantiaca
Caloplaca rinodinae-albae
Cetradonia linearis
Cyttaria septentrionalis
Lethariella togashii
Microglossum atropurpureum
Trichoglossum walteri
Xanthoparmelia beccae
Ascoclavulina sakaii
Geoglossum difforme
Gyromitra korshinskii
Leptogium rivulare
Pseudoplectania melaena
Sarcosoma globosum
Everniastrum nepalense
Mitrulinia sp.
Orbiliopsis callistea
Phaeophyscia hispidula
Poronia punctata
Biscogniauxia bartholomaei
Cordierites acanthophorus
Cordyceps hauturu
Cordyceps kirkii
Thuemenidium sp.
Stable
Stable
Decreasing
Decreasing
Decreasing
Stable
Decreasing
Decreasing
Decreasing
Decreasing
Unknown
Unknown
Decreasing
Stable
Stable
Unknown
Decreasing
Decreasing
Unknown
Decreasing
Decreasing
Unknown
Decreasing
Unknown
Decreasing
Decreasing
Decreasing
Decreasing
Decreasing
Decreasing
Unknown
Decreasing
Decreasing
Decreasing
Decreasing
Decreasing
Unknown
Unknown
Unknown
Unknown
Decreasing
Unknown
Unknown
Unknown
Unknown
Unknown
Lecanoromycetes, Ostropales, Graphidaceae
Lecanoromycetes, Ostropales, Graphidaceae
Lecanoromycetes, Teloschistales, Physciaceae
Lecanoromycetes, Peltigerales, Pannariaceae
Sordariomycetes, Hypocreales, Hypocreaceae
Lecanoromycetes, incertae sedis, Sarrameanaceae
Lecanoromycetes, Lecanorales, Ramalinaceae
Lecanoromycetes, Teloschistales, Physciaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Arthoniomycetes, Arthoniales, Arthoniaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Lecanoromycetes, Lecanorales, Cladoniaceae
Lecanoromycetes, Lecanorales, Cladoniaceae
Lecanoromycetes, Lecanorales, Ramalinaceae
Lecanoromycetes, Lecanorales, Ramalinaceae
Lecanoromycetes, Lecanorales, Ramalinaceae
Lecanoromycetes, Teloschistales, Physciaceae
Lecanoromycetes, Peltigerales, Pannariaceae
Lecanoromycetes, Peltigerales, Lobariaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Pezizomycetes, Pezizales, Pezizaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Incertae sedis, incertae sedis, incertae sedis
Lecanoromycetes, Teloschistales, Teloschistaceae
Lecanoromycetes, Lecanorales, Cladoniaceae
Lecanoromycetes, Cyttariales, Cyttariaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Geoglossomycetes, Geoglossales, Geoglossaceae
Geoglossomycetes, Geoglossales, Geoglossaceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Leotiomycetes, Helotiales, Helotiaceae
Geoglossomycetes, Geoglossales, Geoglossaceae
Pezizomycetes, Pezizales, Discinaceae
Lecanoromycetes, Peltigerales, Collemataceae
Pezizomycetes, Pezizales, Sarcosomataceae
Pezizomycetes, Pezizales, Sarcosomataceae
Lecanoromycetes, Lecanorales, Parmeliaceae
Leotiomycetes, Helotiales, Sclerotiniaceae
Leotiomycetes, Helotiales, incertae sedis
Lecanoromycetes, Teloschistales, Physciaceae
Sordariomycetes, Xylariales, Xylariaceae
Sordariomycetes, Xylariales, Xylariaceae
Leotiomycetes, Helotiales, Helotiaceae
Sordariomycetes, Hypocreales, Cordycipitaceae
Sordariomycetes, Hypocreales, Cordycipitaceae
Geoglossomycetes, Geoglossales, Geoglossaceae
CR critically endangered, EN endangered, VU vulnerable, NT near threatened LC least concern, DD data deficient
13
Fungal Diversity
endemic species of three regions, South America, Europe
and Southeast Asia.
However, the number of threaten teleomorphic ascomycetes
is very low compared with the described number of teleomorphic species in Ascomycota due to the difficulties in screening fungal populations. Most teleomorphic ascomycetes are
only visible when they produce fruiting bodies and may not be
found in the same place every year (Senanayake et al. 2020a).
Some early evolved teleomorphic ascomycetes have become
extinct and there is no documentation for them.
Biodiversity hotspots
Only about 93,000 Ascomycota species have been introduced and documented even more are estimated (Roskov
et al. 2019). Studying the fungi in biodiversity hotspots is
important to determine the undescribed taxa (Hawksworth
and Lücking 2017). Biodiversity hot spots and geographic
and ecological habitats which are poorly or under-studied are
major localities for these undescribed species. Biodiversity
hot spots are designated by IUCN and they are conserved
by government in located country (Marchese 2015). Biodiversity hotspots occupy approximately 1.4% of earth’s land
area, but 60% of Earth’s biodiversity is gathered there (Possingham and Wilson 2005; Marchese 2015).
The global biodiversity hotspots are referred to as areas
featuring exceptional concentrations of endemic species and
experiencing exceptional loss of habitat (Myers et al. 2000).
North America, Europe (France, Germany, Italy, Spain,
Sweden, UK), Japan, India, China, Taiwan, Thailand, Philippines, Australia and Brazil are the leading countries from
which most ascomycetous type collections were obtained
(Species Fungorum 2021). Most of these countries are
located in biodiversity hotspots, but the whole country has
not been preserved as biodiversity hotspots (Matutea and
Sepúlvedab 2019). Although many species have been collected from countries which are located in biodiversity hotspots, it does not mean that fungal diversity is restricted to
the biodiversity hotspots (Marchese 2015). Generally, rules
and regulations for entrance and utilization of reso urces are
strictly controlled in these areas, even for scientific studies.
This is one reason that mycologists could not estimate exact
number of teleomorphic ascomycetes in hotspots. However,
it is predicted that biodiversity hot spots have extremely
favorable conditions for ascomycetes and thus production
of sexual, thick-walled spores is not prominent. Therefore,
species in Ascomycota may occur in vegetative phase or
reproduce asexually.
Hidden species in niches
Morphologically, teleomorphs of ascomycetous fungi are
only recognized when they produce sexual reproductive
organs. Some ascomycetes occur in niches without forming
any visible, distinct fruiting structures and are only obtained
as hyphae (Schardl and Craven 2003). These species are ecologically cryptic and difficult to screen by traditional phenotypic approaches. Environmental DNA analyses can reveal
such species (Wu et al. 2019a). Additionally, fungal succession may help to obtain those species separate. This has
been defined as the sequential occupation of different ascomycetes or different associations of ascomycetes on the same
substrate or site (Challacombe et al. 2019). This happens
because of a sequence of sporulating fungi on a substrate
by mycelium. However, replacement of one ascomycetous
species by another is not necessary and some ascomycetes
sporulate together on a substrate (Hyde and Jones 2002).
Sometimes, incubation of fresh specimens is necessary to
obtain maximum fungal diversity, especially rare or slow
growing species.
Poorly‑studied fields
Many terrestrial teleomorphic ascomycetes have been
described, but fewer are known from aquatic habitats. Natural groundwater limestone aquifers are challenging and
unexplored fungal habitats (Krauss et al. 2003; Lategan
et al. 2012; Risse-Buhl et al. 2013). Mangrove and wetland
associated teleomorphic ascomycetes are poorly known as
compared to terrestrial taxa (Lee et al. 2019). Therefore,
it is necessary to fully screen marine and aquatic habitats
to explore the fungal diversity. Research on aquatic fungi
in recent years have incorporated molecular techniques to
achieve a better identification of taxa and many new species have been introduced (Zhang et al. 2017; Hyde et al.
2020a; Wei et al. 2020). Most of the earlier studies are based
on observation of morphological characters and culturedependent techniques (Luo et al. 2019). There are unculturable freshwater ascomycetes and hence, culture-independent
techniques need to be followed to understand the diversity
among teleomorphic, freshwater ascomycetes (Hyde et al.
2020b). Clinical mycology is another field which has not
been studied much with few human and animal pathogens
reported (Homei 2006; Pihet et al. 2009; Köhler et al. 2015;
Sullivan et al. 2015).
13
Fungal Diversity
Less‑studied sites in developing countries
Biodiversity hotspots in Asia (except Japan) and South
America are generally poorly studied and not well-known
(Hyde et al. 2020c). Most countries in these two continents
are economically impoverished developing countries and
lack resources and enough funding for fungal research (Subramanian 1986; Jones et al. 2007). Most developing countries are located in tropical and subtropical regions and fungi
thrive in these countries due to the favorable habitat conditions. However, some species rich, biodiversity hotspots in
these countries lack studies on species diversity and richness (Marchese 2015). In fact, the whole planet is not being
evenly screened and therefore, many important teleomorphic
ascomycetes may become extinct prior to description and
documentation.
In 2017, 35% of the world’s described fungal species were
from Asia (Willis 2018). This may have increased within the
last two years (Hyde et al. 2020c). Africa is very different from South America or Asia in biodiversity having dry
climates and different geographical lands, while there are
many rainforests and wet ecological sites in South America.
Therefore, less-studied areas should be examined well to
explore undescribed teleomorphic species in Ascomycota.
Unexamined collections in specimen
repositories and missing specimen records
Traditionally, teleomorphic ascomycetes have been distinguished by different approaches and concepts such as morphology, physiology, biochemistry or reactions to chemical
tests. Morphology was commonly used to introduce teleomorphic ascomycetes and it was improved with the innovation of the microscope. However, there are many poorly
identified or superficially examined collections in fungaria
(Senanayake et al. 2020a). Many reference specimens in fungaria and living cultures in fungal culture collections are not
named to the species level, and in some cases not even to the
generic level (Hawksworth and Lücking 2017).
Additionally, some collections and isolates are connected
to existing species names without comprehensive specimen
examination and sometimes it is possible to find different
fungi in the same specimen if those specimens are examined carefully. It has been suggested that morethan 20,000
teleomorphic ascomycetes have been collected worldwide
but not described yet (Hawksworth and Rossman 1997).
Mycologists often have a large collection of material waiting for formal introduction and description. Some taxonomic
records, descriptions and specimens have been lost in various situations before description (Senanayake et al. 2016).
Therefore, re-visiting early-introduced ascomycetes which
13
are only based on morphology is important (Ariyawansa
et al. 2014).
Some monographic work and some journal issues that
focus on exploration of global fungal biodiversity such as
Mycosphere Notes (ZUAE, Guangzhou, China), Fungal
Diversity Notes (Springer Nature, Switzerland) or Fungal
planet description sheets (CBS, The Netherlands) describe
hundreds of novel teleomorphic ascomycetes annually. However, some species are introduced only based on morphology
and Mega-blast similarities in GenBank without phylogenetic analysis. Most of the new collections are described
without re-visiting morphologically similar earlier species
or genera. If a collection is from known host, then it is necessary to check previous species records on the host and
locality (country or continent) prior to describing a new species (Senanayake et al. 2020b). Careless practices will result
in proliferation of synonyms or invalid names leading to
confusion and instability in nomenclature. Many described
teleomorphic ascomycetes in the pre-molecular era are only
based on morphology and there are older collections in
repositories that need to be examined and described.
Incomplete descriptions of earlier described
species
In earlier studies, specimens were examined by light microscope and 40 was the maximum magnification (Senanayake
et al. 2020a). Lenses of light microscopes were calibrated
manually (Zhang et al. 2016). One unit of the ocular micrometer disc is needed to calibrate against a known length and
measurements were subjective from person to person.
Newer technologies facilitate measurements directly when
photographing and software is already calibrated. Hence,
errors in measurements have been less in recent descriptions. Earlier prologues based on the limited characters and
some important, inconspicuous characters were not examined or not mentioned (Senanayake et al. 2018). Therefore,
new collections of existing species are mistakenly described
as new species. A prologue should include information of
examined materials (Seifert and Rossman 2010). However,
some specimens lack these details or are incomplete. Earlier
prologues usually do not contain culture characters. Therefore, incomplete descriptions of earlier introduced species
make complications in species identification.
Synanamorphism
Synanamorphism is a teleomorphic species produces several
morphologically distinct anamorphs and these synanamorphs have been often reported from cultural studies (Crous
et al. 2009; Réblová et al. 2021). Before molecular data
Fungal Diversity
was available for taxonomic studies, species delimitation
was based on the morphology of specimens and cultures.
Pure cultures were used to obtain teleomorphs or anamorphs and then to describe the holomorph. However, some
teleomorphic species produce several anamorphs and those
anamorphs are described as different species. Molecular
phylogenies of some studies proved that some teleomorphic
species produce two or more anamorphs (Fan et al. 2018;
Réblová and Štěpánek 2018). Therefore, it is assumed that
some synanamorphic species might have been considered as
separate species dueto their distinct morphologies and they
should avoid from the unmber of known species.
Species in Cytosporina Sacc., Libertella Desm., and
Naemospora Sacc. have been reported as anamorphs of
diatrypaceous species (Glawe and Rogers 1984). However,
the morphology of the anamorph is not useful when differentiating taxa in Diatrypaceae either at the genus or species level. There is indistinguishable morphology and many
species produce different types of anamorphs in the same
culture (Rappaz 1987). Additionally, different types of conidiogenesis have been reported in the same Diatrype strain
(Glawe and Rogers 1982). Eutypella parasitica produces
both pycnidia and acervuli on both natural substrata and
cultures (Glawe 1983). Further, the anamorph of Hapalocystis (Sydowiellaceae, Diaporthales) is reported as Stilbospora, Hendersonia or Dothiorella (Wehmeyer 1941; Castlebury et al. 2002), or as stilbospora-like taxa (Barr 1978). A
phoma-like anamorph is reported for H. berkeleyi in culture
(Glawe 1985; Liu et al. 2015; Senanayake et al. 2016).
Absence of molecular data for described
species
Before the 1990s, fungal species were introduced based on
morphology and there are no molecular data for many of
those described species. Classification of those species is
challenging without molecular data (Huang et al. 2021).
Cryptic species, phenotypic plasticity, phenotypic stasis,
pleomorphism, and homoplasy between teleomorphic ascomycetes are difficult to reveal without molecular data (Maharachchikumbura et al. 2014). Hence, the estimated number
of teleomorphic species in Ascomycota should include those
hidden species.
Addition to the phylogenetic relationships, molecular
data reveals the character evolution of teleomorphic species in Ascomycota. Character evolution deals with the
process of evolution of a trait along the branches over a
period of time from a common ancestry (Hongsanan et al.
2018). This explains the history of life, the relationships
among extant species and character states for each species
(Vijaykrishna et al. 2006). Most character evolution studies were carried out after the 1990s (Liu and Hall 2004; Li
et al. 2005; Schoch et al. 2009; Schmitt 2011; Kumar et al.
2012) and these studies have not been obviously conducted
on teleomorphic ascomycetes which lack sequence data.
The unavailability of complete sets of sequence data is the
major issue for absence of character evolution studies. These
studies concur or argue against species delimitation. Hence,
these studies influence the estimate number of teleomorphic
species in Ascomycota.
Apothecia are the primitive fruiting body type of Pezizomycotina. However, the formation of perithecia and cleistothecia is still unclear (Hongsanan et al. 2018). The amyloid
reaction of stromatic tissues occurs in some species in Sordariomycetes such as Hypocreaceae in Hypocreomycetidae,
Xylariaceae in Xylariomycetidae and Cryponectriaceae in
Diaporthomycetidae. However, the reason for the appearance and disappearance of the same character during evolution and when, how and why these characters evolved are
unclear. If there is a complete set of molecular data, character evolution of teleomorphic species in Ascomycota can
be predicted clearly and it provides additional taxonomic
value for a species.
Molecular data unavailability
for both teleomorph and anamorph
of ascomycetes
Phenotypic species were previously introduced based on
morphological characters and later, sequence data obtained
for some of earlier described species. However, many species lack sequence data. If a culture of a teleomorphic
Ascomycota sporulates then, the anamorph can be obtained
and identified. These teleomorphs and anamorphs can be
linked if they are derived from a pure culture. If a specimen contains both ascomata and conidiomata in close to
each other, then cultures obtained from both ascospores and
conidia should be further analysed for molecular and colony
morphology. If the colony morphology in the same media
and at the same maturity is similar and sequence data are
identical, then the conidiomata are anamorph of teleomorphic fungi and it should be named as one species. However,
colony characters and sequence similarity can be used if
teleomorphs and anamorphs are described from different
specimens. The sequence quality should be high and both
forward and reverse directions are needed.
Senanayake et al. (2015) introduced the genus Ciferriascosea Senan., Bhat, Camporesi & K.D. Hyde with two teleomorphic species C. fluctuatimura Senan., Bhat, Camporesi
& K.D. Hyde and C. rectimura Senan., Bhat, Camporesi &
K.D. Hyde and they were accommodated in family Phlogicylindriaceae (Xylariales). The anamorph was not reported
for this genus. Phlogicylindriaceae is typified by anamorphic
genus Phlogicylindrium Crous, Summerb. & Summerell and
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Fungal Diversity
teleomorph is not reported for this genus. Therefore, species in
Ciferriascosea and Phlogicylindrium cannot be morphologically compared. Phylogenetically, species in these two genera
clustered together with high bootstrap support. However, ITS
loci of the type species of Ciferriascosea, C. rectimura showed
10.4% base pair differences from ITS loci of the type species
of Phlogicylindrium, P. eucalypti (Jeewon and Hyde 2016).
Further, 85 base pairs are missing in the ITS loci of Phlogicylindrium eucalypti. Additionally, Ciferriascosea rectimura
was isolated from twigs of Spartium junceum L., while Phlogicylindrium eucalypti was collected from leaves of Eucalyptus
globulus Labill. Therefore, Senanayake et al. (2015) proposed
a new genus Ciferriascosea for this teleomorphic taxon instead
of proposing it as the teleomorph of Phlogicylindrium.
Camarosporidiella caraganicola was introduced based
on a strain derived from the anamorph that was described
by Liu et al. (2015). Wanasinghe et al. (2017) has examined several specimens of the teleomorph of Camarosporidiella caraganicola. Phylogenetically, these strains clustered
together with strong support. Both teleomorph and anamorph were collected from the same host (Caragana frutex (L.)
K.Koch) in the Rostov Region, Russia. Therefore, Wanasinghe et al. (2017) linked these teleomorph and anamorph
together as the holomorph of Camarosporidiella caraganicola by considering the host similarity and statistical support
in phylogeny.
A teleomorph of an astragalicola-like taxon was collected
and obtained in culture (MFLUCC 17˗0832). The multigene sequence analyses showed that this isolate clusters with
Astragalicola amorpha with strong support (Wanasinghe et al.
2018). However, A. amorpha is known from its anamorph
and thus, it is not possible to compare their morphologies. A
comparison of the ITS loci of these two strains revealed 2.12%
nucleotide differences, which justifies these two isolates as
two distinct taxa (Jeewon and Hyde 2016). Therefore, the new
collection was introduced as Astragalicola vasilyevae.
The anamorph of Wojnowiciella dactylidis was illustrated in Liu et al. (2015) and molecular data was provided.
Karunarathna et al. (2017) illustrated the teleomorph for this
species. Both teleomorph and anamorph were collected from
Italy on different host plants but colony characters of both
strains are similar. Phylogenetically, both strains clustered
together with strong support. The ITS sequences obtained
from the teleomorphic strain was identical to anamorphic
strain with 100% base pair similarity. Therefore, Karunarathna et al. (2017) linked these two strains as the holomorph
of Wojnowiciella dactylidis.
13
Teleomorphic ascomycetes described based
on only Internal Transcribed Spacer (ITS)
region
The nuclear ribosomal internal transcribed spacer (ITS)
region is located between the sequences encoding the small
(SSU) and large (LSU) subunits of the ribosomal operon
(White et al. 1990). The ITS region is the formal primary
fungal barcode with the highest probability of correct species identification of a broad group of fungi (Horton and
Bruns 2001; Bridge et al. 2005; Martin and Rygiewicz 2005;
Seifert 2009; Bellemain et al. 2010; Schoch et al. 2012).
More than 100,000 ITS sequences of teleomorphic ascomycetes are deposited in international nucleotide sequence
databases (Baturo-Cieśniewska et al. 2020). Even though,
many sequences of correctly identified species are deposited
in databases, there are some sequences with technical errors,
incorrectly named species, atypical chimeric ITS sequences,
and sequences verified only at the generic level or above
which makes reliable problems in species identification
(Nilsson et al. 2009, 2014; Hongsanan et al. 2018). The ITS
locus in species of some genera has minimal molecular variation (Andrew et al. 2009) and cannot resolve species (de
Hoog and Horré 2002).
Further, the ITS region has additions and deletions in
some groups of fungi and is not equally variable in all
taxa (Nilsson et al. 2008). Most databases are improving
sequence availability. The lack of sufficient ITS sequences
especially in species-rich, morphologically indistinct genera,
can therefore be problematic (Seifert 2009). A high-fidelity,
universal primer pairs for amplification of the EEF1A1
gene has been developed as secondary DNA barcodes for
the fungi (Stielow et al. 2015). However, the availability of
deposited reference sequences of EEF1A1 gene in databases
is poor (Meyer et al. 2019). Further, mycologists who have
deposited sequences in the NCBI database determine the
species name by themselves. The introduction of incorrectly
identified sequences creates in errors because users adopt
incorrect species names (Ko et al. 2011; Nilsson et al. 2012,
2019). In most species, the nucleotide composition of the
ITS sequences in the single species is identical or slightly
varied. The ITS locus has certain interspecific variations
(Nilsson et al. 2008; Talgø et al. 2010). These interspecific
variations in ITS sequences may be higher than 1% due to
technical errors, may be evidence to be a new species (Jeewon and Hyde 2016).
As an example, 245 ITS sequences of Diaporthe eres
obtained from GenBank and those sequences were aligned
using default settings of MAFFT v.7 with Diaporthella
corylina (CBS 121124) as the out group. Maximum likelihood analysis was performed after excluding 105 identical sequences and then resulted in 14 different clades
Fungal Diversity
Fig. 4 Phylogram generated from maximum likelihood analysis based
on ITS sequence data of Diaporthe eres. The tree is rooted with Diaporthella corylina (CBS 121124) and ex-epitype strain of D. eres is
in red bold. Subclades which are representing the different type of
sequences are labeled A to N
13
Fungal Diversity
Fig. 4 (continued)
(Fig. 4) representing 14 types of sequence patterns. The
ex-epitype strain of Diaporthe eres (AR5193) clustered
in clade A, while clades B-E showed around 1% base-pair
difference with sequence pattern of clade A (A-B:1.1%,
A-C: 1.3%, A-D:1.1% A-E:1.3%). However, clades F-N
13
showed more than around 1% base pair variation than
clades A (A-F:4.0%, A-G:3.7%, A-H:3.9%, A-I:4.4%,
A-J:4.4%, A-K:4.6%, A-L:5.2%, A-M:5.0%, A-N:5.4%).
According to the guidence of Jeewon and Hyde (2016), 1%
position ambiguities in ITS with 450 base pairs can be a
Fungal Diversity
new species. In our analysis, B-E types of sequences show
fewer base-pair differences than other patterns of sequence
types showing certain interspecific variations (Nilsson
et al. 2008; Talgø et al. 2010). However, clades F-N show
more base-pair differences than the epitype strain due
to technical errors of sequencing or misidentification of
fungi. Therefore, newly obtained strains should always be
compared with the sequences of ex-type strains and extype strains should be included in phylogenetic trees.
Further, ITS based species identification may not be
suitable when interspecies nucleotide variations are less
(Baturo-Cieśniewska et al. 2020). Sclerotinia sclerotiorum
and S. trifoliorum are causative agents of clover rot disease
and the symptoms are identical (Vleugels et al. 2012). ITS
sequences of Sclerotinia species are almost identical (Freeman et al. 2002). Sclerotinia sclerotiorum differs from S. trifoliorum by one nucleotide, guanine or thymine in position
120 (Njambere et al. 2008; Baturo-Cieśniewska et al. 2017).
Therefore, ITS based species identification is not suitable
for this situation. Interspecific and intraspecific variation of
ITS sequences is often low for some species. Phacidium
fennicum differs from P. lacerum by two base pairs while
others share identical ITS sequences but are morphologically
distinct (Crous et al. 2014; Tanney and Seifert 2018). Therefore, use of ITS sequences together with morphology and at
least a protein coding gene should be emphasized (Pryor and
Michailides 2002; Schubert et al. 2007; Samson et al. 2014;
Robbertse et al. 2017; Bensch et al. 2018).
Next generation sequencing (NGS)
and meta‑genomic DNA
NGS or high-throughput sequencing methods are the modern sequencing technologies (van der Heijden et al. 2008).
These approaches have several advantages in that an initial knowledge of the genome or genomic features are not
required, single-nucleotide resolution facilitates detection of
related genes, alternatively spliced transcripts, allelic gene
variants and requires less DNA/RNA as input and has higher
reproducibility (Peršoh 2015). Diversity and distribution
patterns of teleomorphic ascomycetes in ecosystems give
important evidence on species number, ecosystem functions
and stability (Schmit and Lodge 2005). Microscopic observations and culture-dependent approaches cannot efficiently
calculate the number of teleomorphic species of Ascomycota
in environmental samples such as soil, water, and air as the
majority of them is generally complex, unculturable and not
visible without a microscope (Mitchell and Zuccaro 2006;
Stewart 2012; Hongsanan et al. 2018).
Authentic physical specimens do not exist for these species and they are therefore known as “dark taxa” (Tedersoo
and Smith 2017; Ryberg and Nilsson 2018). There are no
given any formal names to genus and species level, and
hence types are not designated (Taberlet et al. 2012; Herder
et al. 2014; Hawksworth et al. 2016). Article 40 (Valid publication of names; section 2: Names of new taxa) in Shenzhen
Code states that a new taxon at the rank of genus or below
published on or after 1st January 1958 is valid only when
the type specimen of the name is indicated (Turland et al.
2018). Therefore, high-throughput sequencing approaches
are needed to reasonably characterize and estimate those
fungal communities.
However, there are some limitations in meta-DNA
sequencing and many mycologists disagree with validating
names. Concerns of the mycological community against the
premature introduction of DNA-only based nomenclature
have been presented (Hongsanan et al. 2018; Thines et al.
2018; Zamora et al. 2018). However, DNA meta-barcoding
has the potential to provide a much better understanding
the species diversity and richness in fungal communities
(Heeger et al. 2018) and meta-genomic DNA has provided
clues as to where and which teleomorphic ascomycetes can
be found and this is applicable in estimation of the species
number, but it cannot be used for classification.
Environmental samples are poorly linked to species-based
databases (Hibbett et al. 2011). Therefore, errors in communication have occurred from one publication to another
when using the system databases (Hibbett et al. 2011;
Ryberg and Nilsson 2018). Thus, metagenomics DNA-based
nomenclature can be accepted at least for species number
estimations and fungal diversity studies (Hawksworth et al.
2016; Ryberg and Nilsson 2018). There is uncertainty when
naming a species, if the morphology is indistinct or DNA
sequences fail to resolve species relationships. Eventhough
the species lack morphology, genomic and ecological significance supported this introduction. Khan et al. (2020)
introduced two new species, Archaeorhizomyces victor and
A. secundus (Archaeorhizomycetes, Taphrinomycotina,
Ascomycota) based on the distinct base pairs comparison
of the internal transcribed spacer region ITS1 and ITS2 with
similar taxa obtained from environmental DNA.
Buèe et al. (2009) found 1000 operational taxonomic
units (OTUs) in 4 g of soil. A study on litter decomposition in temperate forests based on meta-DNA confirmed
that Ascomycota species have highest relative abundances
in the later stages of decomposition (Voříšková and Baldrian
2013). Therefore, high-throughput sequencing methods can
be used for fungal diversity estimates. Further, species estimates from different studies may not be compared directly
due to differing laboratory standards, protocols and data processing methods (Lindahl et al. 2013; Purahong et al. 2017).
However, determination of a common NGS platform, primer
pairs and re-analyzation of missing or non-matching gene
13
Fungal Diversity
regions may allow comparison of the results of NGS across
different studies and biomes (Nilsson et al. 2011).
How to avoid errors in species identification
and estimation of teleomorphic ascomycetes
Species identification and description gives fundamental
data for prediction of species number. Hence, documentation
of species names in data bases without errors is important.
Here we discuss some guidelines to improve this.
Documentation and personal errors
DNA sequences in public databases are annotated by the
submitting authors and mostly, further validation by the
curators in databases is not often. This approach creates
erroneous taxonomic sequence labels (Kozlov et al. 2016).
The novel sequences are annotated based on existing ones
and mislabeled sequences induce downstream errors.
Some sequences in databases are confusing because the
same species are classified with different names (Ashelford et al. 2005). This may increase synonyms which are
hidden under previous classifications (Bidartondo 2008;
Nilsson et al. 2008; Robbertse et al. 2017). Therefore, the
taxonomic credibility in sequence databases is with errors
(Binder et al. 2005; Bridge et al. 2005; Nilsson et al. 2006).
Blast similarities of sequences sometimes give uncultured
or unverified sequences and names not identified to species
level (Baturo-Cieśniewska et al. 2020). The major reason for
these unidentified or insufficiently identified sequences in
public databases is that most international journals require
all sequences used in the manuscript to be deposited (Ryberg
et al. 2008). Errors in protein coding gene sequences give
wrongly translated protein sequences which are sometime
named as “unverified”. Further, all species in a genus necessary to form a well-supported, monophyletic clade with the
type strain of the type species in the genus and the newly
introduce species must be in this monophyletic clade without
distantly cluster from generic type.
Senanayake et al. (2016) studied the taxonomy and phylogeny of phomatospora-like taxa and showed that Paramicrothyrium chinensis H.X. Wu & K.D. Hyde has 99%
similarity to Phomatospora biseriata. Wu et al. (2011)
introduced Paramicrothyrium based on P. chinensis using
morphology and molecular data. However, the combined
LSU and SSU analysis (Wu et al. 2011) showed Paramicrothyrium chinensis as morphologically close to Microthyrium, but phylogenetically distant from Microthyrium. Further, Singtripop et al. (2016) showed that Paramicrothyrium
chinensis (IFRDCC 2258) clusters with Chaetothyrina mangiferae (Micropeltidaceae) with high support. Hence, these
sequences might have some errors (Senanayake et al. 2016).
13
Voglmayr et al. (2018) revealed problems in some
xylarialean sequences. The LSU part included in the ITS
sequences of KP297406 and KP297396 which were deposited as Anthostomella helicofissa and A. forlicesenica
respectively, in NCBI searches show obvious variations
in the sequence alignment. Further, the LSU sequence of
KP340547 was revealed as xylarialean by BLAST searches;
however, it has 60 nucleotides difference in LSU sequences
with other Anthostomella species. A BLAST search of LSU
and RPB2 sequence of KP340538 and KP340524 from
MFLUCC 14-0007 (A. forlicesenica) revealed various pleosporalean taxa as the closest matches. Therefore, most
scientific journals have recently made it a prerequisite that
the sequence data and the alignments of novel taxa must be
provided for review to have better quality control.
There is a 1% intra-species variation within Aspergillus
niger (Henry et al. 2000). The ITS sequence of Aspergillus
niger (MK461010) is identical to ITS sequences of Alternaria alternata (Baturo-Cieśniewska et al. 2020). Further,
the ITS sequences of Aspergillus niger (MK461010) shows
10% inter-species variation of ITS sequences of other A.
niger strains. This probably suggests a misnomer of the
MK461010 sequence. This happens because the person who
deposits sequences in the NCBI database determines the
species affiliation. The deposition of incorrectly identified
sequences creates errors and mistakes because users obtain
incorrect species names (Ko et al. 2011; Nilsson et al. 2012).
About 65% of the GenBank entries are mislabeled sequences
or from poorly characterized vouchers (Leray et al. 2019).
Common, standard data base for taxonomic
information of fungi
Recently, most scientific data are documented in websites
(Hyde et al. 2020b). Websites gives quick and easy access,
sorting and filtering facilities and safeness for data. Currently, there are several databases for molecular, taxonomic
and ecological information of teleomorphic ascomycetes
(Rossman 1994). There is no standardized protocol for introducing new teleomorphic species in Ascomycota (Seifert
and Rossman 2010; Senanayake et al. 2020a). However,
there are some formal requirements for proposing names
that are imposed by the Nomenclatural Codes (Turland et al.
2018). All the taxonomic information must be available in
public for future researchers including taxonomic characters, descriptions, prologues, sequence data, phylogenetic
analysis, type specimens, ex-type cultures and ecological
data. However, the mycologists have their own preferences
and different policies when selecting databases to deposit
taxonomic information. Therefore, taxonomic data are scattered in various databanks and this makes difficulties when
comparing phenotypic characters. An introduction of “combined platform for taxonomic information of teleomorphic
Fungal Diversity
ascomycetes” with a unique identity number for each species
is essential. Then, the entire phenotypic, genotypic, chemotaxonomic, ecological, evolutionary, biological and classification data of a teleomorphic species in Ascomycota can
be obtained and compared easily. All the data of species can
be updated with the newly obtain information. This kind of
universal database would minimize errors due to poor updating and also personal arguments. Hibbett et al. (2016) listed
commonly used databases and tools for fungal classification
and identification.
The number of described, and accepted species in Ascomycota has been counted since 1943 through Ainsworth and
Bisby’s Dictionary of the Fungi and currently, the Catalogue
of Life web site (https://www.catalogueoflife.org/index?taxon
Key=F), which is updated annually through the Species Fungorum, The Integrated Taxonomic Information System and
The Global Information System for Lichenized and nonlichenized Ascomycetes outputs.
A standard approach to demarcate species
boundaries and describe teleomorphic species
in Ascomycota
As there are different approaches to designate a species,
mycologysts use different methods according to their requrements and those approaches demarcate species boundaries
based on different criteria. Hence, the species numbers vary
from one to another and this results complications in species
identification and descriptions in Ascomycota (Maharachchikumbura et al. 2021). Therefore, it is necessary to designate
a common criterion for species introduction and description
(Pažoutová et al. 2013; Kamil et al. 2018). The number of
described teleomorphic ascomycetes has increased with the
use of molecular techniques for species delimitation. Most
teleomorphic ascomycetes introduced earlier than the 1990s
lack molecular data from their type collections (Wu et al.
2019a). Hence, it is impossible to compare genotypic characters of fresh collections with genotypic characters of available
materials. Therefore, mycologists may misinterpret those collections as new species. Introduction of previously described
teleomorphic ascomycetes as new species with molecular data
has increased the number of teleomorphic species in the Ascomycota. Further, this value has accelerated with the ending of
the separate naming of teleomorph and anamorph of the same
species. Therefore, it is necessary to follow well-defined, sets
of criteria when introducing a new species and obtain molecular data from the new collection revealing the genetically close
taxa. Sequence data should be obtained from both conserved
and variable gene regions and all the genes or loci are blasted
with GenBank for base pair similarities. Morphological characters could be compared with all phylogenetically similar taxa
to ensure that it is not an existing species and also it should be
compare with all species in the genus which molecular data
unavailable. Ecological data, colony characters, and secondary metabolites provide additional information. Sporulation
or conidiation of cultures is important. Describing both teleomorph and anamorph characters and linked them as one is
required.
Discussion and conclusion
Assessment of the actual number of teleomorphic species in
Ascomycota and their diversity is important for systematics,
resource utilization, industrial production and environmental
management (Maharachchikumbura et al. 2021). Traditionally, species estimations were based on the numbers of fungi
recorded on particular plants and insects. There are several
estimations such as around 0.1 million by Bisby and Ainsworth (1943) to 2.2–3.8 million by Hawksworth and Lücking (2017). Traditional species estimation approaches have
several drawbacks. The major limitation of these estimates
is that they only focus on the fungi that either produces fruiting bodies, which can be identified using a microscope, or
those that can be easily cultured on artificial media (Duong
et al. 2006). Many ascomycetous endophytes do not sporulate in culture (White and Cole 1986), while some hostspecific species need host tissues to sporulate. Therefore,
it is a bias estimate of teleomorphic ascomycetes based on
microscopic examination of fruiting structures on substrata
or media (Guo et al. 2001; Promputtha et al. 2004). Several
studies have shown that most species are singletons found
only once (Anslan et al. 2016) or found in only a few locations (Wu et al. 2013) however with plants and animals,
some teleomorphic ascomycetes are endemic species. Even
though fungi are ubiquitous, individual fungal species are
restricted to a specific niche. Therefore, the distribution of
teleomorphic ascomycetes is distinct and probably the fungal species number could be much higher than the current
estimates (O’Brien et al. 2005; Blackwell 2011).
The development of molecular techniques, such as next
generation sequencing has helped identify the previously
undescribed diversity in the teleomorphic ascomycetes
(Raja et al. 2017; Wilson et al. 2019). Obtaining molecular sequence data from environmental samples from a wide
range of localities and substrata has generated a new source
of data for estimating species numbers (Wu et al. 2013).
The numerous sequences obtained through high-throughput
sequencing do not have any close matches with described
fungi in sequence databases (Lindahl et al. 2013) and this
shows that the current estimated species number is too low.
The occurrence of more uncultured teleomorphic ascomycetes also supports that the diversity of fungal species
is in underestimation (Hawksworth 1991). However, the
main problem in estimates is merging data obtained from
traditional culture-based approaches with data obtained
13
Fungal Diversity
from high-throughput sequencing methods (Buée et al.
2009; Kubartová et al. 2012; Dissanayake et al. 2018). Several studies disagree to use sequences derived from highthroughput sequencing methods because of various reasons
such as NGS detects only the predominant fungi in a sample
instead of all available species (Dissanayake et al. 2018),
specific PCR primers need to be developed for some ascomycetes, and the ITS locus is longer than the NGS sequences
(Kruse et al. 2017). Further, many ascomycetous species are
not well-separated into well-resolved species with a single
gene region. Therefore, NGS are suitable to separate fungi
only at the genus level (Purahong et al. 2017).
OTUs are also detected by other molecular approaches
based on DNA sequence data such as TGGE (thermal gradient gel electrophoresis), DGGE (denaturing gradient gel
electrophoresis), SSCP (single-strand conformation polymorphism), RFLP (restriction fragment length polymorphism), TRFLP (terminal restriction fragment length polymorphism), ARDRA (amplified ribosomal DNA restriction
analysis), 454 pyrosequencing and illumina MiSeq sequencing can be used to establish a better estimate of species
numbers (Hawksworth and Lücking 2017). These methods
provide species-specific sequences and therefore, the OTUs
generated in these methods reveal an enormous, unprecedented magnitude of fungal diversity.
Wu et al. (2019a, b) estimated that the total fungal diversity is about 12 million species based on meta-DNA and
culture-independent methods. In culture-dependent methods,
teleomorphic species in Ascomycota are mostly collected
and isolated based on their conspicuous fruiting bodies.
However, more fungal species are detected by cultureindependent approaches and the fungal species detected
by culture-independent and culture-dependent approaches
often do not overlap (Zhang et al. 2010). Therefore, cultureindependent methods are important in estimates of the teleomorphic species in Ascomycota. Hence, neither culturedependent nor culture-independent method can accurately
determinethe structure of a given community. Because of
the intrinsic selectivity of each method, the probability of a
given species being detected often differs with the method
(Zhang et al. 2010).
The above-mentioned problems could be avoided with
appropriate morpho-molecular phylogenetic approaches and
proper taxon sampling. Furthermore, polyphasic taxonomy
should be the preferred approach when introducing new teleomorphic species in Ascomycota, as the conclusions of one
method may be eliminated or justified with evidence from
the other methods (Maharachchikumbura et al. 2021). Furthermore, molecular data of more extype strains should be
included in the phylogenetic analyses as these would provide
more clarification where misidentified pleomorphic fungi
could be correctly placed. This has not been followed universally by mycologists, thus leading to misidentifications
13
of many fungal species. The best approach to avoid errors
in future taxonomic studies of teleomorphic ascomycetes is
to focus on increasing missing sequence data and linking
those data to digital voucher specimens (Lücking et al. 2020;
Jayawardene et al. 2021).
Acknowledgements Ning Xie thanks National Key R&D Program
of China (2021YFA0910800), National Natural Science Foundation
of China (No. 31601014), Basic and applied basic research fund of
Guangdong Province (2121A1515012166), Stability Support Project
for Universities in Shenzhen (20200812173625001) and Project of
DEGP (2019KTSCX150) for funding. Senanayake thanks to Paul
Kirk, Samantha C. Karunarathna for data contribution. S.N. Wijesinghe
would like to acknowledge Thailand Science Research and Innovation
(TSRI) grant for Macrofungi diversity research from the LancangMekong Watershed and Surrounding areas (Grant No. DBG6280009).
Dhanushka Wanasinghe thanks the CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (number
2021FYB0005), the Postdoctoral Fund from Human Resources and
Social Security Bureau of Yunnan Province and the National Science
Foundation of China. Saowaluck Tibpromma would like to thank the
International Postdoctoral Exchange Fellowship Program (Number
Y9180822S1), CAS President’s International Fellowship Initiative
(PIFI) (Number 2020PC0009), China Postdoctoral Science Foundation
and the Yunnan Human Resources, and Social Security Department
Foundation for funding her postdoctoral research. Rungtiwa Phookamsak thanks to CAS President’s International Fellowship Initiative (PIFI)
for young staff (Grant No. 2019FYC0003) and “High-level Talent Support Plan” Young Top Talent Special Project of Yunnan Province.
Declarations
Conflict of interest The authors declare that there is no confict of interest.
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long
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Authors and Affiliations
Indunil C. Senanayake1,2 · Dhandevi Pem3,4 · Achala R. Rathnayaka3,4 · Subodini N. Wijesinghe3,4
Saowaluck Tibpromma5,7 · Dhanushka N. Wanasinghe5,6,7 · Rungtiwa Phookamsak5,6,7 ·
Nuwan D. Kularathnage2,3,4 · Deecksha Gomdola3,4 · Dulanjalee Harishchandra3,4,8 ·
Lakmali S. Dissanayake9 · Mei‑mei Xiang2 · Anusha H. Ekanayaka3,4,5 · Eric H. C. McKenzie10 ·
Kevin D. Hyde2,3,4,5 · Hao‑xing Zhang1 · Ning Xie1
1
Guangdong Provincial Key Laboratory for Plant Epigenetics,
College of Life Science and Oceanography, Shenzhen
University, Nanhai Avenue, Nanshan, Shenzhen 3688,
Guangdong, China
2
Innovative Institute for Plant Health, Zhongkai University
of Agriculture and Engineering, Haizhu District,
Guangzhou 510225, China
3
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
4
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
5
Honghe Center for Mountain Futures, Kunming
Institute of Botany, Chinese Academy of Sciences,
Honghe County 654400, Yunnan, China
6
Centre for Mountain Futures (CMF), Kunming Institute
of Botany, Kunming 650201, Yunnan, China
13
·
7
CIFOR-ICRAF China Program, World Agroforestry
(ICRAF), Kunming 650201, Yunnan, China
8
Beijing Key Laboratory of Environment Friendly
Management On Fruit Diseases and Pests in North
China, Institute of Plant and Environment Protection,
Beijing Academy of Agriculture and Forestry Sciences,
Beijing 100097, China
9
Engineering Research Center of the Utilization
for Characteristic Bio-Pharmaceutical Resources
in Southwest, Ministry of Education, Guizhou University,
Guiyang 550025, Guizhou, China
10
Manaaki Whenua-Landcare Research, Auckland Mail Centre,
Private Bag 92170, Auckland 1142, New Zealand