Corresponding author: Georgios I. Zervakis (
Academic editor: M.P. Martín
Fryssouli V, Zervakis GI, Polemis E, Typas MA (2020) A global meta-analysis of ITS rDNA sequences from material belonging to the genus
The genus
Still, the species concept in
The internal transcribed spacer (
Although the
On the basis of the discrepancies and shortcomings noted before, the objectives of this study were: (i) to perform a thorough metadata analysis on the basis of a global dataset of
Dried specimens were obtained on loan from the
We studied 54 specimens in the form of either dried material or pure cultures. They represented well-established
Mycelia for DNA extraction were produced in static potato dextrose (Difco, USA) cultures. Following a 10–15 day incubation period at 25 °C, the mycelia were harvested by filtration and either directly processed for DNA extraction or stored at -20 °C. Mycelium or dried basidiome samples were pulverised by a micropestle in the presence of sterile sand and liquid nitrogen. Total genomic DNA was subsequently extracted through the silica Plant II DNA Extraction Miniprep System (Macherey and Nagel, Germany) by following the standard CTAB protocol provided by the manufacturer with minor modifications, i.e. the lysis step was extended to 1 h at 65 °C and the precipitation step to 1 h at room temperature, while the final elution step was performed at 65 °C for 1 h (
Sequences of the
Amplified fragments were examined by electrophoresis on 1% agarose gels. PCR products of the expected size were purified by microcentrifugation using the PureLink PCR purification kit (Invitrogen/Thermo Fisher Scientific, USA), according to manufacturer’s protocol. Purified amplicons were processed for bidirectional Sanger sequencing at CEMIA (University of Thessaly, Greece;
An initial dataset was compiled by retrieving and examining all publicly available
Summarised information on the
CLADES/Species (no. of sequences per taxon) | Sequences original labelling (no. of sequences per taxon) | Geographic origin of sequenced material | Host type | |
---|---|---|---|---|
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China |
|
100%/1.00 | |
|
|
|||
|
Canada, Germany, India, Pakistan, UK, USA | |||
|
Canada, Estonia, USA |
|
||
|
Belgium, Czech Republic, France, Greece, Poland, Slovakia, Slovenia |
|
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|
uncultured soil fungus (2), |
Estonia, UK | 93%/1.00 | |
|
Algeria, Argentina, Armenia, Belgium, Bulgaria, China, Czech Republic, Finland, France, Greece, India, Iran, Italy, Norway, Russia, Slovakia, Spain, South Korea, Sweden, Thailand, UK, USA, commercial | |||
|
China (Tibet), Nepal, Pakistan | 81%/1.00 | ||
|
|
|||
|
South Africa |
|
89%/- | |
|
China, Gabon, Myanmar | 99%/1.00 | ||
|
Philippines, Taiwan |
|
91%/0.99 | |
|
Australia, China, India |
|
||
|
Gabon, Cameroon |
|
-/0.98 | |
|
India |
|
100%/1.00 | |
|
Brazil, China, India |
|
||
|
Argentina, Brazil, Martinique, Mexico, USA |
|
||
|
Brazil, Colombia, Costa Rica, Cuba, French Guiana, Mexico, USA |
|
77%/- | |
|
China | 100%/1.00 | ||
|
Belgium, Bulgaria, China, Czech Republic, Egypt, France, Greece, India, Iran, Iraq, Italy, Netherlands, Poland, Slovakia, South Africa, South Korea, Tunisia, Turkey, UK |
|
||
|
Malaysia, Taiwan, commercial |
|
99%/1.00 | |
|
India, Turkey |
|
68%/- | |
|
Argentina, USA |
|
98%/1.00 | |
|
China, India, USA |
|
93%/1.00 | |
|
Argentina, China, India, Japan, Russia, South Korea, Taiwan, USA, commercial | 78%/1.00 | ||
|
|
|||
|
Brazil | |||
|
Brazil, Colombia, Cuba, Martinique, Mexico, Panama, USA |
|
100%/1.00 | |
|
Argentina | 99%/1.00 | ||
|
Ghana, India, Senegal |
|
100%/1.00 | |
China, Laos, Vietnam | 100%/1.00 | |||
|
China, Myanmar |
|
100%/1.00 | |
|
China, Indonesia, Malaysia, Thailand |
|
97%/1.00 | |
|
Bangladesh, China, India, Iran, Iraq, Japan, Laos, Malaysia, Myanmar, Nepal, South Korea, Thailand, commercial | 100%/1.00 | ||
|
Mexico, USA, commercial |
|
||
India, USA | 78%/1.00 | |||
|
Brazil, Colombia, Mexico |
|
99%/1.00 | |
|
Cameroon, South Africa |
|
||
|
India |
|
||
|
Australia, Indonesia |
|
89%/0.99 | |
|
Argentina, Brazil, Colombia, Cuba, Martinique, Mexico, USA |
|
93%/1.00 | |
|
China, India, Nepal, Pakistan, Philippines, Taiwan, Thailand | 74%/0.97 | ||
|
India |
|
100%/1.00 | |
|
China, India, Laos, Taiwan, Thailand |
|
||
|
Malaysia | 100%/1.00 | ||
|
|
|||
|
China, USA | 100%/1.00 | ||
|
Nepal | 100%/1.00 | ||
|
uncultured soil fungus (230), |
Antarctica, Armenia, Austria, Bulgaria, Canada, China, Czech Republic, Estonia, France, Germany, Greece, Hungary, India, Japan, Kyrgyzstan, Latvia, Lithuania, Netherlands, Poland, Russia, Slovakia, South Korea, Thailand, UK, USA, commercial | 99%/1.00 | |
|
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|||
|
|
|||
|
China, Laos, Myanmar, Taiwan |
|
99%/1.00 | |
|
|
|||
|
Cameroon |
|
82%/1.00 | |
|
Cameroon, South Africa | |||
|
Cameroon |
|
100%/1.00 | |
|
Ghana, Ivory Coast, South Africa, Thailand | |||
|
China, India, Sri Lanka | 71%/- | ||
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|
|
|||
|
Ghana, India, Senegal, Sri Lanka | 98%/1.00 | ||
|
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|
India, Laos, Malaysia, Vietnam |
|
98%/1.00 | |
|
China, Taiwan, Thailand |
|
100%/1.00 | |
|
|
|||
|
Cameroon, Gabon, Malaysia, South Africa, Tanzania |
|
99%/1.00 | |
|
Brazil | 89%/0.96 | ||
|
Brazil, Ecuador, French Guiana, India, Peru |
|
100%/0.96 | |
|
Australia, China, India, Indonesia, Laos, Malaysia, Myanmar, Taiwan, Thailand, Vietnam | 80%/1.00 | ||
|
|
|||
|
|
|||
|
Australia, China, India, Indonesia, Malaysia | |||
|
Gabon |
|
100%/1.00 | |
|
|
|||
|
USA | 100%/1.00 | ||
|
Colombia |
|
100%/1.00 | |
|
Cameroon, India | 100%/1.00 | ||
|
China, Indonesia, Japan, Malaysia, Taiwan, Thailand, Vietnam |
|
||
|
Indonesia |
|
98%/1.00 | |
|
|
|||
|
|
|||
|
China, Malaysia, Myanmar, Thailand |
|
99%/1.00 | |
|
|
|||
|
Brazil, Colombia, Costa Rica, Ecuador, French Guyana, Mexico, Peru, USA |
|
87%/0.99 | |
|
Argentina, Brazil, Colombia, Cuba, Puerto Rico, USA | 96%/1.00 | ||
|
India | 69%/1.00 | ||
|
South Africa |
|
92%/1.00 | |
|
China, Japan, Laos, Pakistan, South Korea, Taiwan, Thailand | |||
|
Australia, Cambodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Papua New Guinea, Sri Lanka, Thailand, USA, Vietnam | -/0.99 | ||
|
Australia, Indonesia | 96%/1.00 | ||
|
Indonesia, Malaysia | 100%/1.00 | ||
|
|
|||
|
South Africa |
|
100%/1.00 | |
|
China | 100%/1.00 | ||
|
||||
|
Czech Republic, Greece, Slovakia, UK |
|
93%/- | |
|
-/ |
|||
|
Ecuador | |||
|
Ecuador, Panama | 100%/1.00 | ||
|
Argentina |
|
100%/1.00 | |
|
China, India, Laos, New Zealand, Papua New Guinea, Taiwan, Thailand, Vietnam |
|
95%/1.00 | |
|
Argentina, Australia, Brazil, Chile, Costa Rica, India, New Zealand, South Africa, UK, USA | |||
|
|
|||
|
USA | 100%/1.00 | ||
|
China, Japan, Korea | 72%/- | ||
|
Armenia, Belgium, Croatia, France, Georgia, Germany, Greece, India, Iran, Italy, Slovakia, Spain, Tunisia, UK, USA | 89%/1.00 |
Abbreviations used for associated hosts:
The principal phylogenetic analysis of
Hence, multiple alignments of seven different matrices (Table
Phylogenetic analyses were based on Maximum Likelihood (
Datasets of
Datasets constructed and analysed | No. of sequences used/total | Represented entries/total entries | Alignment length | Constant characters* | Parsimony Informative characters* | No. of Rapid Bootstraps | Model substitution AICc TS1/ |
No. of generations | Split frequency | 50% credible trees | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|
440/2119 | 2027/3908 | 713 | 328 | 307 | 504 | -10429.843785 | 37885000 | 0.004398 | 28415 | Figs |
||
pDS1a: Cluster A.1 (expanded) | 120/124 | 297/301 | 571 | 500 | 25 | 600 | -1701.266373 | JC/JC/JC | 13710000 | 0.004400 | 10285 | Suppl. material |
pDS1b: Cluster A.2 (expanded) | 263/274 | 517/528 | 608 | 436 | 86 | 600 | -2929.177334 | TIM2ef+G/ |
63120000 | 0.004399 | 47341 | Suppl. material |
pDS1c: Cluster A.3 (expanded) | 341/641 | 694/1385 | 648 | 376 | 156 | 552 | -4501.116292 | 49780000 | 0.004400 | 37337 | Suppl. material |
|
pDS2/pDS3: Clades B & C (expanded) | 26/292 67/74 | 224/431 88/95 | 607 | 560/459 | 38/89 | 600 | -2652.570067 | K80/JC/JC | 7390000 | 0.004365 | 5545 | Suppl. material |
pDS4: Clade D (expanded) | 292/316 | 449/474 | 631 | 391 | 149 | 504 | -4424.884308 | TPM3uf+I+G/TPM2/TPM1uf+I+G | 40320000 | 0.004398 | 30241 | Suppl. material |
pDS5: Clade E (expanded) | 367/396 | 664/693 | 656 | 386 | 162 | 552 | -5123.351039 | TIM3+G/ |
65585000 | 0.004398 | 49190 | Suppl. material |
Elaboration of
Widely-adopted thresholds for separating amongst species in
Moreover, in order to provide additional information about the variation existing in the
In total, 3970
Almost half (45.3%) of all
Basidiomes of
For inferring the phylogeny of the entire genus, the main dataset (
A comprehensive evaluation of all
Summary of polymorphic regions in ITS1 and ITS2 spacers assessed in
Species/Groups | ITS1 sequence of potential diagnostic value | Length (nt) | Position in the alignment of Suppl. material |
ITS2 sequence of potential diagnostic value | Length (nt) | Position in the alignment of Suppl. material |
---|---|---|---|---|---|---|
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13 | 25–37 | GCCTTTGCGGGTW | 26 | 17–42 | |
|
TGTGAAGCGTGCT | 13 | 25/26–37/38 | TGYRGGCTTGGAC | 26 | 17–42 |
|
13 | 25–37 | AGCCTTGC | 8 | 16–23 | |
|
TGAAGCGYNCCYY | 13 | 27–39 | nd | ||
CGAAGCGTGC | 10 | 27–36 | nd | |||
|
||||||
|
CTTCAGTC | 8 | 16–23 | CTTGTGGGTT | 10 | 20–29 |
|
nd | nd | ||||
|
nd | nd | ||||
|
AACGTCGTKAAGCGGGC | 17 | 21–37 | nd | ||
GGGTCTTTT | 9 | 34–42 | CGTCTTTC | 8 | 60–67 | |
|
GCTCTTTACTGAGCC | 15 | 36–50 | CGGCCGGCTCCTCT | 21 | 65/67–85/87 |
|
15 | 36–50 | TAAATGC1 | 21 | 65/67–85/87 | |
|
AAGCGGCG | 8 | 55/56–62/63 | nd | ||
GGATCGGCGT | 10 | 55–64 | ACAGATCT | 8 | 13–20 | |
|
ACACCTAT | 8 | 84–91 | nd | ||
|
CCACAAACTCTR | 12 | 78–89 | CTTACAAA | 8 | 10–17 |
|
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|
GATTGTCG | 8 | 21–28 | CCATGCCC | 8 | 58/59–65/66 |
|
GGCATTAT | 8 | 21–28 | TTCTCTTA | 8 | 71/72–78/79 |
|
TTGCTGGG | 8 | 39–46 | CTTTTGTGGYTTT | 13 | 18–30 |
|
CAGATTGC | 8 | 19–26 | 10 | 54–63 | |
|
TGCGGAGCGCAT | 12 | 49–60 | CGGCCGTTAT | 10 | 54–63 |
|
GAGTGCAT | 8 | 53–60 | 10 | 54–63 | |
|
CCCTTTAT | 8 | 35–42 | nd | ||
9 | 22–30 | nd | ||||
|
ATCVTAAAA2 | 9 | 22–30 | CTCTTGGCC | 9 | 61–69 |
|
9 | 22–30 | CATTCTTG | 8 | 59–66 | |
|
9 | 22–30 | G(C)AAGCTTTTG | 10–11 | 13–22/23 | |
TCCCAGGA | 8 | 50–56 | CTCCTCTCTT | 10 | 72–81 | |
|
ACCGGGCTTTGCA | 13 | 42–54 | nd | ||
|
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|
GTGCTYTT | 8 | 32–39 | TAAGCTTKTGT | 11 | 14–24 |
|
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|
ATGGATCGCG | 10 | 18–27 | AGGTGTTTG | 9 | 47–55 |
CTTCTTGTC | 9 | 35–43 | TTGCAACC | 8 | 11–18 | |
|
GCTCTTGT | 8 | 34–41 | 8 | 11–18 | |
|
||||||
|
TTWCAGASSGT | 11 | 16–26 | AGGCTATT | 8 | 48–55 |
|
CGTTTTCA | 8 | 70–77 | TCTTTAATA | 9 | 60/62–68/70 |
|
GGAGCTSGT | 9 | 41–49 | GTAAAGGC | 8 | 24–31 |
|
nd | TTTTTARYGRKTTTGTAGG | 19 | 19–37 | ||
|
GTGTAAAA | 8 | 27–34 | ATGGCTWGT | 8 | 24/28/29–32/36/37 |
|
TCGCTCGC | 8 | 34–41 | TCTCTTCA | 8 | 3–10 |
|
TCGTGCGG | 8 | 23–30 | CTTTAACT | 8 | 61–68 |
|
GTTTGACRAGTT | 12 | 40/44–51/55 | ATCTCTTTGY | 10 | 16–25 |
GGCGTGGT | 8 | 24–31 | 10 | 16–25 | ||
|
||||||
|
CTTCAGGTC | 9 | 16–24 | CTTAATYGA | 9 | 21–29 |
GTTTTACG | 8 | 15–22 | ATRAGCTTCT | 10 | 13–22 | |
8 | 15–22 | TATGKGAG | 8 | 23–30 | ||
|
13 | 27–39 | 10 | 60–69 | ||
|
TGARRSGGGCTYG3 | 13 | 27–39 | TCCYTTTACR3 | 10 | 60–69 |
|
13 | 27–39 | 10 | 60–69 | ||
RTTAAACG | 8 | 26–33 | GTCGGACTW4 | 9 | 59–67 | |
|
GGCCCGTTT5 | 9 | 34/35–42/43 | GCCTTTGTC6 | 9 | 57–65 |
ACYGAGCYYGC | 11 | 41–51 | TCTTTGCGGGG | 11 | 19–29 | |
|
CGAAACGKGCTCG | 13 | 27–39 | 11 | 19–29 | |
CCCCATGA | 8 | 83/84–90/91 | GTCTTTACA | 9 | 59–67 | |
|
GGGCCCGTTC | 10 | 33–42 | CTTCTTGCGG | 10 | 18–27 |
|
AGGCCCGTTC | 10 | 33–42 | AGGTTTGTAGGG | 12 | 27/28–38/39 |
The
Summary tree of the genus
In all cases,
On the basis of
Detail from Fig.
Clade A includes 1927 entries distributed across 881 unique
Detail from Fig.
In the context of this work, Cluster A.1 corresponds to a well-supported clade (100%, 1.00; Fig.
The major group of Cluster A.1 corresponds to
Sequences deriving from two U.K. specimens (deposited as
In the past,
Noteworthy cases pertain to sequences under the names of “
In the frame of this study, four
Cluster A.2 (94%, 1.00; Fig.
A major subclade is formed by specimens collected in southeast Asia and Australia growing mostly on angiosperms (93%, 1.00 and 72%, 1.00, Fig.
A new phylogenetic species within Cluster A.2 is hereby proposed and is provisionally named “
Other four entries of dubious identity derive from material originating from Asia (China and India) labelled as
Another major subclade consisting of material originating from the Neotropics is formed by a total of 17 entries. Four of them are singletons, while the other 13 are grouped in six
Another well-supported terminal clade (100%, 1.00; Fig.
Another new phylogenetic species is hereby proposed, provisionally named “
A closely-related and well-supported (93%, 1.00; Fig.
A distinct phylogenetic group (98%, 1.00; Fig.
Another terminal group, although not adequately supported in any of the trees constructed, forms a sister clade to
Cluster A.3 (76%, 1.00; Fig.
Furthermore, one sequence (
Two sister subclades (0.99, Suppl. material
Two new well-supported monophyletic species, provisionally named as “
Three species, i.e.
Six other closely-related species are found in Cluster A.3 (i.e.
In this study,
A second group (A.3.2; 100%, 1.00; Fig.
A strongly-supported terminal subclade in Group A.3.2 (100%, 1.00; Fig.
Furthermore, three singletons from Malaysian material (in this particular case, geographic origin is inferred from the title of the study which appears on the respective GenBank records), initially identified as “
Clade B (96%, 1.00; Fig.
Detail from Fig.
Detail from Fig.
The other two species in Clade B form a well-supported sister clade (95%, 1.00; Fig.
Box plots of
Clade C is strongly supported (100%, 1.00; Fig.
Cluster C.2 is strongly supported (100%, 1.00; Fig.
The other group (100%, 1.00; Fig.
Clade D includes 291 sequences representing 449 individual entries separated into four clusters (Suppl. material
Cluster D.1 is placed at the base of Clade D and it is strongly supported (98%, 1.00; Fig.
Cluster D.2 comprises two distantly-related species, i.e.
Cluster D.3 is well supported (75%, 1.00; Fig.
Another terminal subclade is composed of nine entries (98%, 1.00; Fig.
Five sequences, representing
Finally, a well-represented and supported terminal subclade of Cluster D.3 (80%, 1.00; Fig.
Cluster D.4 is strongly supported (93%, 1.00; Fig.
Group D.4.1 (87%, 1.00; Fig.
Group D.4.2 (91%, 1.00; Fig.
The other two species comprise material from Asia only; the ‘core’ part corresponds to
On the basis of the results presented above, it is apparent that
Clade E is strongly supported (81%, 1.00; Fig.
Cluster E.1 corresponds to a single strongly-supported phylospecies (99%, 1.00; Fig.
Cluster E.2 (84%, 0.99; Fig.
A well-supported terminal subclade (87%, 0.99; Fig.
A strongly-supported (96%, 1.00; Fig.
Cluster E.2 also includes a large terminal subclade (65%, 1.00; Suppl. material
The other terminal clade (92%, 1.00; Fig.
The other two phylogenetic species appearing on the terminal subclade of Cluster E.2 (96%, 1.00; Fig.
Cluster E.3 consists of material corresponding to the laccate taxa
The close phylogenetic position of
Cluster E.4 (0.99; Fig.
A sister group to the former (1.00; Fig.
A sister terminal subclade to the previous group (
Finally, a distinct group included 80 entries deriving from specimens mainly from southeast Asia, South America and Oceania, as well as from central/north America and South Africa. That said, one sequence originated in the vicinity of the Kew Botanical Gardens, UK, such that we suspect it to have been imported with plant material. These sequences were mostly deposited as “
Cluster E.5 (97%, 1.00; Fig.
A sister subclade to the aforementioned clade corresponds to
In conclusion, the plasticity of morphological characters and substantial overlap of alleged diagnostic features is prevalent in the ‘dull’ taxa of this group; consequently, their taxonomic significance is dubious. Moreover, the massive, heavily-agglutinated matrix of the pileal crust in such non-laccate species often obstructs observation of discriminating features in pileal elements. The situation is further aggravated by the loss of type material of widely and commonly used species names (e.g.
As determined from the analysis performed in this study, the combined length of the two spacers (excluding the intercalary
Box plots of
In the ITS1 region (289 sites), 195 (67%) were variable and 159 (55%) were parsimony informative; the ITS2 region (262 sites) included 169 (65%) variable and 137 (52%) parsimony informative sites. This is in accordance with the outcome of previous reports indicating a larger variability for ITS1 in comparison to ITS2 in
The GC content was almost identical in ITS1 and ITS2 spacers when calculated for the entire genus (49.1 ± 1.5% and 49.2 ± 1.7%, respectively) (Fig.
The application of criteria of wide applicability/suitability for delineating taxa in the genus
Intraspecific
Especially as regards taxa of Cluster A.1, average interspecific distances (0.008 ± 0.004) are close to the respective intraspecific values (0.004 ± 0.003); similarly, sequence similarity in interspecific comparisons is high (99.0 ± 0.6; Fig.
On the other hand, species in Clade B present a clear barcoding gap and distinct ‘sequence diameters’ (as defined by
As regards Clades C, D and E, barcoding gaps are quite pronounced and delimitation of many species could be made on the basis of the 98% sequence similarity and the 0.015 genetic distance values previously mentioned and used for the establishment of the phylospecies proposed herein (Fig.
In conclusion,
Members of the genus
Three main lineages of the genus were identified: Clade A, Clade B and Clades C through E. Clades A and E include taxa with a cosmopolitan distribution, while species of Clade B are distributed across the Holarctic region; species of Clades C occur in the Paleotropics and taxa of Clade D exhibit a pantropical distribution. Their subsequent analysis results are consistent with the hypothesis of a Northern Hemisphere origin (tropical Asia) for
Indicative cases of
Palearctic – Eurasian, Old World:
East Asia – Malay Archipelago – Oceania:
East Asia and South Africa (Paleotropic): All taxa of Clade C, as well as
Holoarctic/Nearctic – Palearctic: Within Clade B,
East Asian – North American: Several biogeographic studies evidenced migration of fungi from east Asia to North America via the Bering Land Bridge route (Wu and Mueller 1997;
Neotropical:
Southern Hemisphere: The phylogenetic analysis of Southern Hemisphere species and complexes (species in Clusters A.2 and A.3, Clusters D.3 and D.4 and E.2 and E.4) indicated a restricted gene flow apparently due to geographic isolation, although episodic long-distance dispersal still occurs (
The majority of
As previously stated, one of the major obstacles for exploiting sequences present in public depositories is that many of them are inaccurate; errors in labelling of metadata were estimated to correspond to as much as 20% – or even 30% according to a recent report – of GenBank accessions including also recent deposits (
The study of a large dataset comprising almost four thousand
At a more general level, this study evidences that significant – yet largely untapped – mycological explanatory power resides in the public DNA sequence corpus and we hope that other mycologists will start scrutinising the sequence data available for their fungal groups of expertise. Our results also demonstrated that the so-called environmental sequences – usually ignored in a taxonomic/phylogenetic context – should be included in such pursuits (cf.
We are indebted to the curators of the fungaria of the Bulgarian Academy of Sciences (
This research was performed in the frame of a project (THALIS – UOA – MIS 377062) cofinanced by the European Union (European Social Fund – ESF) and Greek National Funds through the Operational Programme ‘Education and Lifelong Learning’ of the National Strategic Reference Framework (NSRF).
Tables S1–S6
species data
Figure S1
molecular data
Hypervariable regions of potential diagnostic value in ITS1 and ITS2 spacers for the
Figure S2a
molecular data
Phylogenetic reconstruction of the genus
Figure S2b
molecular data
Phylogenetic reconstruction of the genus
Figure S2c
molecular data
Phylogenetic reconstruction of the genus
Figure S2d
molecular data
Phylogenetic reconstruction of the genus
Figure S2e
molecular data
Phylogenetic reconstruction of the genus
Figure S2f
molecular data
Phylogenetic reconstruction of the genus