Fungal Diversity (2022) 117:1–272
https://doi.org/10.1007/s13225-022-00513-0
Fungal diversity notes 1512–1610: taxonomic and phylogenetic
contributions on genera and species of fungal taxa
Ruvishika S. Jayawardena1,2 · Kevin D. Hyde1,2,3 · Song Wang1 · Ya‑Ru Sun1,2,4 · Nakarin Suwannarach5,6 ·
Phongeun Sysouphanthong1,2,7 · Mohamed A. Abdel‑Wahab8 · Faten A. Abdel‑Aziz8 ·
Pranami D. Abeywickrama1,2,9 · Vanessa P. Abreu10 · Alireza Armand1,2 · André Aptroot11 · Dan‑Feng Bao1,12,13 ·
Dominik Begerow14 · Jean‑Michel Bellanger15 · Jadson D. P. Bezerra16 · Digvijayini Bundhun1,2 · Mark S. Calabon1,17 ·
Ting Cao18,19 · Taimy Cantillo20 · João L. V. R. Carvalho21 · Napalai Chaiwan1,2 · Che‑Chih Chen22 ·
Régis Courtecuisse23 · Bao‑Kai Cui24 · Ulrike Damm25 · Cvetomir M. Denchev26,27 · Teodor T. Denchev26,27 ·
Chun Y. Deng28 · Bandarupalli Devadatha29,46 · Nimali I. de Silva5,6 · Lidiane A. dos Santos30 · Nawal K. Dubey31 ·
Sylvain Dumez23 · Himashi S. Ferdinandez32 · André L. Firmino33 · Yusufjon Gafforov34,35 · Achala J. Gajanayake1,2 ·
Deecksha Gomdola1,2 · Sugantha Gunaseelan36 · Shucheng‑He1,2,37 · Zin H. Htet1,2 · Malarvizhi Kaliyaperumal36 ·
Martin Kemler14 · Kezhocuyi Kezo36 · Nuwan D. Kularathnage1,2,3,61 · Marco Leonardi38 · Ji‑Peng Li28 ·
Chunfang Liao1,2,3 · Shun Liu24 · Michael Loizides40 · Thatsanee Luangharn1 · Jian Ma1,2,41 · Hugo Madrid42 ·
S. Mahadevakumar43,62 · Sajeewa S. N. Maharachchikumbura44 · Dimuthu S. Manamgoda32 · María P. Martín45 ·
Niranjan Mekala46,47 · Pierre‑Arthur Moreau23 · Yan‑Hong Mu18,19 · Pasouvang Pahoua1 · Dhandevi Pem1,2 ·
Olinto L. Pereira10 · Wiphawanee Phonrob48 · Chayanard Phukhamsakda1,39 · Mubashar Raza49 ·
Guang‑Cong Ren1,2 · Andrea C. Rinaldi50 · Walter Rossi38 · Binu C. Samarakoon1,2 · Milan C. Samarakoon13 ·
Vemuri V. Sarma51 · Indunil C. Senanayake3,61 · Archana Singh31 · Maria F. Souza11 · Cristina M. Souza‑Motta21 ·
Adriano A. Spielmann11 · Wenxin Su39 · Xia Tang1,2,63 · XingGuo Tian1,2,41,52 · Kasun M. Thambugala53 ·
Naritsada Thongklang1,2 · Danushka S. Tennakoon5,6 · Nopparat Wannathes48 · DingPeng Wei1,13,37 ·
Stéphane Welti23 · Subodini N. Wijesinghe1,2 · Hongde Yang1,2,37 · Yunhui Yang1,2,3 · Hai‑Sheng Yuan18 ·
Huang Zhang37 · Jingyi Zhang1,2,41 · Abhaya Balasuriya1 · Chitrabhanu S. Bhunjun1,2 · Timur S. Bulgakov54 ·
Lei Cai49 · Erio Camporesi55,56,57 · Putarak Chomnunti2 · Y. S. Deepika58 · Mingkwan Doilom3 · Wei‑Jun Duan59,64 ·
Shi‑Ling Han49 · Naruemon Huanraluek1 · E. B. Gareth Jones60 · N. Lakshmidevi58 · Yu Li39 · Saisamorn Lumyong5,6 ·
Zong‑Long Luo41 · Surapong Khuna5,6 · Jaturong Kumla5,6 · Ishara S. Manawasinghe3 · Ausana Mapook1 ·
Wilawan Punyaboon1 · Saowaluck Tibpromma52 · Yong‑Zhong Lu41 · JiYe Yan9 · Yong Wang4
Received: 12 September 2022 / Accepted: 6 December 2022 / Published online: 23 February 2023
© The Author(s) under exclusive licence to Mushroom Research Foundation 2023
Abstract
This article is the 14th in the Fungal Diversity Notes series, wherein we report 98 taxa distributed in two phyla, seven classes,
26 orders and 50 families which are described and illustrated. Taxa in this study were collected from Australia, Brazil, Burkina
Faso, Chile, China, Cyprus, Egypt, France, French Guiana, India, Indonesia, Italy, Laos, Mexico, Russia, Sri Lanka, Thailand,
and Vietnam. There are 59 new taxa, 39 new hosts and new geographical distributions with one new combination. The 59 new
species comprise Angustimassarina kunmingense, Asterina lopi, Asterina brigadeirensis, Bartalinia bidenticola, Bartalinia
caryotae, Buellia pruinocalcarea, Coltricia insularis, Colletotrichum flexuosum, Colletotrichum thasutense, Coniochaeta
caraganae, Coniothyrium yuccicola, Dematipyriforma aquatic, Dematipyriforma globispora, Dematipyriforma nilotica,
Distoseptispora bambusicola, Fulvifomes jawadhuvensis, Fulvifomes malaiyanurensis, Fulvifomes thiruvannamalaiensis,
Fusarium purpurea, Gerronema atrovirens, Gerronema flavum, Gerronema keralense, Gerronema kuruvense, Grammothele
taiwanensis, Hongkongmyces changchunensis, Hypoxylon inaequale, Kirschsteiniothelia acutisporum, Kirschsteiniothelia
crustaceum, Kirschsteiniothelia extensum, Kirschsteiniothelia septemseptatum, Kirschsteiniothelia spatiosum, Lecanora
Handling Editor: Jian-Kui Liu.
Extended author information available on the last page of the article
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Fungal Diversity (2022) 117:1–272
immersocalcarea, Lepiota subthailandica, Lindgomyces guizhouensis, Marthe asmius pallidoaurantiacus, Marasmius
tangerinus, Neovaginatispora mangiferae, Pararamichloridium aquisubtropicum, Pestalotiopsis piraubensis, Phacidium
chinaum, Phaeoisaria goiasensis, Phaeoseptum thailandicum, Pleurothecium aquisubtropicum, Pseudocercospora vernoniae, Pyrenophora verruculosa, Rhachomyces cruralis, Rhachomyces hyperommae, Rhachomyces magrinii, Rhachomyces
platyprosophi, Rhizomarasmius cunninghamietorum, Skeletocutis cangshanensis, Skeletocutis subchrysella, Sporisorium
anadelphiae-leptocomae, Tetraploa dashaoensis, Tomentella exiguelata, Tomentella fuscoaraneosa, Tricholomopsis lechatii,
Vaginatispora flavispora and Wetmoreana blastidiocalcarea. The new combination is Torula sundara. The 39 new records
on hosts and geographical distribution comprise Apiospora guiyangensis, Aplosporella artocarpi, Ascochyta medicaginicola,
Astrocystis bambusicola, Athelia rolfsii, Bambusicola bambusae, Bipolaris luttrellii, Botryosphaeria dothidea, Chlorophyllum squamulosum, Colletotrichum aeschynomenes, Colletotrichum pandanicola, Coprinopsis cinerea, Corylicola italica,
Curvularia alcornii, Curvularia senegalensis, Diaporthe foeniculina, Diaporthe longicolla, Diaporthe phaseolorum, Diatrypella quercina, Fusarium brachygibbosum, Helicoma aquaticum, Lepiota metulispora, Lepiota pongduadensis, Lepiota
subvenenata, Melanconiella meridionalis, Monotosporella erecta, Nodulosphaeria digitalis, Palmiascoma gregariascomum,
Periconia byssoides, Periconia cortaderiae, Pleopunctum ellipsoideum, Psilocybe keralensis, Scedosporium apiospermum,
Scedosporium dehoogii, Scedosporium marina, Spegazzinia deightonii, Torula fici, Wiesneriomyces laurinus and Xylaria
venosula. All these taxa are supported by morphological and multigene phylogenetic analyses. This article allows the
researchers to publish fungal collections which areimportant for future studies. An updated, accurate and timely report of
fungus-host and fungus-geography is important. We also provide an updated list of fungal taxa published in the previous
fungal diversity notes. In this list, erroneous taxa and synonyms are marked and corrected accordingly.
Keywords 59 New taxa · 39 New records · Ascomycota · Basidiomycota · Dothideomycetes · Lecanoromycetes ·
Leotiomycetes · Sordariomycetes · Ustilaginomycotina
Table of contents
Phylum Ascomycota R.H. Whittaker
Subphylum Pezizomycotina O.E. Erikss. & Winka
Class Dothideomycetes O.E. Erikss. & Winka
Subclass Dothideomycetidae P.M. Kirk et al.
Mycosphaerellales (Nannf.) P.F. Cannon
Mycosphaerellaceae Lindau
1512. Pseudocercospora vernoniae Archana Singh & N.K.
Dubey, sp. nov. (contributed by A. Singh and N.K. Dubey)
Subclass Pleosporomycetidae C.L. Schoch et al.
Kirschsteiniotheliales Hern.-Restr. et al.
Kirschsteiniotheliaceae Boonmee & K.D. Hyde
1513. Kirschsteiniothelia acutisporum S. Wang, Q. Zhao &
K.D. Hyde, sp. nov. (contributed by S. Wang and Y.R. Sun)
1514. Kirschsteiniothelia crustaceum S. Wang, Q. Zhao &
K.D. Hyde, sp. nov. (contributed by S. Wang and Y.R. Sun)
1515. Kirschsteiniothelia extensum. S. Wang, Q. Zhao &
K.D. Hyde, sp. nov. (contributed by S. Wang, K.D. Hyde
and Y.R. Sun)
1516. Kirschsteiniothelia septemseptatum S. Wang, Q.
Zhao & K.D. Hyde, sp. nov. (contributed by S. Wang and
Y.R. Sun)
13
1517. Kirschsteiniothelia spatiosum S. Wang, Q. Zhao &
K.D. Hyde, sp. nov. (contributed by S. Wang and Y.R. Sun)
Pleosporales Luttrell ex M.E. Barr
Amorosiaceae Thambug. & K.D. Hyde
1518. Angustimassarina kunmingense H.D. Yang & K.D.
Hyde, sp. nov. (contributed by H. Yang, K.D. Hyde and C.
Bhunjun)
Bambusicolaceae D.Q. Dai & K.D. Hyde
1519. Bambusicola bambusae D.Q. Dai & K.D. Hyde, new
host/substrate record from Thailand (contributed by D.F.
Bao and K.D. Hyde)
1520. Corylicola italica Wijesinghe, Camporesi, Yong
Wang bis & K.D. Hyde, new host record from Italy (contributed by P. Pahoua and E. Camporesi)
1521. Palmiascoma gregariascomum Phookamsak & K.D.
Hyde, new host record from Thailand (contributed by M.C.
Samarakoon)
Coniothyriaceae W.B. Cooke
1522. Coniothyrium yuccicola Chaiwan, Jayaward., Bulgakov & K.D. Hyde, sp. nov. (Contributed by N. Chaiwan
and T.S. Bulgakov)
Fungal Diversity (2022) 117:1–272
Didymellaceae Gruyter, Aveskamp & Verkley
1523. Ascochyta medicaginicola Q. Chen & L. Cai, new
host record from Russia (contributed by N. Chaiwan and
T.S. Bulgakov)
Didymosphaeriaceae Munk
1524. Spegazzinia deightonii (S. Hughes) Subram., new
host record from Thailand (contributed by D. Bundhun and
B.C. Samarakoon)
Lindgomycetaceae K. Hiray. et al.
1525. Hongkongmyces changchunensis Phukhams., W.X.
Su, & Y. Li, sp. nov. (contributed by C. Phukhamsakda, K.D.
Hyde and W.X. Su)
1526. Lindgomyces guizhouensis J. Ma, Y.Z. Lu & K.D.
Hyde, sp. nov. (contributed by J. Ma, J. Y. Zhang and Y.Z.
Lu)
Lophiostomataceae Sacc
1527. Neovaginatispora mangiferae Tennakoon, M.S. Calabon, E.B.G. Jones, K.D. Hyde, sp. nov. (contributed by M.
S. Calabon, K.D. Hyde and D.S Tennakoon)
1528. Vaginatispora flavispora M.S. Calabon, E.B.G. Jones,
K.D. Hyde, sp. nov. (contributed by M. S. Calabon, K.D.
Hyde and E.B.G. Jones)
Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde
1529. Phaeoseptum thailandicum Samarak. & K.D. Hyde,
sp. nov. (contributed by M. C. Samarakoon and K.D. Hyde)
1530. Pleopunctum ellipsoideum N.G. Liu, K.D. Hyde &
J.K. Liu, new host record from Thailand (contributed by
Y.R. Sun)
Phaeosphaeriaceae M.E. Barr
1531. Nodulosphaeria digitalis W.J. Li, Camporesi, Bhat
& K.D. Hyde, new host record from Italy (contributed by
D. Bundhun)
Pleosporaceae Nitschke
1532. Bipolaris luttrellii Alcorn, new host record from
China (contributed by K.M. Thambugala and R.S.
Jayawardena)
1533. Curvularia alcornii Manamgoda, L. Cai & K. D.
Hyde, new host record Sri Lanka (contributed by H.S. Ferdinandez and D.S. Manamgoda)
1534. Curvularia senegalensis (Speg.) Subram., new host
record from Sri Lanka (contributed by H.S. Ferdinandez
and D.S. Manamgoda)
1535. Pyrenophora verruculosa Madrid & Cantillo, sp. nov.
(contributed by H. Madrid and T. Cantillo)
3
Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray
1536. Tetraploa dashaoensis C.F. Liao & Doilom, sp. nov.
(contributed by C.F. Liao, K.D. Hyde and M. Doilom)
Torulaceae Corda
1537. Torula fici Crous, new host record from Taiwan
and Thailand (contributed by B.C. Samarakoon, D.S. Tennakoon and P. Chomnunti)
1538. Torula sundara (Subram.) Y.R. Sun, Yong Wang
bis & K.D. Hyde, comb. nov. (contributed by Y.R. Sun)
Periconiaceae (Sacc.) Nann.,
1539. Periconia byssoides Pers., new host and geographical record from India (contributed by S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and S. S. N.
Maharachchikumbura)
1540. Periconia cortaderiae Thambugala & K.D. Hyde,
new host and geographical record from Russia (contributed by B.C. Samarakoon and T.S. Bulgakov)
Tubeufiales Boonmee & K.D. Hyde
Tubeufiaceae M.E. Barr
1541. Helicoma aquaticum Y.Z. Lu, J.C. Kang & K.D.
Hyde, new host record from Thailand (contributed by
X. Tang)
Wiesneriomycetaceae Suetrong, Rungjind., Somrith. &
E.B.G. Jones
1542. Wiesneriomyces laurinus (Tassi) P.M. Kirk, new
host record from China (contributed by Y. Yang and I.S.
Manawasinghe)
Dothideomycetes orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss
Asterinaceae Hansf
1543. Asterina brigadeirensis A.L. Firmino & O.L.
Pereira, sp. nov. (contributed by O. L. Pereira and A. L.
Firmino)
1544. Asterina lopi A.L. Firmino & O.L. Pereira, sp. nov.
(contributed by O. L. Pereira and A. L. Firmino)
Botryosphaeriales C.L. Schoch, Crous & Shoemaker
Aplosporellaceae Slippers, Boissin & Crous
1545. Aplosporella artocarpi Trakun., L. Lombard &
Crous, new host record from Thailand (contributed by Z.
H. Htet and A. Mapook)
Botryosphaeriaceae Theiss. & Syd. [as 'Botryosphaeriacae']
1546. Botryosphaeria dothidea (Moug.) Ces. & De Not.,
new geographical and habitat record from China (contributed by H. Yang and R.S. Jayawardena)
13
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Class Laboulbeniomycetes Engler
Laboulbeniales Lindau
Laboulbeniaceae G. Winter
1547. Rhachomyces cruralis W. Rossi & M. Leonardi, sp.
nov. (contributed by W. Rossi & M. Leonardi)
1548. Rhachomyces hyperommae W. Rossi & M. Leonardi,
sp. nov. (contributed by W. Rossi & M. Leonardi)
1549. Rhachomyces magrinii W. Rossi & M. Leonardi, sp.
nov. (contributed by W. Rossi & M. Leonardi)
1550. Rhachomyces platyprosophi W. Rossi & M. Leonardi,
sp. nov. (contributed by W. Rossi & M. Leonardi)
Class Lecanoromycetes O.E. Erikss. & Winka
Subclass Lecanoromycetidae P.M. Kirk et al.
Caliciales Bessey
Caliciaceae Chevall
1551. Buellia pruinocalcarea Aptroot, M.F. Souza &
Spielmann, sp. nov. (contributed by Aptroot, Souza and
Spielmann)
Lecanorales Nannf
Lecanoraceae Körb
1552. Lecanora immersocalcarea Aptroot, M.F. Souza &
Spielmann, sp. nov. (contributed by Aptroot, Souza and
Spielmann)
Teloschistales D. Hawksw. & O.E. Erikss
Teloschistaceae Zahlbr
1553. Wetmoreana blastidiocalcarea Aptroot, M.F. Souza
& Spielmann, sp. nov. (contributed by Aptroot, Souza and
Spielmann)
Class Leotiomycetes O.E. Erikss. & Winka
Phacidiales C.E. Bessey
Phacidiaceae Fr
1554. Phacidium chinense G.C. Ren & K.D. Hyde, sp. nov.
(contributed by G.C. Ren and K.D. Hyde)
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan., Maharachch. &
K.D. Hyde
Diaporthaceae Höhn. ex Wehm
1555. Diaporthe foeniculina (Sacc.) Udayanga & Castl.,
new host record from Italy (contributed by P. D. Abeywickrama and E. Camporesi)
1556. Diaporthe longicolla (Hobbs) J.M. Santos, Vrandečić
& A.J.L. Phillips, new host record from India (contributed
by S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and
S. S. N. Maharachchikumbura)
1557. Diaporthe phaseolorum (Cooke & Ellis)
Sacc., new host record from India (contributed by S.
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Fungal Diversity (2022) 117:1–272
Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and S. S.
N. Maharachchikumbura)
Melanconiellaceae Senan., Maharachch. & K.D. Hyde
1558. Melanconiella meridionalisVoglmayr & Jaklitsch,
new host and geographical record from Italy (contributed
by N. I. de Silva and E. Camporesi)
Pararamichloridiales Crous
Pararamichloridiaceae Crous
1559. Pararamichloridium aquisubtropicum J.Y. Zhang,
Y.Z. Lu & K.D. Hyde, sp. nov. (contributed by J.Y. Zhang,
J.Ma, Y.Z. Lu, and K.D. Hyde)
Distoseptisporales Z.L. Luo, K.D. Hyde & H.Y. Su
Distoseptisporaceae K.D. Hyde & McKenzie
1560. Distoseptispora bambusicola X. Tang, Jayaward., J.C
Kang & K.D. Hyde sp. nov. (contributed by X. Tang).
Glomerellales Chadef. ex Réblová et al
Glomerellaceae Locq. ex Seifert & W. Gams
1561. Colletotrichum aeschynomenes B.S. Weir & P.R.
Johnst., new host record from Thailand (contributed by D.
Gomdola and R.S. Jayawardena)
1562. Colletotrichum flexuosum Damm, sp. nov. (contributed by U. Damm)
1563. Colletotrichum pandanicola Tibpromma & K.D.
Hyde, new host records from India and Thailand, geographical record from India (contributed by S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi, S. S. N. Maharachchikumbura and R.S. Jayawardena)
1564. Colletotrichum thasutense Armand, K.D. Hyde,
Jayaward., sp. nov. (contributed by A. Armand and R.S.
Jayawardena)
Hypocreales Lindau
Nectriaceae Tul. & C. Tul
1565. Fusarium brachygibbosum Padwick, new host record
from India (contributed by S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and S.S.N. Maharahchikumbura)
1566. Fusarium purpurea S.L. Han, M. Raza, W.J. Duan
& L. Cai, sp. nov. (contributed by S.L. Han and M. Raza)
Microascales Luttr
Microascaceae Luttr. ex Malloch
1567. Scedosporium apiospermum Sacc. ex Castell. &
Chalm., a new host record from Thailand (contributed by
A. J. Gajanayake)
1568. Scedosporium dehoogii Gilgado, new record from
India (contributed by Devadatha and Sarma)
1569. Scedosporium marina Devadatha & V.V Sarma, sp.
nov. (contributed by Devadatha and Sarma)
Fungal Diversity (2022) 117:1–272
Hypocreomycetidae incertae sedis (Rhexoacrodictys and
Dematipyriforma clade)
1570. Dematipyriforma aquatica Abdel-Aziz &AbdelWahab, sp. nov. (contributed by Abdel-Aziz and
Abdel-Wahab)
1571. Dematipyriforma globispora Abdel-Aziz &AbdelWahab, sp. nov. (contributed by Abdel-Aziz and
Abdel-Wahab)
1572. Dematipyriforma nilotica Abdel-Aziz &AbdelWahab, sp. nov. (contributed by Abdel-Aziz and
Abdel-Wahab)
Subclass Savoryellomycetidae Hongsanan, K.D. Hyde &
Maharachch
Coniochaetales Huhndorf, A.N. Mill. & F.A. Fernández
Coniochaetaceae Malloch and Cain
1573. Coniochaeta caraganae D. Pem, Bulgakov & K.D.
Hyde, sp. nov. (Contributed by D. Pem, T.S. Bulgakov and
M. Raza)
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
1574. Rhexoacrodictys erecta (Ellis & Everh.) W.A. Baker
& Morgan-Jones, in Baker, Partridge & Morgan-Jones,
Mycotaxon 82: 99 (2002) new host record from Thailand
(contributed by X.G. Tian and S. Tibpromma)
1575. Phaeoisaria goiasensis H.M. Silva, A.D. Cavalcanti
& J.D.P. Bezerra, sp. nov. (contributed by H.M. Silva, A.D.
Cavalcanti and J.D.P. Bezerra)
1576. Pleurothecium aquisubtropicum J. Ma, Y.Z. Lu &
K.D. Hyde, sp. nov (contributed by J. Ma, J.Y. Zhang and
Y.Z. Lu)
Subclass Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D Hawksw & OE Erikss
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor &
M.E. Barr
1577. Apiospora guiyangensis Samarak., Jian K. Liu &
K.D. Hyde, new host record from China (contributed by
D.P. Wei)
Sporocadaceae Corda
1578. Bartalinia bidenticola Htet, Mapook & K.D. Hyde,
sp.nov (contributed by Z. H. Htet, K.D. Hyde and A.
Mapook)
1579. Bartalinia caryotae Senan., Kular. & K.D. Hyde,
sp. nov. (contributed by I.C. Senanayake and N. D.
Kularathnage)
1580. Pestalotiopsis piraubensis V.P. Abreu & O.L. Pereira,
sp. nov. (contributed by V.P. Abreu and O.L. Pereira)
5
Xylariales Nannf
Diatrypaceae Nitschke
1581. Diatrypella quercina (Pers.) Cooke, new host record
from Russia (contributed by S. N. Wijesinghe and T.S.
Bulgakov)
Hypoxylaceae DC
1582. Hypoxylon inaequale S.C. He & Jayaward., sp. nov
(contributed by S.C. He)
Xylariaceae Tul. & C. Tul
1583. Astrocystis bambusicola R.H. Perera & K.D. Hyde,
new host record from China (contributed by D.P. Wei)
1584. Xylaria venosula Speg., new geographical record
from India (contributed by M. Niranjan and V. V. Sarma)
Phylum Basidiomycota R.T. Moore
Subphylum Agaricomycotina Doweld
Class Agaricomycetes Doweld
Agaricales Underw
Agaricaceae Chevall
1585. Chlorophyllum squamulosum A.K. Dutta, Soumili
Bera & K. Acharya, new record from Thailand (contributed
by J. Kumla and N. Suwannarach)
1586. Lepiota metulispora (Berk. & Broome) Sacc., new
record from Laos (contributed by P. Sysouphanthong and
N. Thongklang)
1587. Lepiota pongduadensis Sysou., new record from Laos
(contributed by P. Sysouphanthong and N. Thongklang)
1588. Lepiota subthailandica Sysouph., K.D. Hyde &
Thongkl., sp. nov (contributed by P. Sysouphanthong and
N. Thongklang)
1589. Lepiota subvenenata Hai J. Li, Y.Z. Zhang & C.Y.
Sun, new record from Laos (contributed by P. Sysouphanthong and N. Thongklang)
Atheliales Jülich
Atheliaceae Jülich
1590. Athelia rolfsii (Curzi) C.C. Tu & Kimbr., new record
from India (contributed by S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and S. S. N. Maharachchikumbura)
Hymenochaetales Oberw
Hymenochaetaceae Donk
1591. Coltricia insularis P.-A. Moreau, Bellanger, Loizides
& A. Rinaldi, sp. nov. (contributed by P.-A. Moreau, Bellanger, Loizides and A. Rinaldi)
1592. Fulvifomes jawadhuvensis Kezo, K., Gunaseelan, S.,
& Kaliyaperumal, M., sp. nov. (contributed by K. Kezo, S.
Gunaseelan, M. Kaliyaperumal and T. Luangharn)
13
6
1593. Fulvifomes malaiyanurensis Gunaseelan, S., Kezo,
K. & Kaliyaperumal, M., sp. nov. (contributed by contributed by K. Kezo, S. Gunaseelan, M. Kaliyaperumal and T.
Luangharn)
1594. Fulvifomes thiruvannamalaiensis Gunaseelan, S.,
Kezo, K. and Kaliyaperumal, M., sp. nov. (contributed by
contributed by K. Kezo, S. Gunaseelan, M. Kaliyaperumal
and T. Luangharn)
Hymenogastraceae Vittad
1595. Psilocybe keralensis K.A. Thomas, Manim. &
Guzmán, new record from Thailand (contributed by N.
Suwannarach and J. Kumla)
Marasmiaceae Roze ex Kühner
1596. Marasmius pallidoaurantiacus Wannathes, N.
Suwannarach, J. Kumla & S. Lumyong, sp. nov. (contributed by N. Wannathes, N. Suwannarach, J. Kumla and S.
Lumyong)
1597. Marasmius tangerinus Wannathes, N. Suwannarach,
J. Kumla & Lumyong, sp. nov. (contributed by N. Wannathes, N. Suwannarach, J. Kumla and S. Lumyong)
Physalacriaceae Corner
1598. Rhizomarasmius cunninghamietorum Chun Y.
Deng, J.P. Li & Gafforov, sp. nov. (contributed by Chun
Y. Deng, J.P. Li and Y. Gafforov)
Fungal Diversity (2022) 117:1–272
Agaricales genera incertae sedis
1605. Gerronema atrovirens Wannathes, N. Suwannarach,
J. Kumla, Phonrob & S. Lumyong, sp. nov. (contributed by
N Wannathes, N Suwannarach J Kumla and S Lumyong)
1606. Gerronema flavum Wannathes, N. Suwannarach, J.
Kumla, Phonrob & S. Lumyong, sp. nov. (contributed by N
Wannathes, N Suwannarach J Kumla and S Lumyong)
1607. Gerronema keralense K. P. D. Latha & Manim, new
record from Thailand (contributed by N Wannathes, N
Suwannarach J Kumla, S Khuna, W Phonrob and S Tabtan)
1608. Gerronema kuruvense K. P. D. Latha & Manim, new
record from Thailand (contributed by N Wannathes, N
Suwannarach J Kumla, S Khuna, W Phonrob and S Tabtan)
1609. Tricholomopsis lechatii Courtec., S. Dumez, S.
Welti & P.-A. Moreau, sp. nov. (contributed by Courtec., S.
Dumez, S. Welti and P.-A. Moreau)
Subphylum Ustilaginomycotina Doweld
Class Ustilaginomycetes R. Bauer et al.
Ustilaginales G. Winter
Ustilaginaceae Tul & C. Tul
1610. Sporisorium anadelphiae-leptocomae T. Denchev,
Denchev, Kemler, M.P. Martín & Begerow, sp. nov. (contributed by T. Denchev, Denchev, Kemler, M.P. Martín and
Begerow)
Introduction
Polyporales Gäum
Polyporaceae Fr. ex Corda
1599. Grammothele taiwanensis C.C. Chen, sp. nov. (contributed by C.C. Chen)
Incrustoporiaceae Jülich
1600. Skeletocutis cangshanensis B.K. Cui & Shun Liu,
sp. nov. (contributed by B.K. Cui and Shun Liu)
1601. Skeletocutis subchrysella B.K. Cui & Shun Liu, sp.
nov. (contributed by B.K. Cui and Shun Liu)
Psathyrellaceae Vilgalys, Moncalvo & Redhead,
1602. Coprinopsis cinerea (Schaeff.) Redhead, Vilgalys
& Moncalvo, new record from India (contributed by
S. Mahadevakumar, Y.S. Deepika, N. Lakshmidevi and
S.S.N. Maharachchikumbura)
Thelephorales Corner ex Oberw
Thelephoraceae Chevall
1603. Tomentella exiguelata Y.H. Mu & H.S. Yuan, sp.
nov. (contributed Y.H. Mu, T. Cao and H.S. Yuan)
1604. Tomentella fuscoaraneosa Y.H. Mu & H.S. Yuan,
sp. nov. (contributed Y.H. Mu, T. Cao and H.S. Yuan).
13
Fungi play a key role in many biological processes, influencing ecosystems (Schimann et al. 2017). They are saprobes,
epiphytes, endophytes, animal and plant pathogens or symbionts (Chethana et al. 2021a, b). High species diversity in
fungi exhibits a huge variation in morphology, lifestyles and
the mode of dispersal (Hyde et al. 2018). Fungi are also
important in biotechnological applications (Hyde et al.
2019).
The current estimate of fungal diversity is highly uncertain, ranging from 1.5 to 12 million species (Wu et al. 2019;
Hyde et al. 2021; Bhunjun et al. 2022). Of this massive
number, only around 150,000 species have been named and
classified to date. With the introduction of DNA-based techniques in species delimitation, the newly described taxa per
year have dramatically increased. Whether these newly introduced taxa are novel is another challenge the mycologists
face. With only 10% of fungi being named and classified,
many species remain to be discovered (Hyde et al. 2021).
Some species are poorly described and lack molecular data.
This can be overcome if we collect, isolate, sequence and
provide new data on fungi from different hosts and habitats.
Identification of new taxa, recollection of already known
Fungal Diversity (2022) 117:1–272
taxa, the establishment of reference specimens and epitypification or neo-typification of taxa with fresh material
and cultures are necessary as they contribute to providing a
stable taxonomy for fungi Chethana et al. (2021a) as well as
for carrying out assays to identify any potential compounds
that can be harnessed at the industrial level. Identification
and documentation of the host and the geographical range
of a fungus can be particularly important in disease management (Dugan et al. 2009).
In order to provide an outlet for the mycologists to
publish their findings in mycology, different publication
series such as AJOM new records and collections of fungi
(Hyde et al. 2019; Chethana et al. 2021b), Fungal Diversity notes (Liu et al. 2015; Ariyawansa et al. 2015; Hyde
et al. 2017, 2019, 2020; Tibpromma et al. 2018; Wanasinghe et al. 2018; Phookamsak et al. 2019; Boonmee et al.
2021), Fungal planet (Crous et al. 2015a, b, c, 2017, 2018)
and Mycosphere notes (Thambugala et al. 2015; Hyde et al.
2018, 2021; Jayawardena et al. 2018; Manawasinghe et al.
2022), are now available. As a result, numerous new taxa,
geographical and host records, new combinations, and reference data were introduced along with morphological and
multigene analyses.
This is the 14th in the series of Fungal Diversity Notes
with entries mainly collected from Australia, Brazil, Burkina Faso, Chile, China, Cyprus, Egypt, France, French
Guiana, India, Indonesia, Italy, Laos, Mexico, Russia, Sri
Lanka, Thailand, and Vietnam. We aim to provide new data
including morphological, geographical and sequence data
for a stable taxonomy and phylogeny, which become significantly important for the accurate identification of fungi
as suggested by Cao et al. (2021), Chethana et al. (2021a),
Manawasinghe et al. (2019), Maharachchikumbura et al.
(2021), Jayawardena et al. (2021b) and Pem et al. (2021).
We provide a detailed description and an updated tree for the
genus or family of each entry. The ‘notes’ under each entry
discuss how the new taxa are established, including the host
and geographical ranges. The data compiled in this study can
be used by future researchers for a better understanding of
the taxonomy of each different group of fungi.
7
Indonesia, Italy, Laos, Mexico, Russia, Sri Lanka, Thailand,
and Vietnam. Taxa were described and illustrated based on
morphological features, coupled with phylogenetic analyses
performed by maximum likelihood (ML), maximum parsimony (MP) and Bayesian posterior probability (BYPP) criteria. Colour codes followed the Methuen Handbook of Colour
(Kornerup and Wanscher 1978). Phylogenetic analyses were
performed based on details outlined by Dissanayake et al.
(2020). Details of each analysis are given in Supplementary
Table 1. The pairwise homoplasy index (PHI) test was carried
out when necessary, using Split Trees as described by Quaedvlieg et al. (2014) to determine the recombination level within
phylogenetically closely related species. The new taxa are justified based on the guidelines of Cao et al. (2021), Chethana
et al. (2021a, b), Manawasinghe et al. (2021), Maharachchikumbura et al. (2021), Jayawardena et al. (2021b) and Pem
et al. (2021).
Results
Ascomycota R.H. Whittaker
Notes: We follow the latest treatments and updated accounts
of Ascomycota in Wijayawardene et al. (2020, 2022).
Subphylum Pezizomycotina O.E. Erikss. & Winka
Class Dothideomycetes O.E. Erikss. & Winka
Notes: We follow the latest treatments and updated accounts
of Dothideomycetes in Hongsanan et al. (2020a, b) and
Wijayawardene et al. (2020, 2022).
Subclass Dothideomycetidae P.M. Kirk, P.F. Cannon,
J.C. David & Stalpers ex C.L. Schoch, Spatafora, Crous &
Shoemaker
Materials and methods
Mycosphaerellales (Nannf.) P.F. Cannon
Notes: Abollahzadeh et al. (2020) based on LSU, tef1 and
rpb2 sequence data revalidated Mycosphaerellales as a
separate order. Mycosphaerellales include species that are
saprobes, ectophytes, plant pathogens and lichenised fungi.
This order includes eight families viz. Cystocoleaceae, Dissoconiaceae, Extremaceae, Mycosphaerellaceae, Neodevriesiaceae, Phaeothecoidiellaceae, Schizothyriaceae and Teratosphaeriaceae (see Abdollahzadeh et al. 2020).
Materials and methods follow the previous fungal diversity
notes (Hyde et al. 2016, 2020a, b, c; Tibpromma et al. 2017;
Wanasinghe et al. 2018; Phookamsak et al. 2019; Boonmee et al. 2021 and Senanayake et al. 2020). When specific
details are available for material and methods they are given
in the ‘notes’ section of each taxon. Taxa described in this
study were collected from Australia, Brazil, Burkina Faso,
Chile, China, Cyprus, Egypt, France, French Guiana, India,
Mycosphaerellaceae Lindau, Nat. Pflanzenfamilien:
421(1897)
Notes: Mycosphaerellaceae was established by Lindau
(1896) with Mycosphaerella as the type genus. This is one
of the largest families including asexual morphs, asexual
holomorphs or species with mycosphaerella-like sexual
morphs. The majority of them are parasitic or saprobic on
plants, fungi and lichens (Hyde et al. 2013). Wijayawardene
13
8
Fungal Diversity (2022) 117:1–272
et al. (2022) accepted a total of 119 genera having molecular
data under Mycosphaerellaceae.
wide range of climatic conditions including cool temperate,
sub-tropical and tropical regions (Crous et al. 2014).
Pseudocercospora Speg., Anales del Museo Nacional de
Historia Natural Buenos Aires 20 (13): 438 (1910)
Notes: Pseudocercospora was established by Spegazzini
(1910) with P. vitis as the type genus. The genus is characterized by conidiophores solitary, fasciculate, synnematal
or arranged in sporodochia, conidia coloured, scars unthickened or slightly thickened (Crous and Braun 2003; Crous
et al. 2014). They are mostly plant pathogenic fungi associated with leaf and fruit spots and are widely distributed in a
Pseudocercospora vernoniae Archana Singh & N.K. Dubey,
sp. nov
Mycobank Number: MB 834618, Facesoffungi number: FoF
07979, Figs. 1, 2
Etymology: Based to the host genus from which the taxon
was isolated
Holotype: AMH:10043
Asexual morph: on leaf spots of Vernonia cineria, hypophyllous later amphiphyllous, 2–5 mm, angular, vein limited,
Fig. 1 Pseudocercospora vernoniae (AMH: 10043, Holotype)
a Habit of infected plant Vernonia cineria. b Symptoms on the
upper leaf surface. c Symptoms
on the lower leaf surface. d Culture after 3 weeks. e–o Conidia.
k Conjugating conidia. p–q
Fascicle of conidiophores. Scale
bars: e–j = 20 µm, k–o = 10 µm
13
Fungal Diversity (2022) 117:1–272
9
Fig. 2 Scanning Electron
Micrographs of Pseudocercospora vernoniae. a Fasciles
of conidiophores arising from
stomata of the host plant.
b Single fascicle of conidiophores. c Conidia attached to
conidiophores. d Conidiogenous
cells with scars. Scale bars:
a–c = 10 µm, d = 2 µm
discrete and later forming irregular larger patches, grayish
brown on lower surface and dark blackish- brown on upper
surface. Caespituli hypophyllous later amphiphyllous, dark
brown, erumpent. Stromata substomatal, few cells to welldeveloped, made up of oval to round 3–5 µm wide pseudoparenchymatous cells, median to dark brown. Conidiophores
fasciculate, unbranched or rarely branched, geniculate,
1–8-septate, light brown 13.6–40.3 (50) × 3.5–5.5 µm. Conidiogenous cells integrated, polyblastic, cicaterised. Conidia
septate (1–7), catenate in branched chains, straight to curved,
cylindrical, constricted at septa, olivaceous brown, subcylindrical, base obclavate to obconico truncate, tip subacute
to obtuse 21.7–44.8 (92) × 4.5–5.5 µm. Sexual morph: Not
observed.
Culture characteristics: Conidia germinating on Potato
Dextrose Agar (PDA). Colonies very slow growing, velvety,
greyish brown; reaching 2–5 mm diam., in 28 days at 27
°C, margin circular to irregular, reverse blackish brown raising centrally, of dense cottony mycelium and hard texture.
Mycelium smooth, branched, asexual and sexual spores not
formed within 60 days.
Material examined: India, Sonebhadra U.P., on living
leaves of Vernonia cineria (L.) Less (Asteraceae), Dec 2017,
AMH: 10043 (Holotype), culture ex type NFCCI: 4441.
GenBank numbers: MN691042 (LSU); MN691041 (ITS);
MT106617 (act); MT106618 (tef1)
Notes: Pseudocercospora species are mostly host-specific
(with few exceptions) related to a single host species, host
genus or closely related host genera (Braun et al. 2013; Crous
et al. 2013). Two species of Pseudocercospora has been
reported earlier on Vernonia, Pseudocercospora cinereae
(Deighton 1976) and Pseudocercospora vernoniacearum
(Shukla et al. 1982). Pseudocercospora cineriae has dark
brown circular, coalescing leaf spots and P. vernoniacearum
has oval, effuse leaf spots whereas P. vernoniae has grayish
brown, angular and vein limited leaf spots. Conidiophores
are much smaller (14–40 µm) and more septate (1–8) in
P. vernoniae compared to previously described species P.
cinereae (1–3 septate, 40–150 × 3.5–5 µm) and P. vernoniacearum (44–133 × 3.5–5.4 µm). Conidia are simple and
longer in P. cinereae (28.5–145 × 2.8–5.7 µm) and P. vernoniacearum (40–100 × 3.5–5.4 µm). The presence of catenate conidia in branched chains with smaller and variable
in size 21.74– 44.76 × 4.5–5.5 µm) differentiate P. vernoniae
from the previously described species. Molecular analysis
based on combined gene analysis of LSU, ITS, act and tef1
(Fig. 3) reveals that P. vernoniae clusters with P. hakeae
(CBS 144520) with moderate support.
Subclass Pleosporomycetidae C.L. Schoch, Spatafora,
Crous & Shoemaker
Kirschsteiniotheliales Hern. -Restr., R.F. Castañeda,
Gené & Crous
Kirschsteiniotheliales was introduced by (HernandezRestrepo et al. 2017) based on phylogenetic analysis.
Kirschsteiniotheliales consists with Kirschsteiniotheliaceae,
and two genera incertae sedis, viz. Brachysporiella, Taeniolella (Hongsanan et al. 2020a; Wijayawardene et al. 2020)
Kirschsteiniotheliaceae Boonmee & K.D. Hyde, in Boonmee et al., Mycologia 104(3): 705 (2012)
The monotypic family, Kirschsteiniotheliaceae, was introduced by Boonmee et al. (2012) to accommodate Kirschsteiniothelia species based on morphology and phylogenetic
13
10
Fungal Diversity (2022) 117:1–272
Fig. 3 Maximum likelihood tree illustrating the phylogeny of Pseudocercospora vernoniae with related species in Pseudocercospora
based on LSU, ITS, act and tef1 concatenated sequences. Branches
are labelled with ML and MP values ≥ 50% and BYPP ≥ 0.95 are indi-
cated above the node respectively. The ex-types/reference strains are
in bold; the new species is in blue. The tree is rooted with Pallidocercospora heimii (CPC:11716)
analyses. Kirschsteiniotheliaceae species are mostly saprobes on dead wood from terrestrial and aquatic habitats in
tropical and subtropical regions (Boonmee et al. 2012; Su
et al. 2016; Mehrabi et al. 2017; Bao et al. 2018; Sun et al.
2021).
Kirschsteiniothelia acutisporum S. Wang, Q. Zhao & K.D.
Hyde, sp. nov.
Index Fungorum number: IF559759; Facesoffungi number: 1FoF1799; Fig. 4
Etymology: Named after the acute conidia
Holotype: MFLU 21-0127
Saprobic on decaying plant substrates. Sexual morph:
Not observed. Asexual morph: Colonies effuse, scattered,
dark-brown to black, glistening, hairy, sparse. Mycelium
partly superficial, partly immersed in the substratum,
Kirschsteiniothelia D. Hawksw., Bot. J. Linn. Soc. 91: 182
(1985)
We follow the latest treatment and updated accounts of
Kirschsteiniothelia in Sun et al. (2021)
13
Fungal Diversity (2022) 117:1–272
11
Fig. 4 Kirschsteiniothelia
acutisporum (MFLU 21–0127,
holotype) a–c Colonies on dead
wood. d, e Conidiophore with
conidia. f, g Conidiogenous
cells and conidia. h Conidiogenous cell. i–m Conidia. n
Germinating conidium. Scale
bars: a = 100 μm, b = 200 μm,
c–e = 100 μm, f, g = 50 μm,
h = 20 μm, i–n = 50 μm
composed of dark brown, septate, branched hyphae. Conidiophores macronematous, mononematous, solitary, cylindrical, straight or slightly flexuous, dark brown, slightly
tapering towards the apex, 8–12 septate, truncate at the
apex, 180–260 µm ( x̄ = 230 µm, n = 10) long, 7–12.5 μm
( x̄ = 9 µm, n = 10) wide. Conidiogenous cells integrated,
terminal, monoblastic, cylindrical and brown, calyciform.
Conidia acrogenous, solitary, obclavate to obspathulate,
tapering to the apex, rostrate, 7–12-euseptate, mid to dark
brown, becoming pale brown to pale towards the apex,
truncate at the base, 75–120 µm ( x̄ = 92 µm, n = 15) long,
10.5–19.5 μm ( x̄ = 15 µm, n = 15) wide.
Material examined: Thailand, Chiang Mai Province, saprobic on decaying wood at the Mushroom Research Center
(MRC), August 2020, Song Wang, SW231 (MFLU 21-0127,
holotype).
GenBank numbers: ON980758 (LSU); ON980754 (SSU);
OP120780 (ITS); OP009582 (rpb2)
Notes: Kirschsteiniothelia acutisporum shares
similar characteristics with K. fluminicola in having
13
12
macronematous, unbranched, cylindrical, septate, conidiophores and solitary, obclavate, septate, conidia. However,
Kirschsteiniothelia acutisporum differs from K. fluminicola in having a gelatinous rounded sheath at the apex
of shorter and thinner conidia (33–43 × 7.5–8.5 μm vs
47.5–86.5 × 8–10 μm). Kirschsteiniothelia acutisporum
phylogenetically creates an independent branch with
100ML/100MP/1.00BYPP support (Fig. 5).
Kirschsteiniothelia crustaceum S. Wang, Q. Zhao & K.D.
Hyde, sp. nov.
Index Fungorum number: IF559760; Facesoffungi number: FoF11802; Fig. 6
Etymology: Referring to the conidial ‘shell’ shape.
Holotype: MFLU 21-0129
Saprobic on decaying bamboo culms. Sexual morph: Not
observed. Asexual morph: Colonies effuse, scattered, dark
brown to black, glistening, hairy, sparse. Mycelium partly
superficial, partly immersed in the substratum, composed
of dark brown, septate, branched hyphae. Conidiophores
macronematous, mononematous, solitary, cylindrical,
straight or slightly flexuous, brown to dark brown, slightly
tapering towards the apex, 4–8 septate, truncate at the apex,
60–170 µm ( x̄ = 128 µm, n = 15) long, 6.5–10.5 μm ( x̄ = 8 µm,
n = 15) wide. Conidiogenous cells integrated, terminal,
monoblastic, cylindrical and calyciform, brown, 9–16 µm
( x̄ = 12 µm, n = 15) long, 5.5–8 μm ( x̄ = 6.5 µm, n = 15)
wide. Conidia acrogenous, solitary, obclavate to obspathulate, globose to the apex and hyaline to light brown, rostrate,
5–6-euseptate, mid to dark brown, becoming pale brown
to pale towards the apex, truncate at the base, 45–75 µm
( x̄ = 55 µm, n = 20) long, 10–18 μm (x̄ = 14 µm, n = 20) wide.
Material examined: Thailand, Nang Lae, Mueang Chiang
Rai, Chiang Rai Province, saprobic on decaying bamboo,
submerged in a freshwater stream, July 2020, Rongju Xu,
MD71 (MFLU 21–0129, holotype)
GenBank numbers: MW851854 (LSU); MW851849
(ITS)
Notes: Kirschsteiniothelia crustaceum shares similar
morphology with K. rostrata in having macronematous,
unbranched, cylindrical, septate, conidiophores and solitary, obclavate, septate, conidia. However, conidiophores of
Kirschsteiniothelia crustaceum (60–170 × 6.5–10.5 μm) are
much shorter than those of K. rostrata (up to 280 μm long,
12 μm wide). Conidia of K. crustaceum (45–75 × 10–18 μm)
are much shorter than those of K. rostrata (up to 115 μm
long, 15 μm wide) also. The combined LSU, SSU and ITS
phylogenetic analysis show that Kirschsteiniothelia crustaceum represents a sister taxon to K. rostrata with good
separation (89ML/1.00BYPP) (Fig. 5).
Kirschsteiniothelia extensum. S. Wang, Q. Zhao & K.D.
Hyde, sp. nov.
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Fungal Diversity (2022) 117:1–272
Index Fungorum number: IF559761; Facesoffungi number: FoF11803; Fig. 7
Etymology: Referring to the conidiophore extending
characteristic
Holotype: MFLU 21-0130
Saprobic on decaying wood. Sexual morph: Not
observed. Asexual morph: Colonies effuse, scattered,
brown or black, hairy, glistening. Mycelium partly superficial, partly immersed in the substratum, composed
of brown, septate, branched hyphae. Conidiophores
macronematous, mononematous, solitary, cylindrical,
straight or slightly flexuous, dark brown, unbranched,
thick-walled, smooth, slightly tapering towards the apex,
4–9 septate, truncate at the apex, 80–230 µm ( x̄ = 140 µm,
n = 15) long, 6.5–9.5 μm ( x̄ = 7.5 µm, n = 15) wide. Conidiogenous cells integrated, terminal, monoblastic, percurrent, pale brown, cylindrical, 11–19 µm ( x̄ = 15 µm,
n = 15) long, 4–7.5 μm ( x̄ = 6 µm, n = 15) wide. Conidia
acrogenous, solitary, smooth, obclavate, straight or slightly
curved, tapering to the apex, 5–8-euseptate, becoming
pale brown to pale towards the apex, truncate at the base,
45–120 µm ( x̄ = 60 µm, n = 30) long, 5–12 μm ( x̄ = 9 µm,
n = 30) wide.
Material examined: Thailand, Nang Lae, Mueang Chiang
Rai, Chiang Rai Province, saprobic on decaying wood, July
2020, Rongju Xu, MD73 (MFLU 21-0130, holotype).
GenBank numbers: MW851855 (LSU); MW851850
(ITS)
Notes: Kirchsteiniothelia extensum is introduced here
based on both morphology and molecular data. Kirchsteiniothelia extensum forms a distinct clade within Kirschsteiniotheliaceae and is sister to K. submersa (Fig. 5). The
difference between them is that conidiophores of Kirschsteiniothelia extensum (80–230 × 6.5–9.5 μm) are much shorter
than those of K. submersa (220–280 × 6–7 μm)
Kirschsteiniothelia septemseptatum S. Wang, Q. Zhao &
K.D. Hyde, sp. nov.
Index Fungorum number: IF559762; Facesoffungi number: FoF11800; Fig. 8
Etymology: Referring to the number of septa mostly
observed in conidia
Holotype: MFLU 21–0126
Saprobic on decaying wood. Sexual morph: Not
oberved. Asexual morph: Colonies on natural substrate,
scattered or fascicular, effuse, hairy, dark brown to black,
glistening. Mycelium partly superficial, partly immersed in
the host tissue, composed of smooth, light brown, branched,
septate. Conidiophores macronematous, mononematous,
single to loosely fasciculate, erect, straight to slightly flexuous, branched at the apex, dark brown, multiseptate, 9–16
septate, 250–580 µm (x̄ = 415 µm, n = 20) long, 6.5–14.5 μm
(x̄ = 10 µm, n = 20) wide. Conidiogenous cells mostly
Fungal Diversity (2022) 117:1–272
Fig. 5 Phylogram generated from maximum likelihood analysis based
on combined LSU, SSU, ITS sequence data. Forty-six taxa were
included in the combined analyses, which comprised 2,104 characters (LSU = 1–788 bp, SSU = 789–1,632 bp, ITS = 1,633–2,104 bp),
including alignment gaps. Among them, 1,191 characters were constant, 239 characters were singleton sites, and 674 characters were
13
parsimony informative. The best scoring RA × ML tree is presented.
Bootstrap support values for ML and MP ≥ 75% and BYPP ≥ 0.95
are given above the nodes. Pseudorobillarda eucalypti (MFLUCC
12–0422) and P. phragmitis (CBS 398.61) were used as the outgroup
taxa. The newly generated sequences are indicated in red. The ex-type
strains are indicated in bold
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Fungal Diversity (2022) 117:1–272
Fig. 6 Kirschsteiniothelia
crustaceum (MFLU 21–0129,
holotype) a, b Colonies on dead
wood. c Conidiophore with
conidia. d–g Conidiogenous
cells and conidia. h–k Conidia.
l Germinating conidium.
m Culture on MEA. Scale
bars: b–e = 50 μm, f = 20 μm,
g = 50 μm, h–l = 20 μm
polytretic, sometimes monotretic, integrated, discrete, terminal and lateral, calyciform, 2 septate, 9.5–21 µm (x̄ = 16 µm,
n = 20) long, 4–8 μm (x̄ = 6 µm, n = 20) wide. Conidia acrogenous, solitary, dry, olivaceous brown to brown, pale at apex,
obclavate, rostrate, smooth, straight or curved, truncate at
base, 5–8– euseptate, 25–55 μm (x̄ = 41 µm, n = 20) long,
6.5–12.5 µm ( x̄ = 10.5 μm, n = 20) wide.
Material examined: Thailand, ChiangMai Province, saprobic on decaying wood at MRC, July 2020, Song Wang,
SW212, (MFLU 21–0126, holotype)
GenBank numbers: ON980757 (LSU); ON980752 (SSU);
OP120779 (ITS); OP009581 (rpb2)
Notes: Kirschsteiniothelia septemseptatm shares similar
characteristics with K. fluminicola in having macronematous,
13
unbranched, cylindrical, septate, conidiophores and solitary, obclavate, septate, conidia. However, K. cangshanensis differs from K. fluminicola in having a gelatinous
rounded sheath at the apex of shorter and thinner conidia
(33–43 × 7.5–8.5 μm vs 47.5–86.5 × 8–10 μm). In our phylogetic analyses, K. septemseptatum forms an independent
branch with 91ML/95MP/1.00BYPP support (Fig. 5)
Kirschsteiniothelia spatiosum. S. Wang, Q. Zhao & K.D.
Hyde, sp. nov.
Index Fungorum number: IF559763; Facesoffungi number: FoF11801; Fig. 9
Etymology: Referring to the long conidia
Holotype: MFLU 21-0128
Fungal Diversity (2022) 117:1–272
15
Fig. 7 Kirschsteiniothelia
extensum (MFLU 21–0130,
holotype) a, b Colonies on
dead wood. c, d Conidiophore
with conidia. e Conidiophore.
f–h Conidiogenous cells and
conidia. i–k Conidia. l Germinating conidium. m Culture on
MEA. Scale bars: c–e = 50 μm,
f–l = 20 μm
Saprobic on decaying wood. Sexual morph: Not oberved.
Asexual morph: Colonies effuse on natural substrate, scattered or fascicular, hairy, black, glistening. Mycelium partly
immersed, partly superficial in the substrate, composed of
pale brown, ranched hyphae. Conidiophores macronematous, mononematous, solitary or sometimes caespitose,
cylindrical, wide at base, tapering towards apex, straight
or slightly flexuous, smooth, light brown to dark brown,
unbranched, 6–12 septate, 70–128 µm (x̄ = 100 µm, n = 15)
long, 7.5–12.5 μm (x̄ = 9 µm, n = 15) wide. Conidiogenous
cells holoblastic, monoblastic, integrated, terminal, determinate, cylindrical, smooth, mid to dark brown. Conidia
acrogenous, solitary, dry, olivaceous brown to brown, pale at
apex, obclavate, rostrate, smooth, straight or curved, truncate
at base, 8–23– euseptate, sometimes with a mucilaginous
sheath, 90–139 μm (x̄ = 113 µm, n = 15) long, 9.5–16.5 µm
(x̄ = 14 μm, n = 15) wide.
Material examined: Thailand, Chiang Mai Province, saprobic on decaying wood at MRC, August 2020, Song Wang,
SW280 (MFLU 21–0128, holotype)
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Fig. 8 Kirschsteiniothelia septemseptatum (MFLU 21–0126, holotype) a–c Colonies on dead wood. d, e Conidiophore with conidia. f–h Conidiogenous cells and conidia. i–m Conidia. n Germinating conidium. Scale bars: a = 500 μm, b = 200 μm, c = 50 μm, d, e = 100 μm, f–n = 20 μm
GenBank numbers: OP077294 (LSU); ON980753 (SSU)
Notes: In the phylogenetic analyses our strain is
closely realted with K. tectonae (Fig. 5). Kirschsteiniothelia spatiosum shares similar characteristics with
Kirschsteiniothelia tectonae in having macronematous,
13
unbranched, cylindrical, septate, conidiophores and solitary, obclavate, septate, conidia. However, K. spatiosum
differs from K. tectonae in having a gelatinous rounded
sheath at the apex of shorter and thinner conidia and in
having shorter and thinner conidia (90–139 × 9.5–16.5 μm
Fungal Diversity (2022) 117:1–272
17
Fig. 9 Kirschsteiniothelia spatiosum (MFLU 21–0128, holotype) a–c Colonies on dead wood. d, e Conidiophore with conidia. f Conidiogenous
cells and conidia. g Conidiophore. h–l Conidia. m Germinating conidium. Scale bars: a–c = 100 μm, d–g = 50 μm, m = 100 μm
vs 135–150 × 16–19 μm). Kirschsteiniothelia spatiosum
differs from K. tectonae in having shorter conidiophores
(70–128 × 7.5–12.5 μm vs 200 × 4–8 μm).
Pleosporales Luttrell ex M.E. Barr.
Notes: We follow the latest treatments and updated
accounts of Pleoporales in Hongsanan et al. (2020b) and
Wijayawardene et al. (2022).
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18
Amorosiaceae Thambug. & K.D. Hyde
Thambugala et al. (2015) introduced this family to accommodate Amorosia Mantle & D. Hawksw. and Angustimassarina Thambug., Kaz. Tanaka & K.D. Hyde. The family
is characterized by immersed or semi-immersed ascomata
with a short, crest-like papilla, and hyaline ascospores with
a mucilaginous sheath (Thambugala et al. 2015). Wijayawardene et al. (2022) accepted five genera in this family.
Angustimassarina Thambug., Kaz. Tanaka & K.D. Hyde
Fig. 10 Angustimassarina
kunmingense (YHD216, holotype). a, b Ascomata immersed
on host surface. c Section
through ascoma. d, e Peridium.
f, g Mature bitunicate asci (g.
asci stained with Congo red).
h–k Ascospores. l Gemmating
ascospores. m, n Colonies on
PDA. Scale bars: c–d = 100 μm,
e = 50 μm, l = 30 μm,
f–g = 20 μm, i–k = 10 μm,
h = 5 μm
13
Fungal Diversity (2022) 117:1–272
Thambugala et al. (2015) introduced this genus to accommodate fungi that have ascospores resembling Massarina,
while being narrowly fusiform. There are 12 species listed
in the Index Fungorum (accessed on 30 August 2022). In
this study, we introduce a new species from China based on
molecular phylogeny and morphology.
Angustimassarina kunmingense H.D. Yang & K.D. Hyde,
sp. nov.
Fungal Diversity (2022) 117:1–272
19
Index Fungorum number: IF559764; Facesoffungi number: FoF11804; Fig. 10
Etymology: Referring to the collecting site, Kunming
City, Yunnan, China.
Holotype: HKAS123210
Saprobic on dead aerial stem of Camellia semiserrata.
Sexual morph: Ascomata (162–)190–332(–333) × (119–
)142–289(–300) μm (x̅ = 261 × 221 μm, n = 5), scattered,
gregarious, immersed to semi-immersed in the host tissue,
black, globose to subglobose, ostiolate. Ostiole in the centre, crest-like, rounded, papillate, with a pore-like opening.
Peridium 27–56 μm thick, comprised of 5–10 layers of cells of
textura angularis, cells smaller at the base and the apex, and
larger at the side, brown to hyaline. Hamathecium composed
of 1.2–2 μm (x̅ = 1.6 μm, n = 30) wide, numerous, septate,
clamped, unbranched, hyaline, pseudoparaphyses, embedded in a gelatinous matrix, longer than asci. Asci (56–) 60–
74(–77) × (7.2–)7.5–8.7(–9.3) μm (x̅ = 68 × 8.1 μm, n = 20),
8-spored, bitunicate, fissitunicate, cylindric-clavate, with short
pedicel at the base, rounded at the apex with a minute ocular
chamber. Ascospores (18–)20–22(–23) × (3.1)3.3–3.8(–4.1)
μm (x̅ = 20 × 3.5 μm, n = 30), 1–2 overlapping seriate, hyaline, fusiform, dimidiate, widest at the centre and tapering
toward the ends, with 1–3 constricted septate septum, filled
with 1–2 guttules per cell, smooth-walled and surrounded by
a mucilaginous sheath. Asexual morph: Not observed.
Culture characteristics: Ascospores germinating on PDA
within 24 h and producing germ tubes from both ends and
sides. Colonies on PDA reaching 28 mm diam. after 33 days
at 20 °C, nearly circular, flat, dense, radial sulcate, edge
entire, smoke grey to grey-white on the surface, dark brown
on the reverse and becoming grey-white at the margin.
Material examined: China, Yunnan Province, Kunming
City, Panlong District, on Camellia semiserrata C.W. Chi
(Theaceae), 25° 8′ 29.27″ N, 102° 44′ 16.03″, 17 Dec 2021,
Hongde Yang, (HKAS123210, holotype); ex-type living
culture, KUNCC22-10799.
GenBank numbers: ON352672 (ITS); ON352671 (LSU);
ON352675 (SSU); ON364144 (tef1); ON791602 (act);
ON791682 (tub2)
Notes: Species of Angustimassarina are broadly distributed in Belgium, Germany and Italy (Hyde et al. 2020a, b,
c; Phukhamsakda et al. 2020), but, have never been reported
from China. Our collections from China are morphologically and phylogenetically related to Angustimassarina. Our
new species Angustimassarina kunmingense resembles other
Table 1 Synopsis of mainly morphological characteristics of Angustimassarina species
Taxa
Host
Ascomata (µm)
Peridium Asci (µm)
(µm)
Ascospores (µm) Locality References
Angustimassarina
acerina
MFLUCC 14–0505
Acer platanoides
200–350 × 164–183
15–26
92–105 × 7.5–8.6 21–23 × 4.1–4.6
Germany
Thambugala et al. (2015)
A. alni
MFLUCC 15–0184
Alnus glutinosa
160–250 × 130–200
28–44
71–89 × 8–10
19–22 × 3–4
Germany
Tibpromma et al. (2017)
A. arezzoensis
MFLUCC 13–0578
Salvia sp.
169–234 × 166–245
22–41
67–95 × 10–15
19–21 × 5–6
Italy
Tibpromma et al. (2017)
A. camporesii
MFLU 18–0057
Galium sp.
130–240 × 130–190
15–21
62–88 × 10–13
15–18 × 4–5
Italy
Hyde et al. (2020a, b, c)
A. coryli
MFLU 15–2603
Corylus avellana
150–250 × 500–750
8–12
95–110 × 8– 12
21–25 × 4–5
Italy
Hyde et al. (2017)
A. italica
MFLUCC 15–0082
Ilex aquifolium
127–159 × 97–131
23–40
78–103 × 10–12
15–22 × 3–6
Italy
Tibpromma et al. (2017)
A. kunmingense
YHD216
Camellia
japonica
162–333 × 119–300
27–56
56–77 × 7.2– 9.3
18–23 × 3.1– 4.1 China
This study
A. lonicerae
MFLUCC 15–0087
Lonicera sp.
193–203 × 170–220
10–18
55–81 × 9–13
19–25 × 4–7
Italy
Tibpromma et al. (2017)
A. populi
MFLUCC 13–0034
Populus sp.
125–175 × 100–120
14–32
80–95 × 9.5–13
19–22 × 3.2–5.5
Italy
Thambugala et al. (2015)
A. premilcurensis
MFLUCC 15–0074
Carpinus betulus
231–238 × 290–311
20–30
64–93 × 11–15
19–23 × 4–7
Italy
Tibpromma et al. (2017)
A. quercicola
MFLUCC 14–0506
Quercus robur
200–250 × 150–265
14–27
60–94 × 8.8–13
17–21 × 4–6
Germany
Thambugala et al. (2015)
A. rosarum
MFLU 17–1513
Clematis
viticella, Rosa
canina
221–306 × 267–400
14–40
77–85 × 10–16
17–23 × 4–4.5
Italy
Wanasinghe et al. (2018)
A. sylvatica
MFLU 15–2603
Fagus sylvatica
180–260 × 150–200
8–12
95–110 × 8– 12
21–25 × 4–5
Italy
Hyde et al. (2019)
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Fungal Diversity (2022) 117:1–272
Angustimassarina species in terms of ascomata, asci and
ascospores (Table 1) and the new species was isolated from
similar habitat to other Angustimassarina species (Thambugala et al. 2015). However, the taxon is charactered by slender asci and ascospores. The megablast search of the ITS and
tef1 sequences show the highest similarity with Angustimassarina populi (457/463, 98%) and Angustimassarina populi
(827/830, 99%), respectively. In the phylogenetic analysis,
Angustimassarina kunmingense formed a well-supported
monophyletic clade basal to Angustimassarina species
(97ML/1.00BYPP). Our phylogenetic tree was constructed
using multi gene loci (ITS, SSU, LSU and tef1, Fig. 11).
However, most taxa were not strongly supported. This could
suggest that additional markers are required to achieve a
Fig. 11 Phylogram generated from maximum likelihood analysis based on combined ITS, SSU, LSU and tef1 sequence data of
Angustimassarina. Twenty strains were included in the analysis
of the combined loci which comprised 2700 characters. The tree is
rooted with Guttulispora crataegi (MFLUCC 13–0442) and Lophiostoma caulium (KT794). Bootstrap support values ≥ 50% in ML
and BYPP ≥ 0.95 are given at the nodes. The ex-types and reference
strains are in bold; the new isolate is in blue
13
Fungal Diversity (2022) 117:1–272
more accurate identity, thus we also provide protein gene
act and tub2 herein.
Bambusicolaceae D.Q. Dai & K.D. Hyde, in Hyde et al.,
Fungal Diversity 63: 49 (2013)
Notes: Bambusicolaceae was placed in Dothideomycetes
by Hyde et al. (2013) to accommodate Bambusicola (Dai
et al. 2012; Liu et al. 2015; Jayasiri et al. 2019; Yang et al.
2019; Bhunjun et al. 2021; Calabon et al. 2022). Four genera viz. Bambusicola, Corylicola, Leucaenicola and Palmiascoma are accepted in this family (Wijayawardene et al.
2022). Bambusicolaceae are characterized by solitary, scattered, immersed, semi-immersed to erumpent and conical or
globose to subglobose ascomata, anastomosing, branching
interascal filaments, cylindrical to clavate asci with a short
furcate or rounded to obtuse pedicel and slightly broad-fusiform or clavate to ellipsoidal, hyaline or yellowish to brown,
single-septate ascospores with a gelatinous sheath (Dai et al.
2012; Hyde et al. 2013; Liu et al. 2015; Dai et al. 2017).
Bambusicola D.Q. Dai & K.D. Hyde, in Dai, Bhat, Liu,
Chukeatirote, Zhao & Hyde, Cryptog. Mycol. 33(3): 367
(2012)
Notes: Bambusicola is a well-studied genus, established
by Dai et al. (2012). There are 15 species accepted in the
genus and all species have sequence data in GenBank (Dai
et al. 2012, 2015, 2017; Thambugala et al. 2015; Yang et al.
2018; Dong et al. 2020; Monkai et al. 2021). Both sexual
and asexual morphs of Bambusicola are reported (Dai et al.
2012, 2015, 2017; Thambugala et al. 2015; Yang et al. 2018;
Dong et al. 2020; Monkai et al. 2021). The sexual morph
of Bambusicola is characterized by gregarious, immersed
or semi-immersed, globose to subglobose, uni- to multiloculate, coriaceous ascomata, bitunicate, cylindrical or
cylindric-clavate, short pedicellate asci with a shallow or
well-developed chamber and fusiform, septate, hyaline to
pale brown ascospores mostly surrounded by a gelatinous
sheath. The asexual morph of Bambusicola is characterized
by pycnothyrial, immersed to semi-immersed, acerose or
subglobose, pyriform or irregular, uni- to multi-loculate conidiomata, holoblastic, annellidic, discrete, cylindrical conidiogenous cells and cylindrical to ellipsoidal, pale brown to
brown, septate conidia (Dai et al. 2012, 2017; Thambugala
et al. 2015; Dong et al. 2020; Monkai et al. 2021).
Bambusicola species have been reported from both terrestrial and freshwater habitats in China and Thailand (Dai
et al. 2012, 2015, 2017; Thambugala et al. 2015; Yang et al.
2018; Dong et al. 2020). Most Bambusicola species are
reported as saprobes on bamboo. In this study, we report a
new record of Bambusicola bambusae on submerged decaying wood from freshwater habitats for the first time.
21
Bambusicola bambusae D.Q. Dai & K.D. Hyde, Cryptog.
Mycol. 33(3): 372 (2012)
Index Fungorum number: IF 801046; Facesofungi number: FoF11797; Fig. 12
Saprobic on decaying wood in a freshwater stream.
Ascomata 135–175 µm high × 190–245 µm diam.
( x̄ = 155 × 216 µm, n = 10), solitary, scattered to gregarious,
immersed under the host tissue, conical in section, brown to
dark brown, coriaceous, subglobose, ostiolate. Ostiole crestlike, central, elongated to papillate, with a pore-like opening,
plugged by hyaline, filamentous hyphae. Peridium comprising host and fungal tissues, 17–31 μm thick, composed of
brown to dark brown cells of textura angularis intermingled
with host cells. Hamathecium composed of numerous, filamentous, hyaline, septate, branched, 1.0–1.5 μm, pseudoparaphyses. Asci 55–75 × 7.5–9.5 μm ( x̄ = 66.3 × 8.5 μm, n = 20),
8-spored, bitunicate, fissitunicate, cylindrical, with a shallow apical chamber and a short furcate pedicel. Ascospores
19–21 × 4.0–4.5 μm ( x̄ = 20 × 4.5 μm, n = 30), 2–3-seriate,
1-septate, constricted at the septum, slightly broad fusiform, tapering towards the ends, occasionally with large
upper cell, with narrowly rounded ends, hyaline, guttulate,
smooth-walled.
Culture characteristics: Ascospores germinating on
PDA within 24 h and germ tubes produced from both ends.
Colonies growing on PDA, reaching a diam. of 20–25 mm
after 20 d at 25 °C, surface smooth to velvety, with entire to
slightly undulate edge, greenish in the centre, white at the
edge; reverse dark greenish to black in the centre, white at
the edge.
Material examined: Thailand, Tao Ngoi, Sakon Nakhon,
on decaying wood submerged in a river, 12 November 2017,
D.F. Bao, B110 (MFLU 22–0080), living culture, MFLUCC
22–0021.
Host/Substrate: Bamboo (Poaceae) (Dai et al. 2012);
decaying wood submerged in a river (this study)
Distribution: Thailand (Dai et al. 2012; this study)
GenBank numbers: ON764309 (ITS); ON764310 (LSU);
ON764313(SSU); ON788004 (rpb2)
Notes: In the phylogenetic analysis, our new isolate
B110 clustered with the ex-type strain of Bambusicola
bambusae (MFLUCC 11–0614) with 98% ML/1.00 BYPP
support (Fig. 13). The morphology of our collection is
almost identical to the holotype of Bambusicola bambusae except for the size of ascomata and the sheath
of the ascospores. The ascomata of our collection are
smaller than the holotype (190–245 vs. 450–70 µm diam)
and the holotype of B. bambusae has ascospores with
a thick sheath (Dai et al. 2012), whereas, the sheath of
ascospores were not observed in our collection. A comparison of the ITS and rpb2 gene regions of MFLUCC
11–0614 and B110 revealed 0 and 3 base pair differences and therefore we identified our new collection as
13
22
Fig. 12 Bambusicola bambusae (MFLU 22–0080, new record).
a–c ascomata on wood d section of ascoma e peridium, f, g ostiole. h pseudoparaphyses. i–k asci. l–p ascospores. q Germinating
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Fungal Diversity (2022) 117:1–272
ascospore. r, s culture on PDA from surface and reverse. Scale bars:
b, f = 100 μm, e = 50 μm, g = 30 μm, h–k = 20 μm, l–q = 10 μm
Fungal Diversity (2022) 117:1–272
23
Fig. 13 Phylogram generated from ML analysis, based on combined ITS, LSU, SSU, tef1 and rpb2 sequence data for Bambusicolaceae. The combined dataset comprises 37 strains with 4617 characters including gaps (LSU: 854 bp, SSU: 1016 bp, ITS: 805 bp,
tef1: 950 bp, rpb2: 992 bp). The tree is rooted with Murilentithe-
cium lonicerae (MFLUCC 18–0675) and M. clematidis (MFLUCC
14–0561). Maximum likelihood bootstrap values ≥ 75% and baysian
BYPP ≥ 0.95 are displayed on the nodes, respectively. Newly introduced taxa are indicated in red. Ex-type and representative strains are
in bold
Bambusicola bambusae as recommended by Pem et al.
(2021). Bambusicola bambusae was described by Dai
et al. (2012), it was collected on bamboo from terrestrial
habitats in Thailand. Our collection was from freshwater
habitats and this is the first time this species reported
from freshwater habitats.
Corylicola was introduced by Wijesinghe et al. (2020).
This genus is characterized by uniseriate, fusiform to ellipsoidal, yellowish to pale brown, single-septate, echinulate
ascospores, accumulating as yellowish-brown masses at the
apices of ascomatal neck (Wijesinghe et al. 2020). We provide a new host record of Corylicola italica from Rubus sp.
in Italy.
Corylicola Wijesinghe, Camporesi, Yong Wang bis &
K.D. Hyde, in Wijesinghe et al., Biodiversity Data Journal
8(e55957): 8 (2020)
Corylicola italica Wijesinghe, Camporesi, Yong Wang bis
& K.D. Hyde, in Wijesinghe et al., Biodiversity Data Journal
8(e55957): 8 (2020).
13
24
Fig. 14 Corylicola italica (MFLU 20–0251, new host record). a–b
Appearance of ascomata on a twig of Rubus sp. c–d Section through
ascomata. e Close-up of ostiole. f Pseudoparaphyses. g Peridium.
h–j Asci. k–n. Ascospores. o. Culture characteristics on PDA after
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Fungal Diversity (2022) 117:1–272
20 days from above. p Culture characteristics on PDA after 20 days
from below. Scale bars: c–d = 50 μm, e–g = 20 μm, h–j = 10 μm,
k–n = 5 μm
Fungal Diversity (2022) 117:1–272
Index Fungorum number: IF557768; Facesofungi number: FoF08684; Fig. 14
Saprobic on a dead branch of Rubus sp. Sexual
morph: Ascomata 109–141 high, 91.5–106 µm diam.
(x̄ = 128.5 × 101.5 µm; n = 4), solitary, scattered, immersed,
erumpent at maturity, raised as brown to dark spots on the
substrate, globose to subglobose, coriaceous, uni-loculate
with an ostiole. Ostiole 46–68 µm wide, central, papillate,
lined with hyaline periphyses. Peridium composed of two
layers, unequally thickened, 15–29 µm wide comprising
brown, blackish to dark brown cells of textura angularis
fused with host tissues, inner layer comprising hyaline cells
of textura prismatica. Hamathecium comprising numerous pseudoparaphyses 1–2 µm wide (x̄ = 1.6 µm, n = 6),
filamentous, cellular, with distinct septa, not constricted
at the septa, branching and anastomosing above the asci.
Asci 52–74 × 4–6 µm (x̄ = 61 × 5 µm, n = 5), 8-spored, bitunicate, fissitunicate, cylindrical, short distinct pedicel with
furcate ends, apically rounded, well-developed ocular chamber. Ascospores 10–12 × 3–4 µm (x̄ = 10 × 3.6 µm, n = 11),
overlapping, uni-seriate, fusiform to ellipsoidal, 1-septate
straight, hyaline and yellowish when young, becoming pale
brown at maturity. Asexual morph: Not observed.
Culture characteristics: Spore germinating on PDA
within 24 h from singles pore isolation. Colonies on PDA
reaching 10 mm diam. after 20 days at 20 °C, circular, submerged, crenated edge, flat with dense, brown to whitish in
the middle, grey at the edges from upper and reverse brownish-black in the lower surface of the colony.
Material examined: Italy, Forlì-Cesena Province near
Meldola, on dead aerial branches of Rubus sp. (Rosaceae),
4 February 2020, Erio Camporesi IT-4596C (MFLU
20–0251); living culture MFLUCC 21-0118.
Host/Substrate: Corylus avellana (Betulaceae) (Wijesinghe et al. 2020); Rubus sp. (Rosaceae) (this study)
Distribution: Italy (Wijesinghe et al. 2020; this study).
GenBank numbers: OM471788 (ITS), OM630433 (tef1).
Notes: Wijesinghe et al. (2020) reported this species from
Corylus avellana. Morphologically our collection resembles
the ex-type strain of this species. Based on our phylogenetic
analyses, our strain MFLUCC 21-0118 clustered together
with MFLU 19–0500 and MFLUCC 20–0111 (Fig. 13) with
100/ML and1.00/BYPP support. Therefore, we introduce our
collection as a new host record.
Palmiascoma Phook. & K.D. Hyde, in Liu et al., Fungal
Diversity: https://doi.org/10.1007/s13225-015-0324-y, [65]
(2015).
Notes: Palmiascoma was introduced by Liu et al. (2015)
and is typified by P. gregariascomum collected from a dead
frond of a palm. Palmiascoma is similar to Didymosphaeria
in having didymosporous, brown, and echinulate ascospores,
but differs in phylogeny. Monkai et al. (2021) introduced the
25
second species into the genus as P. qujingense isolated from
dead twigs of Fagaceae sp. in Yunnan, China.
Palmiascoma gregariascomum Phookamsak & K.D. Hyde,
in Liu et al., Fungal Diversity: https:// doi. org/ 10. 1007/
s13225-015-0324-y, [65] (2015).
Index Fungorum number: IF550927; Facesoffungi number: FoF00429; Fig. 15
Saprobic on dead twigs of Rosa sp. Sexual morph:
See Liu et al. (2015). Asexual morph: Conidiomata
140–200 μm high, 130–220 μm diam. (x̅ = 172 × 165 μm,
n = 5), pycnidial, solitary or aggregated, immersed, erumpent
neck, visible as black, uni- to multi-loculate, globose to
subglobose, rarely irregular, glabrous, ostiole central, with
minute papilla. Conidiomata walls 14–38 μm (x̅ = 27 μm,
n = 8), wide, thick-walled, of equal thickness, composed
of several layers of hyaline to dark brown, pseudoparenchymatous cells, outer layers comprising 4–5 cell layers of
6–12 × 2–5 μm (x̅ = 8.7 × 3.4 μm, n = 15), thick-walled, dark
brown to black, organized in a textura angularis to textura
prismatica cells, inner layers comprising 2–3 layers of
3–8 × 2–4 μm (x̅ = 5.3 × 3.2 μm, n = 15), thin-walled, hyaline,
organized in a textura angularis. Conidiophores arising from
basal cavity of conidiomata mostly reduced to conidiogenous
cells. Conidiogenous cells 5–7 × 1–2 μm (x̅ = 5.8 × 1.6 μm,
n = 25), holoblastic, phialidic, discrete, ampulliform to cylindrical, hyaline, aseptate, smooth-walled, guttulate. Conidia
3.2–4.5 × 1.7–2.4 μm (x̅ = 3.7 × 2.1 μm, n = 35), in culture
Conidia 3.4–4.7 × 1.6–2.5 μm (x̅ = 4.1 × 2.1 μm, n = 35),
solitary, one-celled, oblong to ellipsoidal, with rounded or
obtuse ends, initially hyaline, becoming brown at maturity,
smooth-walled.
Culture characteristics: Colonies on PDA fast growing,
33–37 mm diam. after 2 weeks at 25–30 °C, greenish-grey
to grey, forming white tufts on surface, slightly radiating;
reverse brown to dark brown at the margin, dark brown to
black in the centre; medium dense, circular, flattened to
slightly raised, dull to rough with entire edge, fairy fluffy to
velvety, slightly radially furrowed.
Material examined: Thailand, Mueang, Chiang Rai District, Chiang Rai 57100, (20° 03′ 24.7′′ N, 99° 52′ 23.5′′ E),
dead twigs of Rosa sp. (Rosaceae), 20 August 2017, MC.
Samarakoon, SAMC070 (MFLU 18–0845, HKAS 102350),
living culture MFLUCC 18-0505.
Hosts: on dead frond of palm (Liu et al. 2015), Rosa sp.
(this study)
Distribution: Thailand (Liu et al. 2015; this study)
GenBank numbers: OM293742 (LSU), OM293753
(SSU), OM305060 (tef1), OM305066 (tub2)
Notes: Our new collection of Palmiascoma gregariascomum is described on dead twigs of Rosa species from
Thailand. We found the asexual morph of the taxon with
a similar range of conidiogenous cells (5–7 × 1–2 μm
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Fungal Diversity (2022) 117:1–272
Fig. 15 Palmiascoma gregariascomum (MFLU 18–0845, new host
record) a-c Conidiomata on the substrate. d Vertical section of conidioma. e Peridium. f–h Conidiogenous cells and conidiogenesis.
i,o Conidia (o in culture), j Top view of culture in PDA. k Reverse
view of culture. l,m Conidiomata on PDA. n Peridium. Scale bars:
a,l = 1000 µm, b = 500 µm, c,m = 200 µm, d = 50 µm, e = 20 µm,
n = 10 µm, f-i,o = 5 µm
vs 5–12 × 2–4 μm) and conidia (3.2–4.5 × 1.7–2.4 μm,
3.4–4.7 × 1.6–2.5 μm vs 4–6 × 2–3 μm) in morphologies
compared to the type species (MFLU 11-0211). In multigene phylogeny, our strain clusters with MFLU 11–0211
with high statistical (81/ML, 1.00 BYPP) support. Based
on similar morphology and phylogenetic analyses, here we
provide a new host record of Palmiascoma gregariascomum
on Rosa sp. from Thailand (Fig. 16).
Coniothyriaceae W.B. Cooke, Revta Biol., Lisb. 12: 289
(1983) [1980–1983]
Notes: Coniothyriaceae was introduced by Cooke (1983)
to accommodate species of Coniothyrium. Kirk et al.
(2008) synonymized Coniothyriaceae with Leptosphaeriaceae. De Gruyter et al. (2013) based on morphology
and phylogenetic analyses showed that the type species C.
palmarum is distinct from Leptosphaeriaceae and reinstated Coniothyriaceae in Pleosporales. Wijayawardene
13
Fungal Diversity (2022) 117:1–272
27
Fig. 16 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU and tef1 sequenced data for Bambusicola and allied genera Twenty-two strains are included in the combined sequence analyses, which comprise 2761 characters with gaps.
Murilentithecium clematidis (MFLUCC 14–0562) is used as the
outgroup taxon. Tree topology of the ML analysis was similar to the
BYPP. Bootstrap support values for ML ≥ 50% and BYPP ≥ 0.95 are
given above the nodes. New strain is in red. Ex-type and representative strains are in bold
et al. (2022) accepted Coniothyrium, Foliophoma, Neoconiothyrium, Ochrocladosporium and Staurosphaeria in
this family.
250–450 μm (x̅ = 329 μm, n = 10) diam., superficial on or
immersed in the host. Conidiophores not present. Conidiogenous cells lining entire cavity, hyaline, cylindrical, 8 ± 15 (x̅ = 11.9 μm) 3 ± 8 μm (x̅ = 5.1 μm), longer in
culture than on host plants. Conidiogenesis holoblastic,
proliferating percurrently. Conidia cylindrical, broadly
rounded at apex, initially somewhat truncate at base,
produced deep within the conidiogenous cells, secession
rhexolytic, outer wall of conidiogenous cell often remaining on conidium, except at base, eventually disintegrating,
olivaceous brown, 3-septate, lightly punctate, (2–)6.6–10.
5(–14) × (1–)2–5(–6) μm (x̅ = 7.3 × 3.6 μm, n = 40) brown,
smooth-walled or verruculose, aseptate, curved, both sides
gradually tapering towards the round to slightly acute apex
and truncate base, guttulate. Sexual morph: Not observed.
Material examined: Russia, Donetsk People's Republic,
Donetsk City, Donetsk Botanical Garden, flowerbed, on
dying peduncle stem and live leaves of Yucca filamentosa
L. (Asparagaceae), 20 May 2017, Timur S. Bulgakov, DNK108 (MFLU 17–2529, holotype); ex-type living culture
MFLUCC 18–0456.
GenBank numbers: OM235094 (SSU); OM235097(LSU)
Coniothyrium Corda, Icon. fung. (Prague) 4: 38 (1840)
The genus is typified with C. palmarum Corda. In earlier
studies Contiothyrium was considered as the asexual morph
of Leptosphaeria, Mycosphaerella and Massarina (Sivanesan 1984). However, later studies based on molecular data
transferred many species from Contiothyrium (Verkley et al.
2014; Hongsanan et al. 2020b). De Gruyter et al. (2013)
reinstated Coniothyriaceae and included Coniothyrium as
the family type.
Coniothyrium yuccicola Chaiwan, Jayaward., & K.D. Hyde,
sp. nov.
Index Fungorum number: IF559467; Facesoffungi number: FoF08170; Fig. 17
Etymology: Referring to the host Yucca
Holotype: MFLU 17–2529
Pathogenic on living leaves and peduncle stems
of Yucca filamentosa. Asexual morph: Conidiomata
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Fig. 17 Coniothyrium yuccicola (MFLU 17–2529, holotype) a
Specimen with conidiomata. b Black acervuli. c Brown setae. d Conidiophores with basal parts of setae. e Hyaline conidiogenous cells. f
Conidiomata on PDA. g Hyaline conidia. h Germinating conidium. i
Appressoria. j Reverse view of the colony. k Upper view of the colony. Scale bars: a = 1000 μm, b = 500 μm, c = 20 μm, d = 15 μm, e,
f = 10 μm
Notes: Coniothyrium yuccicola is an asexual morph. Based
on our phylogenetic tree this species is closely related to C.
concentricum (Fig. 18). Conidia of this species are brown
aseptate, bacilliform, ellipsoid and often thick-walled (Fig. 17).
Three Coniothyrium species are recorded on Yucca species
(Farr and Rossman 2022): Coniothryrium bartholomaei from
the USA (Oregon), C. herbarum from the USA (California),
and C. yuccae from Argentina. Coniothyrium bartholomaei
was reported as a plant pathogen that caused leaf spots of
Yucca in Oregon (USA) (Pscheidt and Ocamb 2018; Barr
1992). Coniothyrium herbarum is known from USA (California) on the leaves of several closely related plants: Dracaena
indivisa, Sansevieria sp. and Yucca angustifolia (Cash 1952),
however, this species is invalid. Coniothyrium yuccae was
found on dead leaves of Yucca gloriosa in Argentina (Buenos-Aires) (Farr 1973). Phaeosphaeriopsis yuccae is another
morphologically similar taxon described from living leaves
of Yucca filamentosa from Russia, Rostov region, Botanical
Garden of Southern Federal University (Tibpromma 2017).
(Hongsanan et al. 2020b). Forty-four genera are accepted in
this family (Wijayawardene et al. 2022)
Didymellaceae Gruyter, Aveskamp & Verkley, Mycol. Res.
113(4): 516 (2009)
Members of this family have a wide host range and have
different life modes: endophytic, pathogenic and saprobic
13
Ascochyta Lib., Pl. crypt. Arduenna, fasc. (Liège) 1(Praef.):
8 (1830)
Notes: Ascochyta was introduced by Libert (1830) with
A. pisi as the type species. Species of Ascochyta are characterized by the globose locules with perithecial protuberances immersed in the stroma (Chen et al. 2015). Species
are mostly endophytes, pathogens and saprobes with a wide
host range and a geographical distribution (Hongsanan et al.
2020b; Farr and Rossman 2022). We provide a new host
record of Ascochyta medicaginicola from Prunus cerasifera
in Russia.
Ascochyta medicaginicola Q. Chen & L. Cai, in Chen et al.,
Stud. Mycol. 82: 187 (2015)
Index Fungorum number: IF814129; Facesoffungi number: FoF08216; Fig. 19
Pathogenic on living twigs of Prunus cerasifera, noticeable as black, circular dots on the host surface. Asexual
morph: Conidiomata 165–190 μm high, 170–210 μm wide,
black, scattered or gregarious, superficial to immersed,
black, subglobose to globose, uniloculate. Ostiolar neck
Fungal Diversity (2022) 117:1–272
29
Fig. 18 Phylogram generated from ML analysis based
on combined LSU and SSU
sequence data of selected
taxa. The combined dataset
comprises 41 strains with 1834
characters including gaps. The
tree is rooted to Aposphaeria
populina (CBS543.70) and
Westerdykella capitulum (CBS
337.65). Maximum Likelihood
bootstrap values ≥ 65% and
BYPP ≥ 0.90 are displayed on
the nodes, respectively. Newly
introduced taxa are indicated in
red. Ex-type and representative
strains are in bold
25–50 μm long, 3–5 μm wide, covered with 1-celled, thickwalled, dark brown to almost black. Peridium 15–25 μm
wide at the base, 30–80 μm wide at the sides, thick, comprising 3–4 layers, outer most layer heavily pigmented,
thick-walled, comprising blackish to dark brown loosely
packed cells of textura angularis, inner layer composed 3–5
layers, pale brown to hyaline, cells towards the inside lighter,
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Fungal Diversity (2022) 117:1–272
Fig. 19 Ascochyta medicaginicola (MFLU 17–2138, new host record) a-c Conidia observed on host substrate. d-e Conidiomata. f–h Conidia i,
k-l Conidia j Conidiodenous cell. Scale bars: a = 500 µm, b-e, g = 100 µm, f, h = 50 µm, i-l = 10 µm
flattened, thick-walled cells of textura angularis. Sexual
morph: Not observed.
13
Material examined: Russia, Rostov region, Shakhty,
near a railroad, on dead twigs of Prunus cerasifera Ehrh.
Fungal Diversity (2022) 117:1–272
31
(Rosaceae), 11 May 2017, Timur S. Bulgakov, T-1832
(MFLU 17–2138); living culture MFLUCC 18–0453.
Hosts: Medicago albus, Medicago sativa, Medicago
sp. (Fabaceae, Hyde et al. 2020), Prunus cerasifera
(Rosaceae, this study), Scabiosa sp. (Caprifoliaceae, Tibpromma et al. 2017) and Trichosanthes dioica (Cucurbitaceae, Sarkar et al. 2018-pathogenicity data are available).
Distribution: Canada, Czech Republic, France, Italy, USA
(Hyde et al. 2020a, b, c), India (Sarkar et al. 2018), Thailand
(this study)
GenBank number: OM235096 (ITS)
Notes: Our collection shares similar morphological
characteristics with the ex-type strain of A. medicaginicola
(Boerema et al. 2004; Chen et al. 2013). The multigene
Fig. 20 Phylogram generated from maximum likelihood analysis
based on combined, ITS, LSU and tub2 sequence data of selected
taxa. Related sequences were obtained from GenBank. Forty-one
strains are included in the analyses, which comprise 633 characters
including gaps. The tree was rooted with Phoma herbarum (CBS
377.92 and CBS 502.91). The maximum likelihood bootstrap (ML)
values > 65%) are given above the nodes. The new isolate in red bold
13
32
phylogenetic analysis shows that our specimen groups in
the Ascochyta medicaginicola clade with 96/0.90 ML/
BYPP support (Fig. 20). Four Ascochyta species have been
recorded based on the morphological description from
Rosaceae plants in Russia (Melnik 2000; Farr and Rossman
2022): Ascochyta idaei on Rubus idaeus in the Leningrad
region, Kursk region, and Stavropol region; A. potentillarum
on Potentilla reptans in Arkhangelsk region (Melnik 2000),
Lipetsk region (Sarycheva et al. 2009), Republic of Crimea
(Ovcharenko 2011) and Voronezh region (Melkumov 2015);
A. pruni on Prunus padus in Leningrad region; Ascochyta
sorbina on Sorbus torminalis in Stavropol region. As these
species were identified based on morphology alone, correct species identification is yet to be done. Our collection
provides the first host record of Ascochyta medicaginicola
on Rosaceae based on both morphological and phylogenetic
data.
Didymosphaeriaceae Munk, Dansk bot. Ark. 15(no. 2): 128
(1953).
Didymosphaeriaceae represents an important family in
Dothideomycetes. The family is typified by Didymosphaeria, with D. epidermidis as the type species (Hongsanan
et al. 2020b). While taxa of Didymosphaeriaceae are often
endophytic, pathogenic or saprobic on various plant hosts
(Gonçalves et al. 2019; Hongsanan et al. 2020b), they can
sometimes also be pathogenic to human beings (Hongsanan
et al. 2020b). Species of Didymosphaeriaceae are mainly
characterised by brown, 1–3-septate or muriform ascospores
and cellular or trabeculate pseudoparaphyses in their sexual
morphs while their asexual morphs are fusicladium-like
or phoma-like (Hyde et al. 2013; Hongsanan et al. 2020b).
After several taxonomic revisions, 32 genera have been
accepted in the family (Hongsanan et al. 2020b).
Spegazzinia Sacc., Spegazzinia: [1] (1879)
Spegazzinia was introduced by Saccardo (1880), with S.
ornata as the type species and it currently comprises hyphomycetous taxa. The genus was initially accommodated in
Apiosporaceae (Sordariomycetes) based on morphology
(Hyde et al. 1998). It was then transferred to Didymosphaeriaceae (Dothideomycetes) based on molecular evidence
(Tanaka et al. 2015). Taxa in this genus are mainly characterised by a distinctive conidiophore ontogeny as well as
two types of conidia (Samarakoon et al. 2020; Hongsanan
et al. 2020b). The latest two taxa added to Spegazzinia are S.
musae, reported as a saprobe on Musa sp., and S. camelliae,
isolated as an endophyte from Camellia sinensis var. assamica (Samarakoon et al. 2020; Suwannarach et al. 2021).
Spegazzinia deightonii (S. Hughes) Subram., J. Indian bot.
Soc. 35: 78 (1956).
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Fungal Diversity (2022) 117:1–272
Index Fungorum Number: IF306062; Facesoffungi number: FoF07238; Fig. 21
Saprobic on palm stem. Asexual morph: Hyphomycetous. Sporodochia powder-like, dark, dense, dry,
1–3.5 mm in diameter. Conidiophores 65–120 × 1–3 μm
(x̅ = 93.5 × 2.4 μm, n = 15), macronematous, micronematous, narrow, subspherical to doliiform, flexuous or erect,
unbranched, hyaline to pale brown, verruculose. Conidiogenous cells 10–20 × 2–4 μm (x̅ = 15.8 × 3 μm, n = 15),
basauxic, terminal, erect, unbranched, hyaline to pale
brown, verruculose, each producing a single, holoblastic
conidium at the conidiophore apex. Conidia two types,
disc-like and stellate; disc-like conidia 20–28 × 17–19 μm
(x̅ = 25.2 × 18.1 μm, n = 20), usually 8-celled, solitary, hyaline when immature, pale to dark brown on maturity, crossseptate, slightly constricted at the septa, with short and blunt
spines at the periphery, frequently accompanied by attached
conidiogenous cells post splitting from the conidiophores;
stellate conidia 18–27 × 16–29 μm (x̅ = 22.6 × 24.3 μm,
n = 20), globose or variously shaped, frequently 4- to
6-celled, solitary, septate, deeply constricted at the septa,
pale to dark brown, comprising spines 4–5 μm long. Sexual
morph: Not observed.
Culture characteristics: Conidia germinating on PDA
within 16–18 h. Colonies growing on PDA, reaching a diameter of 55 mm after 14 days at 25 °C, aerial, moderately
dense, undulate margine, middle grey, periphery olive green
at immature stage and brownish gray at maturity; reverse
greyish white to white.
Material examined: Thailand, Chiang Rai, on dead stem
of palm (Arecaceae), 20 December 2019, Binu C. Samarakoon, E003 MFLU 22-0277), living culture MFLUCC
22-0180.
Hosts: Spegazzinia deightonii has been reported from
several hosts, including, Andropogon glomeratus (Arnold
1986), Areca catechu (Tianyu 2009; Matsushima 1980),
Arundo donax (Tanaka et al. 2011), Bambusa vulgaris
(Camino-Vilaro et al. 2019), Calathea makoyana (Tianyu
2009), Cocos nucifera (Tianyu 2009), Imperata cylindrical
(Thaung 2008), Musa sp. (Samarakoon et al. 2018), Panicum
maximum (Lu et al. 2000; Wong and Hyde 2003), Phoenix
hanceana (Tianyu 2009), Quercus xalapensis (Heredia et al.
1995), Saccharum spontaneum (Mel'nik et al. 2000), Thysanolaena latifolia (Mel'nik et al. 2000) and Tillandsia sp.
(Delgado-Rodriguez et al. 2002).
Distribution: China (Tianyu 2009), Cuba (Arnold 1986;
Delgado-Rodriguez et al. 2002; Camino-Vilaro et al. 2019),
Hong Kong (Lu et al. 2000; Wong and Hyde 2003), Japan
(Tanaka et al. 2011), Mexico (Heredia et al. 1995), Myanmar
(Thaung 2008), Philippines (Whitton et al. 2012), Taiwan
(Matsushima 1980), Thailand (Samarakoon et al. 2020),
United States (Delgado 2008), Vietnam (Mel'nik et al. 2000).
Fungal Diversity (2022) 117:1–272
33
Fig. 21 Spegazzinia deightonii
(MFLU 22-0277, new host
record) a Host. b Close-up
of conidia on host. c Mass of
conidia. d Conidiogenous cell
of stellate conidia. e Stellate
conidium on a conidiophore. f
Stellated conidium. g Disk-liked
conidium. h Disk-like conidium
with attached conidiogenous
cell. Scale bars: c–h = 10 μm
GenBank numbers: ON885254 (SSU); ON873996(LSU);
ON873998(ITS); ON885741 (tef1)
Notes: Our isolate (MFLUCC 22-0180) clusters with
other strains of Spegazzinia deightonii with 95% ML, statistical support in the multi-loci phylogenetic tree (Fig. 22).
Pairwise comparison of DNA sequence data shows insignificant differences among our strain (MFLUCC 22-0180) and
the other strains of S. deightonii, following which our isolate
is considered as S. deightonii. Furthermore, the isolate in
the present study shares similar features with other strains
of S. deightonii, including two types of pale to dark brown
conidia which are multi-cellular, constricted at the septa and
comprise either long (stellate conidia) or short (disc-like
conidia) spines (Ellis 1961; Tanaka et al. 2011; Samarakoon
et al. 2018). Small differences in sizes may be accounted for
by host variations. Spegazzinia deightonii has been reported
on a palm substrate in China, Taiwan and the United States
(Matsushima 1980; Delgado 2008; Tianyu 2009). We recovered the species from Thailand and thus report it as a new
record on Palm from Thailand. Recently this species was
also reported to occur in Thailand, in the same Province and
area, on Musa sp. (Samarakoon et al. 2020).
Lindgomycetaceae K. Hiray., Kaz. Tanaka & Shearer, in
Hirayama et al., Mycologia 102(3): 733 (2010)
Notes: Lindgomycetaceae was introduced by Hirayama et al. (2010) with Lindgomyces (L. ingoldianus) as
the generic type. Most Lindgomycetaceae members have
been recorded from freshwater habitats and the freshwater
isolates form a distinct lineage from other Dothideomycetes taxa based on ribosomal sequence data (Zhang et al.
2013; Hyde et al. 2013; Dong et al. 2021). There are seven
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34
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Fungal Diversity (2022) 117:1–272
◂ Fig. 22 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, ITS and tef sequence data of selected
taxa in Didymosphaeriaceae. Bootstrap support values for ML ≥ 65%
and BYPP ≥ 0.95 are indicated above or below the branches. Ex-type
strains are in bold. The new isolate is in blue. The tree is rooted with
Laburnicola zaaminensis (TASM 6152) and L. murtfirmis (MFLUCC
16-0290).
genera in the family, including Hongkongmyces which is
also associated with human diseases (Tsang et al. 2014;
Linqiang et al. 2020). The genus forms a close relationship to Aquimassariosphaeria, Clohesyomyces, Lolia, and
Trematosphaeria in the phylogenetic analyses of the ribosomal DNA and protein-coding genes such as tef (AbdelAziz and Abdel-Wahab 2010; Hyde et al. 2016). We,
therefore, encourage further taxonomic studies to include
protein-coding regions for a better resolution of taxa in
Lindgomycetaceae (Fig. 24).
Hongkongmyces Tsang et al., Medical Mycol. 52(7): 740
(2014)
Hongkongmyces was introduced as a monotypic genus
associated with human infections, and typified by H. pedis
(Tsang et al. 2014). Six species H. aquaticus, H. brunneosporus, H. kokensis, H. pedis, H. snookiorum, and H. thailandicus are listed in Species Fungorum (2022a, b). The
species are commonly associated with freshwater habitats
as saprobes. Hyde et al. (2016) introduced the second species of Hongkongmyces based on sexual morph characters,
which was collected on submerged wood in a stream in Thailand. Later, an asexual morph of H. snookiorum was isolated
from submerged wood (Crous et al. 2018). Hongkongmyces snookiorum was reported as an opportunistic fungal
infection in a transplant patient (Deng et al. 2020). Schoch
et al. (2012) and Aime et al. (2021) recommend using the
fungal barcode locus such as the ITS region as well as any
additional secondary barcode locus such as protein-coding
region for the introduction of new species. Most Hongkongmyces species lack barcodes in public databases, and therefore, delineating species in the genus is a challenging task.
In this paper, we introduce a new species of Hongkongmyces
which was collected in a freshwater pond in Jilin Province of
China based on comprehensive morphology and phylogeny
analyses (Figs. 23 and 24).
Hongkongmyces changchunensis Phukhams., W.X. Su, &
Y. Li, sp. nov.
Index Fungorum number: IF559493; Facesoffungi number: FoF10725; Fig. 23
Etymology: The epithet reflects the locality, Changchun.
Holotype: HMJAU 60185.
Saprobic on decaying Betula twigs submerged in a
freshwater stream. Sexual morph: Not observed. Asexual
morph: Conidiomata 77–188 × 85–174 μm diam, pycnidia
35
scattered, semi-immersed, globose or ellipsoidal, black,
coriaceous, ostiolate. Peridium 10–28 μm thick, composed of large, irregular, and dark brown cells arranged in
a textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2.6–23.5 × 1.6–4.9 μm
(x̅ = 10 × 3 μm, n = 20), enteroblastic, phialidic, determinate,
cylindrical to subcylindrical, hyaline, thin-walled. Conidia
10–18 × 7–13 μm (x̅ = 13 × 10 μm, n = 43), ellipsoidal, obovoid or irregular, aseptate, guttulate, hyaline, thin-walled,
lacking mucilaginous sheath, turning brown at senescence
stage.
Culture Characteristics: Colonies on PDA reaching
20 mm in 7 days at 20 °C, flat with entire margin, circular,
grey to black, aerial mycelium becoming grey towards the
edge; reverse grey-olivaceous to black, smooth.
Material examined: China, Jilin Province, Changchun
District, Jingyuetan National Scenic Areas, on the submerged twigs in a stream of Betula sp. (Betulaceae), 12 May
2021, Chayanard Phukhamsakda (SWX32), (HMJAU 60185,
holotype); ex-type living culture, CCMJ5008.
GenBank numbers: OL897173 (LSU), OL891809(SSU),
OL996122 (ITS), OL944603 (tef1), OL944507(rpb2).
Notes: Phylogenetic analyses of LSU, SSU, ITS and tef
sequence data show that Hongkongmyces changchunensis
(CCMJ5008) is related to the type species Hongkongmyces
thailandicus (MFLUCC 16–0406), with good support (92%
ML/0.93 BYPP; Fig. 23). Hongkongmyces changchunensis
was collected from submerged substrates in a freshwater
habitat in Jilin Province of China. Both species were collected from fresh water habitats, but from different country (Hyde et al. 2016). Hongkongmyces changchunensis is
similar to Hongkongmyces aquaticus in having phialidic
conidiogenous cells and hyaline, variable shaped conidia,
but it lacks sympodial proliferations (Dong et al. 2020).
Hongkongmyces changchunensis is characterized by wet,
globular, mass of conidia on natural substrates after drying and smooth surfaced conidia containing multiple small
bubbles. Herein, we propose a new species based on both
morphology and phylogenetic analyses.
Lindgomyces K. Hiray., Kaz. Tanaka & Shearer, in Hirayama et al., Mycologia 102(3): 733 (2010)
Notes: Lindgomyces was introduced in Lindgomycetaceae by Hirayama et al. (2010) based on the type species L.
ingoldianus (Shearer & K.D. Hyde) K. Hiray., Kaz. Tanaka
& Shearer. The morphology of the species of Lindgomyces
is similar to Massarina eburnea and Lophiostoma macrostomum, but they differ from ascomata and ascospores (Raja
et al. 2013; Hirayama et al. 2010). There are 14 epithets
listed in Lindgomyces in Index Fungorum (2022a, b). Keys
to the species of Lindgomyces were provided by Dong et al.
(2020). In this study, we describe a novel species, Lindgomyces guizhouensis based on the unique morphological features
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Fungal Diversity (2022) 117:1–272
Fig. 23 Hongkongmyces changchunensis (HMJAU 60185, holotype) a, b Appearance of conidiomata on host surface. c Vertical
section through conidioma. d Ostiole canal. e Section of partial con-
idioma peridium. f–h Conidiogenous cells and developing conidia.
i–l Developmental state of conidia. Scale bars: a = 500, b = 200 µm,
c = 100 µm, d, e = 50, f, l = 20 µm, g–k = 10 µm
and multi–gene phylogenetic analysis of a combined LSU
and ITS sequence data (Fig. 26).
erumpent, subglobose to broadly conical, with a centrally
located ostiole. Peridium 44–79 μm thick, thick–walled,
composed of pale brown cells of textura angularis.
Hamathecium 1–2 μm wide, septate, branched, pseudoparaphyses above the asci. Asci 110–154 × 15–28 μm, bitunicate, fissitunicate, apically rounded with an indistinct
ocular chamber, subsessile, broadly cylindrical to clavate.
Ascospores 30–51 × 8–12 μm, (x̅ = 44.7 × 10.5 μm, n = 25),
obliquely uniseriate to 2–3–seriate, fusiform, slightly curved
with narrowly rounded ends, 1–septate, 2–4 guttules in each
cell, wall gray while in the ascus, smooth or slightly verruculose. Sexual morph: Not observed.
Lindgomyces guizhouensis J. Mai, Y.Z. Lu & K.D. Hyde,
sp. nov.
Index Fungorum number: IF559506; Facesoffungi number: FoF 10676; Fig. 25
Etymology: Referring to collecting site in Guizhou Province, China.
Holotypus: GZAAS 21–0383
Saprobic on decaying wood in terrestrial habitats. Asexual morph: Ascomata 396–548 μm high × 466–514 μm
diam., superficial, black, smooth, scattered, solitary,
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37
Fig. 24 Phylogram generated from Bayesian 50% majority-rule
consensus phylogram based on combined LSU, SSU, ITS and tef
sequence data of Lindgomycetaceae. The topology and clade stability
of the combined gene analyses was compared to the single gene anal-
yses. The tree is rooted with members of the Aigialaceae. Bootstrap
values ≥ 50% (MP and ML) and BYPP ≥ 0.90 are given at the nodes.
The type-delivered strains are in bold; the new isolates are in blue
Culture characteristics: Colonies on PDA reaching
43 mm in 20 days at 25 °C, flat, filiform, gray to near–black
from center to edge, with moderate aerial mycelium, smooth,
irregular; In reverse, gray to pale brown.
Material examined: China, Guizhou Province, Longli,
on decaying wood submerged in a freshwater stream, 2
September 2020, Jian Ma, LLSB06 (GZAAS 21–0383,
holotype); ex–type living culture, GZCC 21–0669.
GenBank numbers: OM339435 (ITS), OM339432 (LSU).
Notes: In a BLASTn search of NCBI GenBank, the closest match of LSU and ITS sequence data for Lindgomyces
guizhouensis is 99.54% and 96.96% similar to L. pseudomadisonensis (KT 2742). The multi–loci phylogenetic
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Fig. 25 Lindgomyces guizhouensis (GZAAS 21–0383, holotype). a–c Superficial ascomata
on unidentified plant substrate.
d Vertical section of an ascoma.
e Peridium. f Hamathecium g
Asci with hamathecium h–i
Asci j–n Ascospores o Germinating ascospore p–q Colony
on PDA from above and below.
Scale bars: d, e = 100 μm,
f–o = 20 μm
analyses of the combined LSU and ITS sequence dataset
confirmed the new strain obtained belonging to Lindgomyces, where it is sister to L. griseosporus with 99% ML and
1.00 BYPP support (Fig. 25). Morphologically, L. guizhouensis is also most similar to L. griseosporus in having
superficial, black with roughened surface ascomata; a thick
13
peridium; bitunicate, clavate, subsessile asci and fusiform,
1-septate, guttules ascospores. However, our new collection differs from L. griseosporus in having larger ascomata (396–548 × 466–514 μm vs 240–290 × 320–350 μm),
fragile peridium and smaller asci (110–154 × 15–28 μm vs
Fungal Diversity (2022) 117:1–272
39
Fig. 26 Phylogram generated
from maximum likelihood analysis based on combined LSU
and ITS sequence data. Twenty–
two taxa were included in the
combined analyses, which comprised 2096 characters (LSU:
841, ITS: 1255) after alignment.
Bootstrap support values for
ML ≥ 50% and BYPP ≥ 0.95
are given above the nodes.
Hongkongmyces aquaticus
(MFLUCC 18–1150), H. pedis
(HKU35) and H. thailandica
(MFLUCC 16–0406) were used
as the outgroup taxa. The newly
generated sequence is indicated
in blue. The ex–type strains are
indicated in bold
140–180 × 24–30). Therefore, we introduce Lindgomyces
guizhouensis as a new species.
Lophiostomataceae Sacc.
Lophiostomataceae, typified by Lophiostoma, was first
erected by Nitschke (1869) as ″Lophiostomeae″, but it was
established by Saccardo (1883) as ″Lophiostomaceae″
and placed in Pleosporales. Lophiostomataceae species are
mostly characterized by slot-like ostiole on apex of a flattened neck that are usually saprobes that grow on herbaceous and woody plants from terrestrial and aquatic environments (Holm and Holm 1988; Mugambi and Huhndorf
2009; Thambugala et al. 2015; Hashimoto et al. 2018).
The historical account of Lophiostomataceae and recent
generic notes of Lophiostomataceae were provided by Hyde
et al. (2013) and Hongsanan et al. (2020a, b), respectively.
Wijayawardene et al. (2022) outlined and accepted 30 genera
of Lophiostomataceae with Lophiostoma as speciose genera
(ca. 100). Calabon et al. (2022) listed 20 species under 10
genera of Lophiostomataceae from freshwater environments.
In this series, we introduced two new species of Lophiostomataceae from terrestrial and freshwater habitats.
Neovaginatispora A. Hashim., K. Hiray. & Kaz. Tanaka.
Notes: Neovaginatispora, typified by N. fuckelii, was
introduced by Hashimoto et al. (2018) based on their
phylogenetic analysis and morphological differences of
peridium (i.e., thinner peridium that is uniformly thick and
composed of two cell layers) compared to Vaginatispora
13
40
(Thambugala et al. 2015). Recent phylogenetic analysis
shows that Neovaginatispora strains form a separate subclade with Lentistoma and Vaginatispora (Bao et al. 2019;
Phukhamsakda et al. 2020; Hyde et al. 2020a, b, c). Two
Neovaginatispora species are accepted, N. fuckelii and N.
clematidis. Neovaginatispora fuckelii has a cosmopolitan distribution that thrives on various hosts in terrestrial
habitats (Wang and Lin 2004; Thambugala et al. 2015;
Hyde et al. 2016; Hyde et al. 2020a, b, c) while Bao et al.
(2019) reported this on freshwater habitats. The second
species, N. clematidis, was introduced by Phukhamsakda
et al. (2020) from the dead stems of Clematis viticella
in Belgium. Neovaginatispora clematidis differs from N.
Fig. 27 Neovaginatispora
mangiferae (MFLU 18–0069,
holotype). a Appearance of
ascomata in host substrate. b
Vertical section of ascoma. c
Peridium. d Pseudoparaphyses. e Ostiole with numerous
periphysis. f, g Bitunicate asci.
h–k Ascospores in different
stages of maturity. Scale bars:
b = 100 µm, c, d, h–k = 5 µm, e,
f, g = 20 µm
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fuckelii on their ascospore morphology (broad fusiform
with a single eusepta). In this study, the third Neovaginatispora species is introduced based on the collection
from dead stems of Mangifera indica in Taiwan.
Neovaginatispora mangiferae Tennakoon, M.S. Calabon,
E.B.G. Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF559843; Facesoffungi number: FoF12720; Fig. 27
Etymology: Name reflects the host Mangifera indica,
from which the holotype was collected.
Holotype: MFLU 18–0069
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Saprobic on decaying stem of Mangifera indica. Sexual
morph: Ascomata 430–500 μm high, 320–350 μm diam.,
scattered, semi-immersed to immersed, papilla erumpent
through host surface, coriaceous to carbonaceous, dark
brown to black, globose to subglobose, ostiolate. Ostiole
crest-like, variable in shape, central, periphysate, broadly
papillate, with an irregular pore-like opening. Peridium
25–40 μm wide, thick, composing two layers of irregular
cells arranged in a textura angularis, outer layer with darker,
light brown to brown flattened cells, inner layer comprising several layers of hyaline cells. Hamathecium 1.5–3 μm
wide, massive, long cylindrical cellular, anastomosed, cellular pseudoparaphyses, hyaline, septate with small guttules.
Asci 60–80 × 7–9 μm (x̅ = 60.9 × 7.2 μm, n = 20), 8-spored,
bitunicate, cylindrical to cylindric-clavate, short pedicellate with furcate to obtuse ends, apically rounded with an
indistinct ocular chamber. Ascospores 13–17 × 3.0–5.5 μm
(x̅ = 15.1 × 3.7 μm, n = 25), biseriate, overlapping, fusiform
to sunbfusoid, strongly constricted at the median septum,
straight or slightly curved, 1-septate at the center, enlarged
near the septum at the upper cell, hyaline, guttulate, smoothwalled, mostly with 4 guttules, with a helmet-shaped to subcylindrical mucilaginous sheath at each end when immature,
invisible at maturity, 2.1–5.6 μm long, 2.2–4.4 μm wide.
Asexual morph: Not observed.
Culture characteristics: Conidia germinating on potato
dextrose agar (PDA) within 24 h. Germ tubes produced from
the apical cell of conidia. Colonies growing on PDA, reaching 25–30 mm in 2 weeks at 25 °C. Mycelia superficial,
medium dense, irregular, flat, slightly raised, surface smooth
with crenate edge, fluffy to velvety with smooth aspects,
zonate with different sector yellowish-brown to moss brown
at the margin brownish-grey at the center; from below, light
moss brown at margin, dark gray at the middle, brown at the
center, no pigmentation and sporulation.
Material examined: Taiwan, Chiayi, Fanlu Township
area, Dahu village, dead stems of Mangifera indica (Anacardiaceae), 5 August 2017, D.S. Tennakoon, DTW 018C
(18–0069, holotype), ex-type living culture, MFLUCC
17–2652.
GenBank numbers: MG931027(LSU), MG931030 (SSU),
MG931033 (ITS).
Notes: The isolate MFLUCC 17–2652 has formerly
identified as Neovaginatispora fuckelii by Tennakoon
et al. (2018) but in our phylogenetic analysis, it clustered
with Neovaginatispora sp. MFLUCC 11–0577, and forms
a separate subclade with N. fuckelii and N. clematidis
with 100% MP, 1.00 BYPP support (Fig. 28). The pairwise nucleotide comparison of LSU and ITS of N. mangiferae MFLUCC 17–2652 show 2 bp (0/24%, 838 bp) and
1 bp (0.23%, 442 bp) differences with Neovaginatispora
sp. MFLUCC 11–0577, respectively, with no differences
on SSU sequence data. For this reason, we named the
41
unidentified Neovaginatispora species (strain MFLUCC
11–0577) as another strain of N. mangiferae. In pairwise
nucleotide comparisons of Neovaginatispora mangiferae
MFLUCC 17–2652 with N. fuckelii MFLUCC 17–1334,
there is a nucleotide difference of 6.94% (30/432 bp) in
ITS, and 2.71% (22/813 bp) in LSU genes. A pairwise
nucleotide comparisons of LSU, SSU, and ITS sequence
data of N. mangiferae MFLUCC 17–2652 and N. clematidis reveals 22 bp (2.72%, 810 bp), 6 bp (0.64%, 938 bp),
and 27 bp (7.03%, 384 bp) differences, respectively. Neovaginatispora mangiferae differs from N. fuckelii KT 634
by having larger ascomata (430–500 × 320–350 μm vs
150–180 × 200–250 μm) and thicker peridium (25–40 μm
vs 15–25 μm), and periphyses in ostiole. The asci and
ascospores are similar in size but ascospores of N. mangiferae is fusiform with acute ends but N. fuckelii has a fusiform ascospores with obtuse ends (Thambugala et al. 2015;
Tennakoon et al. 2018; Bao et al. 2019). Neovaginatispora
mangiferae has larger ascomata (430–500 × 320–350 μm
vs 145–250 × 108–160 μm), shorter asci (60–80 × 7–9 μm
vs. 53–105 × 9–12 µm), and smaller ascospores
(13–17 × 3.0–5.5 μm vs. 16–19 × 5–7 µm) compared to N.
clematidis (Phukhamsakda et al. 2020).
Vaginatispora K.D. Hyde, amended
Saprobic on submerged wood, intertidal wood, dead
twigs, endocarp or fallen fruit pericarp. Sexual morph:
Ascomata solitary or scattered, immersed to erumpent, uniloculate, subglobose, glabrous, dark brown to black. Ostiolar
neck slit-like, elongate, laterally compressed, composed of
globose to elongate, brown to black cells, with a pore-like
opening and hyaline periphyses. Peridium unequal in thickness, two-layered, outer layer comprising somewhat flattened
cells, fusing and indistinguishable from the host tissues,
inner layer comprising lightly pigmented to hyaline cells.
Pseudoparaphyses numerous, cellular, hypha-like, hyaline,
septate, anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical to clavate, with a short or long
pedicel, apically round with an ocular chamber. Ascospores
uni- to bi-seriate, narrowly ellipsoidal or fusiform, straight
to slightly curved, hyaline when immature, becoming yellow when mature, 1-septate, occasionally producing pseudosepta, septum mostly median, upper cell slightly broader
than lower cell, smooth, thin-walled, with or without bipolar appendages or entire sheath. Asexual morph: Hyphomycetous. Hyphae septate, hyaline to lightly pigmented,
mostly smooth, thick-walled, moniliform. Chlamydospores
numerous, mostly in chains, globose to subglobose, smooth,
initially hyaline then lightly pigmented at maturity, arising
from the mycelium, formed intercalarily or terminally.
Type species: Vaginatispora aquatica K.D. Hyde
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◂ Fig. 28 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, ITS, tef1, and rpb2 sequence data
representing Lophiostomataceae (Pleosporales). Eighty-one strains
are included in the combined analyses which comprised 3726 characters (741 characters for LSU, 973 characters for SSU, 526 characters for ITS, 1001 characters for tef1, and 1011 characters for rpb2)
after alignment. Angustimassarina acerina (MFLUCC 14–0505) and
Angustimassarina populi (MFLUCC 13–0034) in Amorosiaceae (Pleosporales) were used as the outgroup taxa. Bootstrap support values
for ML ≥ 75% are given above the nodes (left side) and BYPP ≥ 0.95
are given above the nodes (right side). Ex-type strains are in bold and
newly generated sequences are in blue
Notes: Hyde (1995) introduced Vaginatispora to accommodate Vaginatispora aquatica K.D. Hyde (= Lophiostoma
vaginatispora Huang Zhang & K.D. Hyde), which was previously placed in Massarinaceae. Eight species are included
in this genus and members are characterized by depressed
globose ascomata, immersed beneath a blackened neck, with
a slot-like ostiole, numerous filamentous pseudoparaphyses,
cylindrical to clavate asci and narrowly ellipsoidal, hyaline,
1-septate ascospores with a mucilaginous collar around its
equator, having large guttules in each cell, and a spreading
papilionaceous sheath (Thambugala et al. 2015; Hashimoto
et al. 2018). We followed the treatment of Dong et al. (2020)
and Hongsanan et al. (2020a, b) in this genus.
Vaginatispora lignicola M.S. Calabon, E.B.G. Jones & K.D.
Hyde, sp. nov.
Index Fungorum number: IF559844; Facesoffungi number: FoF12721; Fig. 29
Etymology: Referring to this taxon dwelling on wood
Holotype: MFLU 22–0116
Saprobic on decaying wood submerged in freshwater
habitats. Sexual morph: Ascomata 200–360 × 220–415 µm
diam. (x̅ = 269 × 294 µm; n = 10), scattered to gregarious, immersed to semi-immersed, erumpent at maturity, coriaceous, black, subglobose, ostiolate. Ostiole
slot-like, central, elongated, pore-like opening, plugged
by hyaline, filamentous hyphae, periphysate. Peridium
40–65 µm wide, circular, symmetric, outermost layer
heavily pigmented, comprising a blackish to dark brown
amorphous layer, flattened and loosely packed cells of
textura angularis, inner layer composed of hyaline cell
layers, flattened, thick-walled cells of textura angularis.
Hamathecium 1.3–2.5 µm wide (x̅ = 1.9 µm, n = 30) comprising numerous, filamentous, branched, septate, pseudoparaphyses. Asci 125–145 × 15–30 µm (x̅ = 137 × 23 µm,
n = 20), 8-spored, bitunicate, fissitunicate, cylindricalclavate, short-pedicellate, apex rounded with a minute ocular chamber. Ascospores 50–72 × 14–17 µm
(x̅ = 62 × 16 µm, n = 30), uniseriate to bi-seriate, overlapping, straight or slightly-curved, initially hyaline, becoming yellowish at maturity, fusiform, mostly with narrow
acute ends, 1–3-septate, strongly constricted at the septa,
43
smooth-walled, with numerous small guttules and 2 distinct large guttules. Asexual morph: Hyphae 2–5 μm
wide, septate, hyaline to lightly pigmented, mostly smooth,
thick-walled, moniliform. Chlamydospores 6–19 × 4.5–16
(x̅ = 10.9 × 8.4, n = 50), numerous, mostly in short chains,
globose to subglobose, smooth, initially hyaline then
becoming light brown at maturity, arising from the mycelium, formed intercalarily or terminally.
Culture characteristics: Conidia germinating on malt
extract agar (MEA) within 24 h. Germ tubes produced
from the basal and apical cell of conidia. Colonies growing
on MEA, reaching 20–25 mm in 2 weeks at 25 °C. Mycelia superficial, circular, with entire margin, flat, smooth,
from above greyish brown to grey, from below dark grey
to black.
Material examined: Thailand, Tak Province, Tha Sing
Yang, Ban Mae Ja Wang on decaying wood submerged in
a freshwater river, 17 October 2019, N. Padaruth, CC43
(MFLU 22–0116, holotype), ex-type living culture,
MFLUCC 22–0078.
GenBank numbers: MW287233 (LSU),
MW287229(SSU), MW260329(ITS), MW512605(tef1),
OP251197(rpb2)
Notes: Three species of Vaginatispora are recorded in
freshwater habitats: V. aquatica (Hyde 1995; Tsui et al.
2000; Zhang et al. 2014a, b), V. armatispora (Hu et al. 2010;
Bao et al. 2019; Hyde et al. 2019), and V. nypae (Hyde et al.
2020a, b, c; Boonmee et al. 2021). In this series, we introduce another novel species of Vaginatispora, V. lignicola,
from a freshwater habitat in Thailand. Vaginatispora lignicola (MFLUCC 22–0078) confirms with the generic ascomatal morphology (erumpent, uniloculate, glabrous, slotlike ostiole, with numerous hyaline periphyses) but differs
from the known members of the genus in having ascospores
without a wide papilionaceous sheath or distinct hyaline
appendages at both ends. Based on the phylogenetic analysis
of combined LSU, SSU, ITS, tef1, and rpb2 sequence data,
V. lignicola is basal to V. aquatica with high bootstrap support (97 ML and 1.00 BYPP) (Fig. 28). The former differs
from V. aquatica in the measurement (50–72 × 14–17 µm
vs 36–48 × 11–16) and color of the ascospores (initially
hyaline then becoming yellowish at maturity vs hyaline),
and the absence of a sheath when stained in Indian ink
wherein the latter have wide papilionaceous sheaths (Hyde
1995). The novel taxon differs from the freshwater strains
of V. armatispora and V. nypae by having larger ascospores
(50–72 × 14–17 µm vs 22–30 × 5.5–8 µm vs 26–29 × 6–7 µm)
without a mucilaginous sheath and hyaline appendages at
both ends (Bao et al. 2019; Hyde et al. 2019; Boonmee
et al. 2021). Multi-locus phylogenetic analyses showed
that V. lignicola is a distinct taxon in Vaginatispora with
49 bp (9.32%, 526 bp) and 14 bp (1.74%, 804) nucleotide
differences in the ITS and LSU sequence data of the type
13
44
Fig. 29 Vaginatispora lignicola (MFLU 22–0116, holotype). a
Host twig. b, c Appearance of erumpent ascomata in host substrate. d Vertical section of ascoma. e Slot-like ostiole with numerous periphysis. f Peridium. g Pseudoparaphyses. h Bitunicate asci.
i Ocular chamber. j–n Ascospores in different stages of maturity. o
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Germinated ascospore. p Culture on MEA. q–s Hyphae. (t) Terminally and (u) intercalary swollen cells. v–x Moniliform hyphae with
constricted septa. y–ab Multicellular bodies. Scale bars: a = 2 mm,
b–d = 200 µm, e, g = 20 µm, f = 50 µm, h, j = 100 µm, k–o, y,
z = 50 µm, q–x, aa = 20 µm
Fungal Diversity (2022) 117:1–272
species, V. aquatica, respectively. The present work is the
first report also of the morphological characteristics of the
asexual morph of Vaginatispora. A simplified key to species
of Vaginatispora is provided herein.
Key to species of Vaginatispora
1 Ascospores yellow at maturity, without sheath or
appendages……………………………………V. lignicola
1 Ascospores hyaline, with either sheath or appendages……………………………………………………… 2
2 Ascospores with wide papilionaceous sheath, lacking
appendages………………………………………V. aquatica
2 Ascospores with or without a mucilaginous sheath, with
distinct hyaline appendages………………………………3
3 Ascospores < 25 μm long………………V. scabrispora
3 Ascospores > 25 μm long……………………………4
4 Ascomata lack slit-like ostiole………………V. nypae
4 Ascomata with slit-like opening……………………5
5 Appendages 2–8 μm long…………V. microarmatispora
5 Appendages 6–8 μm long……………………………6
6 Ascomata > 450 μm wide, peridium up to > 40 μm
thick……………………………………………V. amygdali
6 Ascomata < 450 μm wide, peridium up to < 40 μm
thick………………………………………………………7
7 Ascospores up to < 40 μm long…………V. armatispora
7 Ascospores up to > 40 μm long………………………8
8 Ascospores 40–45 × 10–15 μm………………………
……………………………………………V. appendiculata
8 Ascospores 23–45 × 6–9 μm…………………V. palmae
Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde,
Mycosphere 9(2): 323 (2018).
Notes: Phaeoseptaceae was introduced by Hyde et al.
(2018) to accommodate Phaeoseptum, Lignosphaeria
and Neolophiostoma. The LSU-SSU-rpb2-tef1 multigene
phylogeny in Hyde et al. (2018) showed a well-supported
Phaeoseptaceae clade sister to Halotthiaceae. The species
in Phaeoseptaceae share immersed or erumpent ascomata,
dark brown to black outer peridium, cylindrical, branched,
septate pseudoparaphyses, 8-spored, bitunicate, cylindrical-clavate, long pedicellate asci, and brown, muriform
ascospores. Based on morphology and phylogeny, Thambugala et al. (2015) treated Lignosphaeria in Dothideomycetes,
genera incertae sedis, and Liu et al. (2019a, b, c, d) treated
Neolophiostoma in Halotthiaceae. In a recent revision of
Dothideomycetes, Hongsanan et al. (2020b) accepted only
Phaeoseptum and Pleopunctum in Phaeoseptaceae which
was followed by Wijayawardena et al. (2020, 2022).
Phaeoseptum Ying Zhang, J. Fourn. & K.D. Hyde, in
Zhang, Fournier, Phookamsak, Bahkali & Hyde, Mycologia 105(3): 606 (2013)
Notes: Zhang et al. (2013) introduced Phaeoseptum,
which differs from Mauritiana in having dictyosporous
45
ascospores. There are six Phaeoseptum species: P. aquaticum (on driftwood of Salix sp., France), P. carolshearerianum (on decaying wood of Avicennia marina, India), P.
hydei (on dead twigs of Delonix regia, Thailand), P. mali
(on dead stems of Malus halliana, China), P. manglicola (on
decaying wood of Avicennia marina, India) and P. terricola
(on dead wood, Thailand).
Phaeoseptum thailandicum Samarak. & K.D. Hyde, sp. nov
Index Fungorum number: IF559754; Facesoffungi number: FoF11798; Fig. 30
Etymology: The specific epithet reflects the name of Thailand, where the species was collected.
Holotype: MFLU 19–2136
Saprobic on dead branches in terrestrial habitats. Sexual
morph: Ascomata 270–350 μm high, 160–305 μm diam.
(x̅ = 309 × 230.3 µm, n = 10), scattered to gregarious, fully
immersed under a small blackened pseudoclypeus, if appearing as black, elongated regions on host surface 540–915 μm
(x̅ = 640 µm, n = 8) length; ascomata depressed spherical,
laterally flattened. Pseudoclypeus composed of host cells
with black deposits. Peridium 6–22 µm (x̅ = 640 µm, n = 8)
wide, pseudoparenchymatous, of thin-walled cells, at apex
comprising isodiametric angular cells that are more pigmented outwardly, at sides with flattened hyaline cells, at
base of angular pigmented cells. Hamathecium comprising
1.4–2.5 μm (x̅ = 1.9 µm, n = 20), wide septate, cellular pseudoparaphyses, situated between and above the asci, embedded in a gelatinous matrix. Asci 100–155 × 20–28.5 μm
(x̅ = 129 × 23.9 μm, n = 25), 8-spored, rarely 32-spored,
bitunicate, fissitunicate, cylindrical-clavate, with a distinct
pedicel, apically rounded with a minute ocular chamber.
Ascospores 25–35 × 8–11.8 μm (x̅ = 30 × 9.9 μm, n = 30),
uniseriate at base and overlapping 2–3-seriate at apex, pale
to dark brown, broadly fusoid with broadly rounded ends,
slightly curved, 11 (9–12-transversally septate, with a vertical septum in nearly all median cells, not constricted at the
septa, the septa partly pale brown, having at maturity a thickened and heavily pigmented appearance, wall smooth, without sheath or appendages. Asexual morph: Not observed.
Material examined: Thailand, Nan, Pua District, on an
unidentified dicotyledonous dead branch, 29 January 2019,
MC Samarakoon (SAMC216), (MFLU 19-2136, holotype;
HKAS 106993, isotype), Phrae, on an unidentified dicotyledonous dead branch, 24 January 2019, MC Samarakoon
(SAMC203), (MFLU 19–2126; HKAS 106983, paratypes).
GenBank numbers: MFLU 19-2126—OM293748 (ITS),
OM293743 (LSU), OM293754 (SSU), OM305061 (tef1),
OM305067 (tub2)
MFLU 19-2136—OM293749 (ITS), OM293744 (LSU),
OM305056 (rpb2), OM293755 (SSU), OM305062 (tef1),
OM305068 (tub2)
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Fig. 30 Phaeoseptum thailandicum (MFLU 19–2136, holotype)
a,b Substarte, c,d Appearance of ascomata on the host, e,f Vertical
section through ascoma, g Peridium, h Pseudoparaphyses, i–l Asci,
m–r Ascospores. Scale bars: a,b = 1 cm, c,d = 1000 μm, e,f = 15 μm,
g-i = 10 μm, m-r = 5 μm
HKAS 106993—OM293750 (ITS), OM293745 (LSU),
OM305057 (rpb2), OM293756 (SSU), OM305063 (tef1),
OM305069 (tub2)
Notes: Our two new collections of Phaeoseptum thailandicum share similar morphology of Phaeoseptum in having
immersed ascomata under a small blackened pseudoclypeus,
cylindrical-clavate asci with a distinct pedicel and broadly
fusoid, brown ascospores with multi-transverse septa. Combined phylogeny shows that the novel taxon is sister to P.
mali + P. manglicola clade with high statistical support
(Fig. 31).
Notes: Pleopunctum was introduced to accommodate two
hyphomycetous species based on phylogenetic analyses and
divergence time which is the first asexual morph genus in
Phaeoseptaceae (Liu et al. 2019a, b, c, d). There are five Pleopunctum species are accepted in Species Fungorum (2022a,
b) with molecular data and all of them were found from
China and Thailand in terrestrial habitate (Liu et al. 2019a,
b, c, d; Phukhamsakda et al. 2020; Boonmee et al.2021).
Pleopunctum N.G. Liu, K.D. Hyde & J.K. Liu, in Liu et al.,
Mycosphere 10(1): 767 (2019)
13
Pleopunctum ellipsoideum N.G. Liu, K.D. Hyde & J.K. Liu,
in Liu et al., Mycosphere 10(1): 767 (2019).
Index Fungorumnumber: IF556523; Facesoffungi number: FoF06114; Fig. 32
Fungal Diversity (2022) 117:1–272
47
Fig. 31 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and tef1sequence data of Phaeoseptaceae. Bootstrap support values for ML ≥ than 75% and BYPP ≥ 0.95
are given above the nodes. The ex-types are in bold; the new isolates
are in blue. The tree is rooted with Amorosia littoralis (NN 6654) and
Angustimassarina populi (MFLUCC 13–0034)
Saprobic on decaying wood. Asexual morph: Hyphomycetous. Colonies on natural substrate forming sporodochial conidiomata, superficial, black, scattered, velvety,
glistening, orbicular. Mycelium immersed, composed of
branched, septate, subhyaline to pale brown hyphae. Conidiophores and conidiogenous cells not observed. Conidia
33–45 × 16–20 μm ( x̄ = 38 × 18 μm, n = 35), acrogenous, solitary, oval to ellipsoidal, muriform, constricted at septa, yellowish brown to dark brown, broadly obtuse at apex, truncate
at base. Basal cell 6.5–10.5 × 9.5–11.5 μm ( x̄ = 9–10.5 μm,
n = 35), hyaline, elliptical to subglobose, smooth walled.
Sexual morph: Not observed.
Culture characteristics: Conidium germinated on PDA
within 12 h. Colonies on PDA reaching 20 mm in 4 weeks at
26 °C. Mycelia superficial, circular, entire, flat, rough, grey
brown from above, dark brown from below.
Material examined: Thailand, Chiang Mai Province,
Mae Taeng District, MRC, on bamboo culms, 15 July 2020,
Y.R. Sun, M10 (MFLU 21–0091); living culture, MFLUCC
21–0064.
Hosts: decaying wood (Liu et al. 2019a, b, c, d) and bamboo (this study)
Distribution: China (Liu et al. 2019a, b, c, d) and Thailand (this study)
GenBank numbers: OM250079 (ITS), OM258687 (LSU)
Notes: Pleopunctum ellipsoideum was isolated from
decaying wood in China by Liu et al. (2019a, b, c, d). The
morphological characters of our collection are the same as
in P. ellipsoideum (MFLUCC 19-0390). Phylogenetic analysis based on a combined LSU, ITS and tef1 sequence data
indicated that our isolate and P. ellipsoideum (MFLUCC
19–0390) clustered together with high support (ML 100%
and 0.99 BYPP; Fig. 31). Based on both morphology and
phylogeny, we identified our taxon as P. ellipsoideum. This
is the first geographical and host report of P. ellipsoideum
on bamboo in Thailand.
13
48
Fungal Diversity (2022) 117:1–272
Fig. 32 Pleopunctum ellipsoideum (MFLU 21–0091, new
record) a, b Colonies on natural substrates c–i Conidia with
basal hyaline cells j Germinated
conidium k, l Colonies on PDA
media. Scale bars: a = 500 μm,
b = 200 μm, c = 50 μm,
d–j = 20 μm
Phaeosphaeriaceae M.E. Barr, Mycologia 71(5): 948
(1979)
Phaeosphaeriaceae has been subjected to various taxonomic changes since its establishment by Barr (1987). The
inception of multi-gene phylogenetic analyses coupled with
morphology has greatly resolved many taxonomic inconsistencies along with the introduction of several novel taxa
13
(Phookamsak et al. 2014, 2017; Li et al. 2016; Crous et al.
2018; Hyde et al. 2020a, b, c). However, confusion in the
placement of many taxa in the family still remains uncertain and problematic (Hongsanan et al. 2020b). Phaeosphaeriaceae at present accommodates more than 80 genera,
with the taxa mainly exhibiting an endophytic, pathogenic
Fungal Diversity (2022) 117:1–272
and saprobic or hyperparasitic lifestyles (Hongsanan et al.
2020b).
Nodulosphaeria Rabenh., Klotzschii Herb. Viv. Mycol., Edn
Nov, Ser. Sec., Cent. 8: no. 725 (in sched.) (1858)
Nodulosphaeria, was introduced by Rabenhorst (1858)
and typified by N. hirta and was accommodated in Phaeosphaeriaceae by Barr (1987). The genus comprises endophytic, saprobic and pathogenic taxa which occur on a
variety of hosts (Mapook et al. 2020; Chaiwan et al. 2019;
Pasouvang et al. 2021). Nodulosphaeria taxa are principally
characterised by ascomata with brown setae at the ostiole
and three-to multi-septate ascospores with a swollen cell and
often with terminal appendages (Shoemaker 1984; Mapook
et al. 2016; Chaiwan et al. 2019). Molecular data for several
Nodulosphaeria species listed in MycoBank (http://www.
mycobank.org/, 07/2022) and Index Fungorum (http://www.
indexfungorum.org/Names/Names.asp, 07/2022) are unavailable. A reference specimen for the type species N. hirta
was recently designated by Mapook et al. (2016), who also
provided molecular data for the same.
Nodulosphaeria digitalis W.J. Li, Camporesi, Bhat & K.D.
Hyde, in Li et al., Mycosphere 6(6): 683 (2015)
Index Fungorum number: IF551664; Faces of fungi number: FoF 01302; Fig. 33
= Nodulosphaeria thalictri D. Pem, Camporesi & K.D.
Hyde, in Hyde et al., Fungal Diversity: https://doi.org/10.
1007/s13225-019-00429-2, [56] (2019)
Saprobic on stem of Solidago virgaurea. Sexual
morph: Ascomata 185–200 µm high, 200–225 µm diam.
(x̅ = 192.3 × 213.1 µm, n = 5), appearing as black dots on host
surface, immersed, unilocular, perithecial, solitary, usually
scattered, globose to subglobose, dark brown, ostiolate. Ostioles 45–55 µm wide (x̅ = 52.2 µm, n = 5), centric, comprising internal dark brown setae. Ascomatal wall 14–23 µm at
the sides and base, 4–5-layered; 25–35 µm near the apex,
5–7-layered; outer layers made up of thick-walled, dark
brown cells of textura angularis, innermost layer comprising
thin-walled, pale brown to hyaline cells of textura angularis
or flattened cells. Pseudoparaphyses 1–3 µm wide, numerous, filiform, hyaline, branched. Asci 55–90 × 8–12 µm
(x̅ = 74.1 × 9.2 µm, n = 40), 8-spored, bitunicate, cylindricclavate, slightly curved, sessile to short-pedicellate, apically rounded with a minute ocular chamber. Ascospores
(17–)20–27 × 3–4 µm (x̅ = 22.6 × 3.7 µm, n = 45), overlapping 1–2-seriate, hyaline when immature, becoming yellowish-brown at maturity, long fusiform, ellipsoidal to subcylindrical, straight or slightly curved, with 4–5 transverse
septa, second cell from the apex slightly swollen, constricted
at second septum from the apex, thick- and smooth-walled,
with rounded ends, with hyaline appendages (1–2.5 µm long,
49
2.5–4 µm wide) at both ends. Asexual morph: Illustrated
in Li et al. (2015).
Material examined: Italy, Forlì Cesena, Valico del
Tramazzo, on dead aerial stem of Solidago virgaurea
L. (Asteraceae), 19 June 2021, Erio Camporesi, IT4714
(MFLU 22-0278)
Hosts: Campanula trachelium (Campanulaceae, Chaiwan
et al. 2019), Dactylis sp. (Poaceae, Li et al. 2015), Solidago
virgaurea (Asteraceae, this study), Thalictrum sp. (Ranunculaceae, Hyde et al. 2019)
Distribution: Italy (Li et al. 2015; Chaiwan et al. 2019;
Hyde et al. 2019; this study)
GenBank numbers: ON873995 (LSU); ON873997(ITS);
ON885742 (tef1)
Notes: The strain MFLU 22-0278 clustered with the
strains Nodulosphaeria digitalis MFLUCC 15-2716 (type
strain) and MFLUCC 17-2418, and Nodulosphaeria thalictri
MFLUCC 18-1138 with a good statistical support (99% ML,
0.98 BYPP) in the SSU-LSU-ITS-tef1 phylogenetic tree.
Pairwise comparison among the two strains of N. digitalis
MFLUCC 15-2716 and MFLUCC 17-2418, and N. thalictri
MFLUCC 18-1138 is as follows: one base pair (bp) difference out of 1000 bp (0.1%) in SSU between N. thalictri
MFLUCC 18-1138 and N. digitalis MFLUCC 17-2418;
no bp difference among the three strains out of 844 bp in
LSU and 504 bp in ITS respectively; two bp differences
out of 891 bp (0.2%) in tef1 between N. thalictri MFLUCC
18-1138 and N. digitalis MFLUCC 17-2418. The pairwise
identity therefore reveals insignificant differences among the
three strains (N. digitalis MFLUCC 15-2716 and MFLUCC
17-2418, and N. thalictri MFLUCC 18-1138) according to
Jeewon and Hyde (2016). Therefore, N. thalictri MFLUCC
18-1138 is another strain of N. digitalis.
Morphologically, there are some differences between
‘N. thalictri’ and N. digitalis in terms of the position of
ascomata (immersed or semi-immersed in ‘N. thalictri’
while superficial to semi-immersed in N. digitalis), ostiole
(comprising internal brown to dark brown cells of textura
globulosa in ‘N. thalictri’ while no such observation was
made for N. digitalis) (Chaiwan et al. 2019, Hyde et al.
2019). Furthermore, there are some differences in size of
the morphological structures between ‘N. thalictri’ and N.
digitalis (Table 2) (Chaiwan et al. 2019; Hyde et al. 2019).
However, all these differences may be the result of phenotypic plasticity which has arisen from the need to adapt to
environmental variations and/or different hosts (Hyde et al.
2019). The differences in the morphological characters are
not being reflected by any significant phylogenetic difference
herein. Such occurrence has also been observed for other
taxa (Mapook et al. 2016; Hyde et al. 2019; Pasouvang et al.
2021). Based on the above morpho-phylogenetic approach,
‘N. thalictri’ is considered as N. digitalis.
13
50
Fig. 33 Nodulosphaeria digitalis (MFLU 22-0278). a Ascomata on
dead aerial stem of Solidago virgaurea. b Close-up of ascomata on
host. c Vertical section through an ascoma. d Close-up of an ostiole.
e Vertical section of ascomatal wall. f Pseudoparaphyses. g–i Imma-
13
Fungal Diversity (2022) 117:1–272
ture and mature asci. j–l Immature and mature ascospores (j, l arrows
indicate appendages). Scale bars: a–c = 100 µm, d, g–i = 20 µm, e,
j–l = 10 µm, f = 3 µm
Fungal Diversity (2022) 117:1–272
51
Table 2 Morphological comparison among strains of Nodulosphaeria digitalis with sexual morph described
Strain
Host
Ascomata (μm) Ostioles
N. digitalis MFLUCC 17–2418 Campanula trachelium 141.9 × 154
N. digitalis MFLUCC 18–1138 Thalictrum sp.
263.7 × 217.9
N. digitalis MFLU xx
192.3 × 213.1
Solidago virgaurea
Fig. 34 Phylogram generated from maximum likelihood analysis
based on combined SSU, LSU, ITS and tef sequence data of selected
taxa in Phaeosphaeriaceae. Bootstrap support values for maximum
likelihood ≥ 65% and Bayesian posterior probabilities ≥ 0.95 are indi-
52.5 μm diam
No setae
71.2 µm diam
No setae
52.2 µm diam
Setae present
Asci (μm)
83 × 9
Ascospores
25 × 5 μm;
4-septate
84.1 × 10.1 32.3 × 4.5 μm;
7–8-septate
74.1 × 9.2 22.6 × 3.7 µm;
4–5-septate
Reference
Chaiwan et al. (2019)
Hyde et al. (2019)
This study
cated above or below the branches. Type strains are in bold; the new
isolate is in blue. The tree is rooted with Bhagirathimyces himalayensis NFCCI 4580, Pseudoophiobolus galii MFLUCC 17–2257 and
Pseudoophiobolus mathieui MFLUCC 17–1785
13
52
The strain MFLU 22-0278 in this study clusters with the
three strains of N. digitalis with 99% bootstrap support and
0.98 BYPP. There is no significant difference in the pairwise
comparison among strains MFLU 22-0278 and MFLUCC
15-2716, MFLUCC 17-2418 as well as MFLUCC 18-1138
with regards to the LSU, ITS and tef1 sequence data. Our
strain MFLU 22-0278 resembles N. digitalis MFLUCC
17-2418 and MFLUCC 18-1138 (N. digitalis MFLUCC
15-2716 was described in its asexual morph) in having ascomatal wall made up of cells of textura angularis, sessile or
short-pedicellate asci and septate ascospores with a swollen
cell and terminal appendages (Chaiwan et al. 2019; Hyde
et al. 2019). Our collection differs from N. digitalis MFLUCC
17-2418 and MFLUCC 18-1138 mainly in having ostioles
with internal brown setae (Fig. 34). The morphological characters also vary in size (Table 2). This can be accounted for by
environmental and host variations as stated above. These morpho-phylogenetic analyses therefore support MFLU 22-0278
as another strain of N. digitalis. The latter is herein reported
as a new record from Solidago virgaurea in Italy.
Pleosporaceae Nitschke, Verh. naturh. Ver. preuss. Rheinl.
26: 74 (1869)
This family consists of endophytes, pathogens and saprobes and has a worldwide distribution (Hongsanan et al.
2020b). Twenty-three taxa are accepted in this family
(Wijayawardene et al. 2022)
Bipolaris Shoemaker, Can. J. Bot. 33:882(1959)
Species of this genus are pathogens, saprobes and endophytes on a wide range of hosts with a worldwide distribution
(Jayawardena et al. 2019a). Bhunjun et al. (2020), provided
polyphasic approaches to delineate species in Bipolaris. There
are 140 records in Index Fungorum (2022a, b) for this genus,
however less than 50 species have molecular data.
Bipolaris luttrellii Alcorn, Mycotaxon 39: 378 (1990)
Index Fungorum number: IF127658; Facesoffungi number: FoF01302; Fig. 35
Saprobic on Poaceae sp. Sexual morph: Not observed.
Asexual morph: Conidiophores up to 200 μm long
and 7–10 μm thick, arising singly or in groups of a few
conidia, simple, septate, straight or flexuous, sometimes
geniculate at the upper part, smooth, pale to mid-brown.
Conidiogenous nodes dark brown, distinct. Conidia
(75–)90–125 × 17–26(–)28) μm ( x̄ = 107 × 21 μm, n = 30),
straight or curved, broadly fusiform or obclavate fusiform,
widest near centre, tapering towards rounded ends, pale to
mid brown, 5–9-distoseptate, smooth-walled. Hilum slightly
protuberant, single germ tubes arising from each end.
Material examined: China, Guizhou Province, Guizhou
Academy of Agricultural Sciences, dead leaves of Poaceae
sp., 20 July 2015, Kasun M. Thambugala CN020 (MFLU
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Fungal Diversity (2022) 117:1–272
16-2836), living culture MFLUCC 16–0281, GZCC
15–0045.
Host: Dactyloctenium aegyptium, on dead leaves of unidentified Poaceae host (Alcorn 1990; This study)
Distribution: Australia, China (Alcorn 1990; This study)
GenBank numbers: OQ154965 (ITS)
Notes: Bipolaris luttrelli is only known from the type
specimen before this record. This species is morphologically
similar to B. setarieae (Manamgoda et al. 2014). Bipolaris
luttrelli can be distinguished from its sister taxa by having
fewer conidiogenous loci on the conidiophores and its darker
conidia with pale end cells (Manamgoda et al. 2014). In our
phylogenetic analyses (Fig. 36), our strain clustered with the
ex-type strain of B. luttrelli with a higher bootsrap support.
There are 10 base pair differences between our strain and
the ex-type strain in ITS. Both of these strains lack tef1 gene
region. Therefore, we identified our strain as B. luttrelli and
provides the first report from China.
Curvularia Boedijn, Bull. Jard. bot. Buitenz, 3 Sér. 13(1):
123 (1933)
This is a species-rich genus which comprised of numerous
pathogenic, saprobic and epiphytic fungi (Manamgoda et al.
2012a, b, 2015; Jayawardena et al. 2019a). Phytopathogenic
species have been recorded mostly on poaceous hosts as well
as non-poaceous hosts. Several Curvularia spp. have also
been reported as opportunistic human pathogens on immunocompromised patients (Madrid et al. 2014; Manamgoda
et al. 2011, 2015; Danish Khan et al. 2017; Tóth et al. 2020).
There are more than 200 epithets recorded in Index Fungorum (2022a, b). To date, 130 species of Curvularia have been
accepted within the genus (Ferdinandez et al. 2021).
Curvularia alcornii Manamgoda, L. Cai & K. D. Hyde,
Sydowia 64(2): 259 (2012)
Index Fungorum number: IF800665; Facesoffungi number: FOF10679; Fig. 37
Saprobic on dried leaf of Panicum virgatum. On CMA
Hyphae 4–5 μm, septate, branched. Conidiophores up
to 241 μm long, micronematous to macronematous,
pale brown to dark brown, simple or branched, septate, flexuous, highly geniculate. Conidiogenous cells
(8–)9–17(–19) × (4–)5–6 μm ( x̄ = 13 × 5 μm, n = 10), hyaline to pale brown, smooth–walled, terminal or intercalary,
monotretic to polytretic. Conidia (18–)20–26(–31) × (7–)8–1
0(–11) μm ( x̄ = 23 × 9 μm, n = 30) apical and basal cells hyaline or pale brown, matured conidia brown, straight, rarely
curved, inequilateral ellipsoidal or clavate, dark brown septa,
3–4-distoseptate, enlarged middle cells; hila 1–2 μm distinctly protuberant, darkened. Sexual morph: Not observed.
Culture characteristics: Colonies on PDA reaching
57 mm in 7 days at 25 °C, slightly convex with entire margin, brown centre, sparse aerial mycelium becoming brown
Fungal Diversity (2022) 117:1–272
53
Fig. 35 Bipolaris luttrelli
(MFLU 16–2836, new host
record) a, b, Conidiophores
and conidia on the host c-f.
Conidiophores g–k, Conidia.
Scale bars: c–k = 25 µm
to grey towards the edge, reverse black to dark brown, concentric. Colonies on CMA reaching 60 mm in 7 days at
25 °C, flat with entire margin, pale brown to dark brown,
moderate aerial mycelium becoming dark brown to black
towards the edge, reverse dark brown, concentric. Colonies
on MEA reaching 67 mm in 7 days at 25 °C, flat with entire
margin, dark green, sparse aerial mycelium becoming olivaceous green towards the edge, reverse black, concentric.
Material examined: Sri Lanka, Anuradhapura District,
Thuruwila, N 8° 14′ 50.53859″, E 80° 25′ 9.24013″, on dried
leaf of Panicum virgatum L. (Poaceae), 13 June 2019, D.S.
Manamgoda, USJ-H-075, living culture USJCC-0088.
Hosts: Panicum spp., Pennisetum clandestinum, Oryza
spp. and Zea mays (Manamgoda et al. 2012a, b; Khemmuk
et al. 2016)
Distribution: Australia and Thailand (Farr and Rossman
2022)
GenBank numbers: MZ948821 (ITS), MZ971267
(gadph), MZ971253 (tef1)
Notes: According to the phylogenetic result, isolate
USJCC-0088 is identified as Curvularia alcornii. This taxon
was originally described in Manamgoda et al. (2012a), as a
saprobe on a leaf sample of Zea mays collected in Thailand
(holotype MFLU 12-0397). In this study, the fresh isolate
was identified as a saprobe on a dead leaf of Panicum virgatum (Fig. 38). To our knowledge, this is a new record from
Sri Lanka and a new fungus-host association.
Curvularia senegalensis (Speg.) Subram., Journal of the
Indian Botanical Society. 35(4): 467 (1956)
≡ Brachysporium senegalense Speg., Anales del Museo
Nacional de Historia Natural Buenos Aires 26: 133 (1914)
Index Fungorum number: IF296254; Facesoffungi number: FoF13382; Fig. 39
Saprobic on dead panicle of Zea mays. On PDA Hyphae
3–4 μm, septate, branched. Conidiophores up to 219 μm
long, micronematous to macronematous, pale brown
to dark brown, simple or branched, septate, straight to
13
54
Fungal Diversity (2022) 117:1–272
Fig. 36 One of the most parsimonious trees obtained in the combined
analyses of ITS-gapdh-tef1. Maximum Parsimony values ≥ 70% and
BYPP ≥ 0.90 are indicated above the nodes. The ex-types and refer-
ence strains are in bold; the new record is in blue. The tree is rooted
with Curvularia buchloes (CBS 246.49) and C. subpapendorfii (CBS
656.74)
flexuous, geniculate at the apex. Conidiogenous cells (8–)
9–15(–17) × 3–5 μm ( x̄ = 12 × 4 μm, n = 10), hyaline to pale
brown, smooth–walled, terminal or intercalary, monotretic
to polytretic. Conidia (17–)18–27(–34) × (7–)9–12(–13) μm
( x̄ = 22 × 11 μm, n = 30) apical and basal cells hyaline or pale
brown, matured conidia brown, straight to curved, sometimes clavate, dark brown septa, 3–4-distoseptate, enlarged
13
Fungal Diversity (2022) 117:1–272
55
Pinus caribaea, Pinus khasya, Quercus germana, Saccharum sp., Stigmaphyllon sagraeanum, Tamarindus indica,
Thuja orientalis, Urena lobate, Vigna unguiculata, Zea
mays (Farr and Rossman 2022)
GenBank numbers: MT410577 (ITS), MZ971268
(GADPH), MZ971254 (tef1)
Notes: Isolate USJCC-0025 is identified as Curvularia senegalensis based on morphology and phylogeny
(Fig. 38). The fresh isolate was collected from a dead
panicle of Zea mays. Curvularia senegalensis has so far
recorded in Sri Lanka only from Hevea brasiliensis (Adikaram and Yakandawala 2020). Moreover, it has only been
reported on Zea mays from Brazil, Malaysia and Nigeria.
To our knowledge, this is the first record of Curvularia
senegalensis on Zea mays from Sri Lanka.
Fig. 37 Curvularia alcornii (USJCC–0088, new host and geographical record) a Seven-day old colony on PDA b Seven-day
old colony on CMA c Seven-day old colony on MEA d, e Conidia
attached to conidiophores f, g Conidia. Scale bars: d–g = 10 μm
middle cells; hila 1–2 μm flat, darkened. Sexual morph:
Not observed.
Culture characteristics: Colonies on PDA reaching
73 mm in 7 days at 25 °C, flat with entire margin, grey to
olivaceous black centre, abundant aerial mycelium becoming brown towards the edge, reverse black to dark brown,
concentric. Colonies on CMA reaching 69 mm in 7 days at
25 °C, flat with entire margin, grey to pale brown, reverse
white to pale brown, concentric. Colonies on MEA reaching 83 mm in 7 days at 25 °C, flat with entire margin, dark
brown, abundant aerial mycelium becoming grey towards
the edge, reverse dark brown to pale brown, concentric.
Material examined: Sri Lanka, Matale District, Palapathwela, N 7° 33′ 22.8″, E 80° 36′ 38.2″, on panicle of Zea
mays L. (Poaceae), 08 November 2018, D.S. Manamgoda,
USJ-H-031, living culture USJCC-0025.
Distribution: Australia, Brazil, China, Cuba, Hawaii,
India, Malaysia, Mexico, Myanmar, Nigeria, Samoa, South
Africa, Sri Lanka, Tanzania, Thailand, Texas, United States,
Virginia and West Indies (Farr and Rossman 2022)
Hosts: Andropogon caricosus, Archontophoenix alexandrae, Bauhinia purpurea, Carya illinoensis, Citrullus vulgaris, Cymbopogon flexuosus, Cynodon dactylon, Dichanthium caricosum, Gmelina arborea, Hevea brasiliensis,
Hibiscus cannabinus, Jasminum sambac, Liquidambar
macrophylla, Musa nana, Musa × paradisiaca, Oryza
sativa, Ougeinia oojeinensis, Paspalum notatum, Paspalum paniculatum, Passiflora edulis, Persea Americana,
Pyrenophora Fr., Summa veg. Scand., Sectio Post. (Stockholm): 397 (1849)
Pyrenophora is a species-rich genus in Pleosporales
which encompasses saprobic and phytopathogenic fungi
associated mainly with poaceous hosts. The sexual morphs
are characterized by black, thick-walled, subglobose to pyriform ascomata with an apical ostiole and conspicuous dark
brown setae. Asci are bitunicate, show a large non-amyloid
ring and usually contain eight pale yellowish brown or pale
brown, muriform ascospores surrounded by a mucilaginous
sheath. Asexual morphs show macronematous, brown,
sympodial conidiophores with tretic conidiogenous cells
and dematiaceous, rather straight, distoseptate conidia with
a dark, non-protruding basal scar. Conidial germination
occurs from polar or intermediate cells (Sivanesan 1987).
This genus is monophyletic and genetically clearly distinct
from other pleosporalean genera with a superficially similar conidial apparatus, such as Bipolaris, Exserohilum and
Porocercospora. Multilocus DNA sequence data is currently
available for at least 26 Pyrenophora species (Marin-Felix
et al. 2019).
Pyrenophora verruculosa Madrid & Cantillo, sp. nov.
Mycobank number: 844464; Facesoffungi number:
FoF10420; Fig. 40
Etymology: The name refers to the verruculose conidia
produced by this species
Holotype: SGO 168420
Probably saprobic or pathogenic to an unidentified
member of Poaceae. Sexual morph: Not observed. Asexual morph: Hyphomycetous. Vegetative hyphae septate,
branched, light olivaceous to mid olivaceous brown, thin
to thick-walled, smooth, 2–6 µm wide, anastomosing,
occasionally showing deposits of a mucilaginous dark
brown material. Conidiophores macronematous, mononematous, solitary, septate, simple, slightly flexuous to
strongly geniculate, light olivaceous brown to dark brown,
13
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Fungal Diversity (2022) 117:1–272
Fig. 38 Phylogram generated from parsimony analysis based on combined ITS, gadph and tef1 sequence data of Curvularia. Bootstrap
support values of MP and ML ≥ 50% and BYPP ≥ 0.95 are indicated
above the nodes. The ex-types are in bold; the new records are highlighted in greenish-blue. The tree is rooted with Bipolaris maydis
(CBS137271/C5)
often paler at the apex, smooth to verruculose, with cell
walls often thicker than those of the supporting vegetative
hyphae, 300–1270 × 5–9 µm with subnodulose to nodulose
intercalary swellings up to 11 µm wide. Conidiogenous
cells integrated, terminal and intercalary, mostly subcylindrical, mono- to polytretic, proliferating sympodially,
15–28 µm long. Conidia narrowly clavate, narrowly ellipsoidal to fusiform or subcylindrical, straight to slightly
curved, light olivaceous brown to dark brown, verruculose,
(26–)32–63(–74) × 12–21 µm, 3–5(mostly 4)-distoseptate, often constricted at the uppermost distoseptum, with
a rounded apex and an obconically truncate or rounded
base, basal cell sometimes delimited by a thick, dark septum. Hilum thick and dark. Microsclerotia abundant, mostly
45–240 µm wide.
Culture characteristics: Colonies on water agar with sterilized maize leaves dark brown, hairy, with abundant clumps
of microsclerotia.
Material examined: Chile, El Loa Province, Atacama
Desert, near Calama, isolated from unidentified dead
Poaceae, 15 October 2015, H. Madrid & L. Linaje (SGO
168420, holotype).
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57
Fig. 39 Curvularia senegalensis (USJCC–0025, new host
record) a Host: dead panicle
of Zea mays b Seven day old
colony on PDA c Seven day old
colony on CMA d Seven day
old colony on MEA e Conidiophore f Conidia attached
to conidiophore g Germinating
conidium h, I Conidia Scale
bars: e–i = 10 μm
GenBank numbers: ON722346 (ITS), ON722346 (LSU),
ON736764 (gpdh).
Notes: Due to mobility restrictions during the SARSCoV-2 pandemic, the ex-type strain (HM 201), which had
been preserved in sterile water, could not be properly maintained for several months. Recent attempts to reactivate
the strain have been unsuccessful and the fungus probably
died. However, the holotype was deposited at SGO and extype sequences of ITS, LSU and gpdh are available in GenBank. DNA sequence analyses revealed that Pyrenophora
verruculosa is clearly distinct from all other members of
Pyrenophora represented in GenBank. The closest hits in
BLAST searches with the ITS sequence of strain HM 201
were Pyrenophora novozelandica (CBS 127934) (ex-type,
GenBank MK539997, 96.55% similarity), P. fugax (CBS
509.77) (GenBank MK539985, 95.23% similarity), P. nisikadoi (CBS 190.29) (ex-type, as Bipolaris brizae, GenBank MH855213, 93.57% similarity), and P. nobleae (CBS
259.80) (GenBank MK539994, 91.33% similarity). BLAST
searches with the gpdh sequence showed P. fugax (CBS
509.77) (GenBank AY004822, 95.70% similarity) and P.
nobleae (CBS 966.87) (GenBank AY004824, 88.81% similarity) as the closest matches. With LSU, similar results were
obtained, but with higher similarity percentages, as expected
for this rather conserved locus. In the phylogenetic analysis,
only ITS sequence data was used considering that LSU offers
little resolution for closely related taxa in Pleosporales, and
gpdh is available for a smaller number of species than ITS.
In the ITS-based phylogenetic tree (Fig. 41), P. verruculosa, P. novozelandica and P. fugax formed a clade with 83%
bootstrap support. These close relatives can easily be distinguished from P. verruculosa on the basis of conidial dimensions, i.e. smaller in P. novozelandica, 20.5–58 × 9.5–14 µm,
and longer in P. fugax, 50–170 × 14–24 µm (Ellis 1961;
Sivanesan 1987; Marin-Felix et al. 2019). Clumps of thickwalled, strongly pigmented cells superficially resembling
the microsclerotia of P. verruculosa, have been reported
in other Pyrenophora species, such as P. nisikadoi and P.
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Fungal Diversity (2022) 117:1–272
Fig. 40 Pyrenophora verruculosa (SGO 168420,
holotype). a, c, d Conidia
attached to conidiogenous cells.
b Conidium passively released
from a conidiogenous cell. e–h
Conidia (verruculose ornamentation can be observed in e). i
Hyphae with mucilaginous dark
brown material. j, k Mycelia
with microsclerotia. Scale bars:
a–d = 40 µm, e–i = 20 µm,
j–k = 45 µm
pseudoerythrospila (Marin-Felix et al. 2019). These species, however, are phylogenetically clearly distinct from P.
verruculosa (Fig. 41).
Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray., Stud.
Mycol. 64: 177 (2009)
13
Tetraplosphaeriaceae was established to accommodate five genera, viz Polyplosphaeria, Pseudotetraploa,
Quadricrura, Tetraplosphaeria with Tetraploa sensu
stricto asexual morphs (the type genus) and Triplosphaeria (Tanaka et al. 2011). Hyde et al. (2013) later treated
Tetraplosphaeria as a synonym of Tetraploa which has been
applied previously. Up to now, nine genera are accepted in
Fungal Diversity (2022) 117:1–272
Fig. 41 Maximum likelihood tree based on ITS sequences, showing
the phylogenetic relationships of Pyrenophora species. Bootstrap support values > 70% are shown near the internodes. The tree is rooted
to Bipolaris maydis. GenBank accession numbers of ITS sequences
59
are given, in parentheses, after each strain number. Aauthentic strain,
ex-type strain, ETex-epitype strain, HTholotype material. New species
is in bold
T
13
60
Fungal Diversity (2022) 117:1–272
Fig. 42 Tetraploa dashaoensis (KUN–HKAS 107,636, holotype). a–b. Conidia on host substrate; c–g. Conidia; k. Verrucose conidia; l. Germinating conidium; m. Colony on PDA (left-front, right-reverse); Scale bars: c–f, k = 50 μm, g–j, l = 100 μm
Tetraplosphaeriaceae viz Aquatisphaeria, Byssolophis,
Ernakulamia, Polyplosphaeria, Pseudotetraploa, Quadricrura, Shrungabeeja, Tetraploa and Triplosphaeria
(Wijayawardene et al. 2022). Members of this family are
mostly reported as saprobe from aquatic and terrestrial habitats (Tibpromma et al. 2018; Hongsanan et al. 2020a, b;
Hyde et al. 2020a, b, c; Li et al. 2021a, b).
13
Tetraploa Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 5:
459 (1850) MycoBank: MB10199
= Tetraplosphaeria Kaz. Tanaka & K. Hiray., in Tanaka
et al., Stud. Mycol. 64: 177 (2009)
Notes: Tetraploa was introduced by Berkeley and Broome
(1850) with T. aristata as the type species. The sexual
morph is characterized by small globose ascomata, narrowly
fusiform ascospores and appendage-like sheath. The asexual
Fungal Diversity (2022) 117:1–272
Fig. 43 Phylogram generated from maximum likelihood analysis of
Tetraploa including related genera based on a combined LSU, ITS,
SSU, tub2 and tef1 sequence data. Relevant sequences were referred
from Dong et al. (2020), Hyde et al. (2020a), and Li et al. (2021a, b).
The data set consisted of 3720 characters with gaps. Tree topology
61
of the ML analysis was similar to the MP and BYPP tree topologies.
Bootstrap support values of ML and MP ≥ 75% with BYPP ≥ 0.95 are
given at the nodes. The ex-type cultures are indicated using ″T″; the
new isolate is in blue
13
62
morph belonging to Tetraploa sensu stricto has a columnar
conidial body, with several prominent setose appendages at
the apex (Hyde et al. 2013; Li et al. 2021a, b). Hyde et al.
(2013) transferred species in Tetraplosphaeria to Tetraploa. In this study, we introduce one new species to Tetraploa based on morphology and molecular analysis with full
description and illustration.
Tetraploa dashaoensis C.F. Liao & Doilom, sp. nov.
Index Fungorum number: IF559273; Facesoffungi number: FoF10585; Fig. 42
Etymology: In reference to the location where the fungus
was collected.
Holotype: KUN–HKAS 107636
Saprobic on dead stem of Saccharum arundinaceum. Sexual morph: Not observed. Asexual morph: hyphomycetous.
Colonies superficial, effuse, gregarious, brown to dull green.
Mycelium partly immersed in natural substratum, brunched,
septate, hyaline. Conidiophores absent. Conidiogenous cells
micronematous, integrated, monoblastic, intercalary, short
cylindrical. Conidia 25–43 × 13–36 µm ( x̄ = 31 × 23 µm,
n = 30), solitary, straight, septate, unbranched, mostly
smooth-walled at the base of immature conidia, becoming verrucose at mature conidia, composed of 1–4 columns
at the base, 1–3-septate in each column, with 1–4 apical
appendages. Appendage 148–258 µm long ( x̄ = 186 µm,
n = 30), 6–15 µm wide at the base with dull green, 3–8 µm
wide at the apex with hyaline, usually composed of three
to four appendages, rarely one or two, euseptate, 6–23-septate, smooth, straight or divergent to intertwined; immature appendages 21–81 µm long ( x̄ = 43 µm, n = 30), with
1–5-septate.
Culture characteristics: Conidia germinating on PDA,
germination tube growing from both ends. Colonies on PDA
reaching 10 to 14 mm diam. in 17 days at room temperature (25 ± 2 °C), mycelium dense, floccose, surface smooth,
velutinous spot centre with flat substrate, circle or irregular
margin, light brown to yellow–brown, dark brown towards to
white margin in above; white at centre, dark brown towards
margin in reverse. No pigment production.
Material examined: China, Yunnan Province, Kunming
City, from dead stem of Saccharum arundinaceum (Retz.)
(Poaceae), 4 July 2020, C.F. Liao, (KUN–HKAS 107636,
holotype), ex-type living culture KUMCC 21-0010.
GenBank number: OL473549 (ITS), OL473555 (LSU),
OL473556 (SSU), OL505601 (tub2), OL505599 (tef1).
Notes: Phylogenetic analyses of the combined LSU, ITS,
SSU, tub2 and tef1 sequence data showed that our new collection Tetraploa dashaoensis KUMCC 21-0010 is related
to Tetraploa aquatica (MFLU 19-0995, MFLU 19-0996),
but forms a distinct linage with 98% ML, 83% MP and 0.99
BYPP values (Fig. 43). Tetraploa dashaoensis shares some
morphological characteristics with the phylogenetically
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Fungal Diversity (2022) 117:1–272
closest species T. aquatica in having cylindrical conidiogenous cells, a verrucose conidial base and 1–4 apical
appendages with euseptate. However, T. dashaoensis differs
from T. aquatica in having a different conidial colour (dull
green vs. brown to pale brown), longer conidia (25–43 µm
vs. 22.5–27 µm), longer appendages (148–258 µm vs.
98–134 μm), and more septate appendages (6–23-septate vs.
6–10-septate). In addition, T. dashaoensis has intertwined
appendages, which were absent in T. aquatic (Li et al. 2021a,
b). Phylogenetic analyses and morphological characters support our collection to be different species. Thus, we introduce T. dashaoensis as a new species based on morphology
and phylogeny.
Torulaceae Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze
Deutschl.) 2: 71 (1829)
Notes: Sturm (1829) introduced Torulaceae with Torula
as the type genus. There are six genera accepted in Torulaceae, namely Dendryphion, Neotorula, Rostriconidium,
Rutola, Sporidesmioides and Torula (Hongsanan et al.
2020b). Torulaceae species are characterized by having
erect, micro- or macronematous, straight or flexuous, subcylindrical conidiophores and doliiform to ellipsoid or clavate,
brown, smooth to verruculose conidiogenous cells. Conidia
are subcylindrical, phragmosporous, acrogenous, brown,
dry, and smooth to verrucose, characteristically produced
in branched chains (Hyde et al. 2016; Li et al. 2017a, b;
Mapook et al. 2020). In this study, we follow Mapook et al.
(2020) as the recent treatment for Torulaceae.
Torula Pers, Ann. Bot. (Usteri) 15: 25 (1795)
Notes: Persoon (1794) established Torula and was typified by T. herbarum. Torula species are characterized by
having a terminal or lateral, monoblastic or polyblastic
conidiogenous cells, which have a basally thickened and
heavily melanized wall, with the apex thin-walled and
frequently collapsing and becoming coronate (Crane and
Miller 2016). There are 54 Torula species listed in Species Fungorum (2022a, b), however, only 18 taxa have
molecular data.
Torula fici Crous, IMA Fungus 6: 192 (2015).
Index Fungorum number: IF816154; Facesoffungi number: FoF02712, Fig. 44
Saprobic on the dead pseudo stem parts of Musa sp.
Sexual morph: Not observed. Asexual morph: Colonies
effuse, black, powdery and thread-like. Mycelium slightly
immersed, septate, branched, smooth, with pale brown
hyphae. Conidiophores 2–3 μm long × 1–3 μm diameter
( x̄ = 2.77 × 1.91 μm, n = 10), macronematous, mononematous, solitary, erect, thick–walled, consisted of 2 distinct cells, first cell from the bottom is pale brown to sub
hyaline, wide at the base and narrow at apex, the second
Fungal Diversity (2022) 117:1–272
63
Fig. 44 Torula fici (MFLU
22-0251, new collection).
Fungal colonies on host surface,
b, c, e–p Budding, conidial
formation and mature conidia, d
Conidiogenous cell. Scale Bars:
a = 500 μm, b, i, k, m = 20 μm,
e–h, j, l, n–p = 15 μm, c,
d = 5 μm
cell is pale brown to brown, different from the first cell by
shape, cylindrical to subcylindrical at the base and globose
to subglobose at apex, conidiophore arising from prostrate
hypha. Conidiogenous cells 2–3 μm long × 1–2 μm diam,
( x̄ = 2.18 × 1.33 μm, n = 10) polyblastic, and terminal, dark
brown to black, verruculose, thick-walled, ellipsoid, paler or
sub hyaline at apex, dark and black at the bottom. Conidia
19– 25 μm long 2–3.5 μm wide ( x̄ = 18.56 × 2.88 μm, n = 10)
solitary to catenate, acrogenous, simple, phragmosporous,
dark brown, apical cell often pale brown, minutely verruculose, often 3–4 septate, rounded at both ends, composed of
subglobose cells, slightly constricted at some septa. Immature conidia are sub hyaline to pale brown and arising to
multi-angles from mature conidia. Conidial secession is
rheixolytic.
Culture characteristics: Conidia germinating on PDA
within 18 h and germ tubes produced from the tip cell.
Colonies growing on PDA, reaching 5 cm in 5 days at 25 °C,
mycelium partly immersed to superficial, slightly effuse,
hairy, with regular edge, pink or pale brown.
Material examined: Thailand, Chiang Mai, Mushroom Research Center, on dead plant material of Musa sp.
(Musaceae), 30 December 2018, Binu Samarakoon, E001
(MFLU 22-0251), living culture MFLUCC 22-0176.
Other material examined: Taiwan, Ali shan mountain,
Fanlu Township area, Dahu forest, dead leaves of Ficus
septica (Moraceae), 20 September 2018, D. S. Tennakoon,
HAY029A (MFLU 19-2775); living culture, MFLUCC
20-0167, ibid., 10 July 2019, HAY029B (NCYU 19-0367);
living culture, NCYUCC 19-0248.
Known hosts: Chromolaena odorata (Asteraceae), Ficus
septica (Moraceae), Ficus religiosa (Moraceae), Garcinia
sp. (Clusiaceae), Magnolia grandiflora (Magnoliaceae),
Olea europaea (Oleaceae), Pandanus sp. (Pandanaceae),
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Fungal Diversity (2022) 117:1–272
◂ Fig. 45 The best scoring RAxML tree for combined dataset of LSU,
SSU, ITS, tef1 and rpb2 sequence data. The topology and clade stability of the combined gene analyses was compared to the single gene
analyses. The tree is rooted with Neooccultibambusa chiangraiensis
(MFLUCC 12–0559) and Occultibambusa bambusae (MFLUCC
13–0855). Ex-type strains are in bold and newly generated sequences
are in red. Bootstrap support values for ML ≥ 60% and BYPP ≥ 0.90
are given above the nodes
Musa sp. (Musaceae) (Crous et al. 2015a, b, c, Li et al.
2017a, b; Tibpromma et al. 2018; Jayasiri et al. 2019;
Mapook et al. 2020; Samarakoon et al. 2021; Tennakoon
et al. 2021; this study).
Known distribution: Taiwan (this study), Cuba, South
Africa, Thailand (this study) (Crous et al. 2015a, b, c, Li
et al. 2017a, b; Tibpromma et al. 2018; Jayasiri et al. 2019;
Mapook et al. 2020; Samarakoon et al. 2021; this study).
GenBank numbers: MFLUCC 22-0176 – OP099550
(LSU), OP097673 (SSU), OP099562 (ITS), OP113821
(tef1) MFLUCC 20-0167 – MZ317501 (LSU); MZ317506
(ITS); MZ326657 (tef1); MZ326659 (rpb2) NCYUCC 19-0248 – MZ317502 (LSU); MZ317507 (ITS);
MZ326658 (tef1); MZ326660 (rpb2)
Notes: In the multi-gene phylogeny, our strain (MFLUCC
22-0176) grouped with T. fici strains with moderate statistical support in ML analysis and a higher support in Baysian
analysis (81% ML, 0.98 BYPP) (Fig. 45). Morphological
comparisons revealed that our collection is similar to the
holotype of T. fici (Crous et al. 2015a, b, c) by the distinct
shape of the conidiophores and having 3–4 celled conidia.
In addition, the conidiogenous cells of the type specimen
are identical in shape with our collection. Therefore, based
on both morphology and phylogeny evidence, we identify
our specimen (MFLU 22-0251) as T. fici from Chiang Mai,
Thailand. Torula fici was previously found from China on
Musa sp. by Samarakoon et al. (2021). In this study, we
report T. fici on Musa sp. for the first time from Thailand as a
new collection. In addition, our strains (MFLU 19-2775 and
NCYU 19-0367) share the size range of the conidial characters with T. fici (Crous et al. 2015a, b, c; Tibpromma et al.
2018; Mapook et al. 2020). The phylogeny also indicates
that both strains were nested with other Torula fici strains in
a 100% ML, 1.00 BYPP supported clade (Fig. 45). We conclude that our stains (MFLU 19-2775 and NCYU 19-0367)
also belong to T. fici which was reported on Ficus septica
from Taiwan as a new host record.
Torula sundara (Subram.) Y.R. Sun, Yong Wang bis & K.D.
Hyde, comb.nov.
≡ Dwayabeeja sundara Subram., J. Indian Bot. Soc.37:
56 (1958)
= Pseudotorula sundara (Subram.) J.L. Crane & A. Mill
Index Fungorum number: IF559464; Facesoffungi number: FoF09933; Fig. 46
65
Saprobic on decaying bamboo culms in terrestrial habitat.
Asexual morph: Hyphomycetous. Colonies on natural substrate superficial, powdery, dark brown to black. Mycelium
immersed, composed of branched, septate, pale brown to
brown hyphae. Conidiophores 2.5–4 μm wide, micronematous to semi-macronematous, mononematous, solitary,
erect, sample, straight or slightly flexuous, unbranched,
paler brown to brown, thin-walled, septate, with ampulliform cells, arising from prostrate hypha. Conidiogenous
cells polyblastic, terminal, with the apex thin-walled and
freuently collapsing and becoming coronate, dark brown
to black, ellipsoid to coronal. Conidia two types, short
conidia and long conidia. Short conidia 41–60 × 9–15 μm
( x̄ = 53 × 12 μm, n = 30), acrogenous, phragmosporous, single or in chains, broadly fusiform, yellow brown to dark
brown, 5–10-septate, slightly constricted at some septa,
verruculose. Long conidia acrogenous, phragmosporous,
single, straight or flexuous, cylindrical, up to 50-septate,
constricted at the septa, brown to dark brown, up to 200 μm
long. Sexual morph: Not observed.
Culture characteristics: Conidium germinated on PDA
within 12 h. Colonies on PDA reaching 20 mm in two weeks
at 26 °C. Mycelia superficial, circular, entire, flat, white
from above, pale brown from below.
Material examined: Thailand, Chiang Mai Province,
Mae Taeng District, MRC, on bamboo culms, 10 September 2020, H.W. Shen, M55 (MFLU 21-0089); living culture,
MFLUCC 21-0067).
GenBank numbers: OM276824 (ITS), OM287866 (LSU)
Notes: Dwayabeeja sundara as the type species in Dwayabeeja was introduced by Subramanian (1958). It has dark
blackish-brown colonies and two types of conidia. Crane and
Miller (2016) transferred D. sundara to Pseudotorula based
on catenate phragmoconidia. It was recollected and isolated
from bamboo culms in Chiang Mai Province, Thailand.
Here, we provided sequences for it. Phylogenetic analyses
show it is a distinct clade in Torula and sister to T. acaciae
with high support (ML 100% and 1.00 BYPP). According
to the similarities in morphology along with phylogenetic
analyses, we propose Pseudotorula sundara as a synonym
of Torula sundara.
Periconiaceae (Sacc.) Nann., Repertorio sistematico dei
miceti dell'uomo e degli animali 4: 482 (1934).
Periconiaceae was accommodated as a distinct lineage
in Massarineae by Tanaka et al. (2011) based on morpho
molecular data. Four genera (viz. Bambusistroma, Flavomyces, Noosia and Periconia) are accepted with endophytic,
saprobic and pathogenic nutritional modes (Sarkar et al.
2019; Hongsanan et al. 2020b, Samarakoon et al. 2020;
Wijayawardene et al. 2022).
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Fig. 46 Torula sundara (MFLU
21–0089, new combination).
a, b Colonies on the host c–e
Conidiophores and conidia f–l
Conidia m Germinated conidium. Scale bars: c = 10 μm,
d–m = 20 μm
Periconia Tode, Fung. mecklenb. sel. (Lüneburg) 2: 2
(1791).
Periconia was introduced by Tode (1791) and typified
by P. lichenoides. The asexual morph of the genus has pale
to dark brown branched or unbranched conidiophores (also
referred to as stipe). Conidiogenous cells of Periconia are
monoblastic or polyblastic, and formed at the terminal
ends or intercalary parts of the conidiophore. Periconia
produces spherical, asepete, catenate or solitary conidia in
pale to dark-brown. Both sexual and hypomycetous asexual
morphs are recorded in this genus (Tanaka et al. 2015; Liu
et al. 2017). There are 187 epithets listed in Index Fungorum
(2022a, b). Periconia has been reported as a plant pathogen,
13
endophyte and a common saprobe in terrestrial and aquatic
habitats with a cosmopolitan distribution (Ellis 1961, 1976;
Markovskaja and Kačergius 2014; Liu et al. 2017).
Periconia byssoides Pers., Syn. meth. fung. (Göttingen)1:
18 (1801)
Index Fungorum number: 144538; Faceoffungi number:
FOF09319; Figs. 47, 48
Saprobic on leaves of Vigna unguiculata. Sexual morph:
Not observed. Asexual morph: Conidiophores with conidial heads were observed on foliar lesions of cowpea. Conidiophores observed after 2–3 weeks, simple, micro- and
semi-macronematous, unbranched and branched, initially
Fungal Diversity (2022) 117:1–272
67
Fig. 47 Cowpea leaves showing the growth of Periconia byssoides on the necrotized lesions
Fig. 48 Periconia byssoides (MD2, new host and geographical
record): a–c Conidiophores terminating with heads of conidia d–e
Heads of conidia showing phialides f – Basal part of conidiophores
showing their distinct nature (color and septa) g Close view of the
conidial head showing conidia and phialides h–j Conidia enlarged.
Scale bars: a − g = 50 μm, h − j = 20 μm
13
68
13
Table 3 Comparative account of Periconia species with P. byssoides recorded on Cowpea from India
Species
Conidiophores
Conidia
Periconia byssoides
Periconia cookei
(Mason and Ellis 1953; Ellis
1971)
Periconia shyamala
(Ellis 1971; Storey 2002)
Macronematous and proliferating with elongate apical
cell and with branchlets
Macronematous and proliferating with elongate apical
cell and with branchlets
Macronematous, with numerous proliferations and
swollen apical cell without
branchlets
Macronematous and proliferations and swollen apex with
without apical cell
Macronematous with elongate
apical cell and branchlets
Periconia typhicola
(Ellis 1976)
Periconia prolifica
(Anastasiou 1963; Vrijmoed
et al. 1982)
Macronematous branched
inside the head
Micronematous, semimicronematous with proliferations
Periconia byssoides
(Mason and Ellis 1953; Ellis
1971; Matsushima 1975)
Periconia pseudobyssoides
(Markovskaja and Kačergius
2014)
Color
Habit
Reference
Spherical, verrucose; 8.5–14.4 Pale brown to brown
(18.2) µm in diam
On cowpea leaves
Present study
Spherical, verrucose; 10–12
(17) µm in diam
Pale brown to brown
Terrestrial saprobe
Markovskaja and Kačergius
(2014)
Spherical to verrucose
(12)15–17(20) µm in diam
Golden brown to reddish
brown
Terrestrial saprobe
Markovskaja and Kačergius
(2014)
Spherical, verrucose
13–16 µm in diam
Brown to dark brown
Terrestrial saprobe
Markovskaja and Kačergius
(2014)
Spherical, verrucose
13–16(18)-22(25) µm in
diam
Spherical, verrucose;
11–17 µm in diam
Subglobose, smooth,
7.5–15 µm in diam
Brown to dark brown
Terrestrial parasite or saprobe Markovskaja and Kačergius
(2014)
Pale brown to brown
Terrestrial saprobe
Subhyaline to pale brown
Marine saprobe
Markovskaja and Kačergius
(2014)
Markovskaja and Kačergius
(2014)
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subhyaline to brownish, verruculose, and variable in length.
Conidiogenous cells discrete, determinate, terminal or lateral, subglobose, mono- and polyblastic, smooth to verruculose, pale brown producing global verrucose conidia in acropetal chains (3–4 in number), 11.5–12.5 × 15 − 17 μm diam.
(x̄ = 11.9 × 15.2 μm, n = 20), conidia pale brown to brown
verrucose. Conidiogenous cells were formed on an apical
cell and in the collar region around the septa, sometimes
on short hyaline or subhyaline branchlets. From primary,
hyaline, globose conidiogenous cells numerous secondary
conidiogenous cells arise, which produce short chains of
spherical, commonly verruculose but sometimes verrucose,
pale brown to brown conidia measured 8.5–11.4 μm diam.
Cultural characteristics: Culture on MEA reaching
35 mm after 20 days at 25 °C. Fungal colony appeared cottony with abundant white to orange-white aerial mycelium.
In reverse colony dark olive to dark olivaceous-grey with
concentric dark olivaceous-brown. Hyphae hyaline sometime greyish green, smooth and verruculose later becomes
brown orange.
Material examined: India, Karnataka, Mysore, Doddamaragowdanahally, on foliar lesions of cowpea (Vigna
unguiculata (L.) Walp.- Fabaceae) 18 May 2020, S.
Mahadevakumar, Y.S. Deepika, N. Lakshmidevi (Specimen UOM-IOE 18/21), living culture (MD2).
GenBank numbers: OM811496 (ITS); OM811504
(LSU)
Notes: Detail descriptions and illustrations are presented
in Mason and Ellis (1953) and Markovskaja and Kačergius
(2014). Morphological inspection and measurements of conidiophores and conidia revealed that the fungal specimens
described by Markovskaja and Kaergius (2014) for several
Apiaceae hosts correspond well with the P. byssoides, which
was based on lectotype material (Fries 1832). Phylogenetic
analyses (Fig. 49) This is the first time that P. byssoides is
reported from Fabaceae on Vigna unguiculata representing
a new host record and geographical record (Table 3).
Periconia cortaderiae Thambugala & K.D. Hyde, in Thambugala et al., Mycosphere 8(4): 734 (2017)
Index Fungorum number: IF553165; Facesoffungi number: FoF03226; Fig. 50
Saprobic on decaying leaves of Musa basjoo. Sexual
morph: Not obersved. Asexual morph: hyphomycetous.
Colonies on host black, powdery, conidial masses are clearly
visible on the host. Mycelium sub hyaline to pale brown
or brown branched, having black conidial clusters Conidiophores 60–130 × 3 − 4 μm (x̄ = 76.5 × 3.5 μm, n = 15),
macronematous, mononematous, appear as single or a cluster, erect, rough-walled, sub hyaline, brown to dark brown,
septate at some points, significantly branched and flexuous.
Conidiogenesis can be observed at the apices of the branches
or from the middle of the conidiophores. Conidiogenous
69
cells 3.5–4.5 μm × 2.5–3.5 μm (x̄ = 3.5 × 2.8 μm, n = 10),
annellidic, monoblastic, discrete on the stipe, percurrent proliferations present as scars at the apex of the conidiophore.
Conidia 4–8 × 4–7 μm (x̄ = 6.8 × 6.2 μm, n = 40), appear as
chains or single, globose, one-celled, immature conidia are
hyaline to pale brown, mature conidia are brown to dark
brown, smooth or minutely verruculose, thick- walled.
Culture characteristics: Conidia germinating on PDA
within 36–48 h. Colonies on PDA, reach 15 mm after
18 days at 25 °C, at maturity, unevenly distributed radial furrows or linear marks were observed, surface notably rough
at maturity with crenulate to crenate margin, the colony
is completely black and powdery at maturity, moderately
dense, reverse white to black.
Material examined: Russia, Krasnodar region, Sochi,
Khstinsky City District, M.V. Frunze Health Care Resort,
park, on a dying leafstalk of Musa basjoo Siebold & Zucc.
ex Iinuma (Musaceae), 15 October 2018, Timur S. Bulgakov, BNR-001 (MFLU 18–1896), living culture MFLUCC
22-0178.
Known hosts: Cortaderia sp. (Poaceae, Monocotyledon) (Thambugala et al. 2017); on Caragana arborescens
(Fabaceae, Dicotyledon) (Phookamsak et al. 2019); on Musa
sp. (Musaceae, Monocotyledon) (Samarakoon et al. 2021
and this study).
Distribution: From Russia (This study), from Thailand
(Thambugala et al. 2017, Samarakoon et al. 2021; this
study), from Yunnan, China (Phookamsak et al. 2019)
GenBank numbers: OP097674 (LSU), OP099358 (SSU),
OP099551 (ITS), OP113822 (tef1)
Notes: Based on BLASTn search results of SSU, LSU,
ITS sequence data, MFLUCC 22-0178, showed high similarity to Periconia cortaderiae (MFLUCC 18-0668) as
follows; SSU = 99.76%, LSU = 99.88%, ITS = 99.80%.
Our new collection is similar to the holotype of P. cortaderiae (Thambugala et al. 2017), except the length and the
shape of the conidiophores. Our collection has short conidiophores with respect to the holotype (60–130 × 3–4 μm
vs. 400–800 × 4–9.4 μm). The conidiophore of our strain is
notably curved and branched compared to other collections.
The conidiogenesis is also found as terminal and intercalary
on the conidiophores in our strain. The other collections of
P. cortaderiae only had terminal conidial formation with
respect to our finding. There is no significant difference
in the nucleotide base pair comparison of our strain with
the ex-type strain. In our multigene phylogeny, MFLUCC
22-0178 grouped with P. cortaderiae (MFLUCC 15-0451,
MFLUCC 18-0668, MFLUCC 20-0236) with strong statistical support (ML = 100%, BYPP = 1.00). In this study, we
identify our new collection as P. cortaderiae, from Musa
sp. (Monocotyledon), from Russia for the first time as a new
geographical record.
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13
Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
◂ Fig. 49 Maximum likelihood tree revealed by RAxML from an analy-
sis of a concatenated SSU, LSU, ITS and tef1 sequence dataset of the
species in Periconiaceae, showing the phylogenetic position of Periconia delonicis (MFLUCC 20–0235). Bootstrap supports ≥ 60% and
BYPP ≥ 0.95 are given above the branches as ML/BYPP. The tree is
rooted with Helminthosporium dalbergiae (MAFF 243853) and Massarina cisti (CBS 266.62). Strains generated in this study are indicated in red bold. Ex–type strains are indicated in black bold. The
scale bar 0.02 represents the expected number of nucleotide substitutions per site
Tubeufiales S. Boonmee & K.D. Hyde.
Notes: Tubeufiales includes three families viz. Bezerromycetaceae, Tubeufiaceae and Wiesneriomycetaeae with 56
genera (Wijayawardene et al. 2022).
Tubeufiaceae M.E. Barr, Mycologia 71(5): 948 (1979).
Notes: Barr (1979) defined Tubeufiaceae based exclusively on the generic type, Tubeufia, and treated it in Pleosporales. Based on unique morphology and multigene phylogenetic investigations, Boonmee et al. (2012) formed the
order Tubeufiales and designated Tubeufiaceae as the typical
family. Previously, 19 genera were compared in Tubeufiaceae (Boonmee et al. 2012). Following that, Lu et al. (2018)
reappraised Tubeufiaceae and accepted 43 taxa to this family
based on phylogenetic analyses and morphological evidence.
With the expansion of fungal investigations, the family
Tubeufiaceae presently contains 47 genera (Lu et al. 2018;
Liu et al. 2019a, b, c, d; Wijayawardena et al. 2022).
Helicoma Corda, Icon. fung. (Prague) 1: 15 (1837)
Notes: Helicoma was introduced by Corda (1837) who
treated H. muelleri as the type species. It is one of the earliest described helicosporous hyphomycete genera (Liu et al.
2019a, b, c, d). Helicoma species have been found in tropical and temperate locations (Boonmee et al. 2012, 2014;
Lu et al. 2018; Brahmanage et al. 2017), and there are 59
records in the genus (Lu et al. 2018; Liu et al. 2019a, b, c,
d; Barreto et al. 2021).
Helicoma aquaticum Y.Z. Lu, J.C. Kang & K.D. Hyde, in
Lu et al. Fungal Diversity 92: 174 (2018).
Index Fungorum: 554836; Facesoffungi number:
FoF04712; Fig. 51
Saprobic on decaying fruits of Dipterocapus sp. Asexual morph: Hyphomycetous, helicosporous. Colonies on
the substratum superficial, gregarious, brown. Mycelium
composed of partly immersed, partly superficial, hyaline to
pale brown, septate, hyphae. Conidiophores macronematous,
mononematous, cylindrical, erect, straight, unbranched,
septate, 119–180 μm long, 6.5–7 μm wide, the lower part
brown to dark brown, the upper part pale brown, smoothwalled. Conidiogenous cells holoblastic, mono- to polyblastic, integrated, intercalary, cylindrical, with denticles,
71
arising laterally from the lower part of conidiophores,
10–15 × 5–9 μm (x̅ = 12 × 7 μm, n = 20), brown, smoothwalled. Conidia solitary, pleurogenous, helicoid, tapering
towards apex, rounded at tip, 33–37 μm diam. and conidial
filament 3.5–6 μm wide (x̅ = 5 μm, n = 20), 290–376 μm long
(x̅ = 321 μm, n = 20), 20–27-septate, constricted at septa,
coiled 11/2–41/4 times, becoming loosely coiled or uncoiled
in water sometimes, hyaline to pale brown, smooth-walled.
Sexual morph: Not observed.
Culture characteristics: Colonies growing on PDA at
25 °C, edge undulate, flat, circular, spreading, with fluffy
pale brown air-mycelium, pale brown to brown mycelium.
reverse dark brown, pale brown at the edge side, lobate at
the center, without pigmented.
Material examined: Thailand, Chiang Mai Province,
MRC, on dead fruits of Dipterocarpus sp. (Dipterocarpaceae), 15 August 2019, Xia Tang, Dip 18 (MFLU
21-0176), living culture MFLUCC 21-0141.
GenBank numbers: OM232106 (ITS), OM248446 (LSU),
OM272846 (tef1)
Hosts: Unindentified submerged decaying wood (Lu et al.
2018), dead fruits of Dipterocarpus sp. (Dipterocarpaceae)
(this study).
Distribution: Thailand (Lu et al. 2018, this study).
Notes: Helicoma aquaticum was described and illustrated
by Lu et al. (2018) from an unidentified submerged decaying wood in Thailand. Based on the morphological characteristics, our collection shares similar morphology with
the ex-type strain (Fig. 52). Comparisons of ITS, LSU and
tef1 sequence data between our collection and the holotype
showed that they are 2/ 549 bp (0.3%) of ITS, 9/1201 (0.7%)
of LSU and 9/879 bp (1%) of tef1. We consider that our collection is the same as Helicoma aquaticum following the
guidelines for species delineation proposed by Jeewon and
Hyde (2016). This is the first report of Helicoma aquaticum
on a Dipterocarpus sp. in Thailand.
Wiesneriomycetaceae Suetrong, Rungjind., Somrith. &
E.B.G. Jones
Based on morphology and molecular phylogeny, Suetrong
et al. (2014) introduced Wiesneriomycetaceae as order incertae sedis. Pratibha et al. (2015) placed Wiesneriomycetaceae
in Tubeufiales. Bezerra et al. (2017) introduced Wiesneriomycetales to accommodate Wiesneriomycetaceae based on
morphological characteristics and phylogenetic analyses.
Based on phylogenetic inference and divergence times estimates, Liu et al. (2017) considered Wiesneriomycetales as
a synonym of Tubeufiales. Hongsanan et al. (2020b) and
Wijayawardena et al. (2020, 2022) accepted this family in
Tubeufiales.
Wiesneriomyces Koord., Verh. K. Akad. Wet., tweede section 13(4): 246 (1907).
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72
Fig. 50 Periconia cortaderiae (MFLU 18–1896, new host and geographical record). a − b Colonies on host c − f, i, j Conidiophores
bearing conidia g, h, k Conidiogenesis from terminal and intercalary
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Fungal Diversity (2022) 117:1–272
parts of the conidiophore l-p Conidial chains and conidia. Scale bars:
a = 3 mm, b = 3.5 mm, c = 50 μm, e − j = 15 μm, d, k − p = 5 μm
Fungal Diversity (2022) 117:1–272
73
Fig. 51 Helicoma aquaticum (MFLU 21–0176, new
host record) a, b Host. c, d
Conidiophores. e Conidiophore
with conidiogenous cells. f–h
Conidia. i Germinated conidium. Scale bars: b = 100 μm,
c–e = 50 μm, f–h = 10 μm
Notes: Wiesneriomyces species have setose, branched
conidiophores, single-chain hyaline conidia, with a short
isthmi separating the conidia (Suetrong et al. 2014; Hongsanan et al. 2020b). The sexual morph for this genus is not
yet reported.
Wiesneriomyces laurinus (Tassi) P.M. Kirk, Trans Br
Mycol Soc 82: 748 (1984)
Index Fungorum number: IF107371; Facesoffungi number: FoF09126, Fig. 53
Saprobic on dead leaves of Dimocarpus longan. Sexual
morph: Not observed. Asexual morph: Colonies effuse,
yellowish-brown. Conidiomata sporodochial, solitary,
3–10 setae arising from the margins of the sporodochial
stalk. Setae 140–300 × 3–8 µm (x̄ = 180 × 5 μm, n = 30),
subulate, apex acute, septate, thick-walled, deep brown,
arising on leaf surface. Conidiophores 30–50 × 2–4 µm
(x̄ = 40 × 3 μm, n = 21), emi-mucronematous, close to one
another, brown to sub-hyaline at the base, hyaline towards
the apex, septate, irregularly branched. Conidiogenous cells
6–9 × 2–4 µm (x̄ = 7 × 3 μm, n = 26), located at the conidiophores terminal, cylindrical, hyaline, integrated. Conidia
50–70 × 2–4 μm (x̄ = 60 × 3 μm, n = 30), solitary, clumped
together into a semi-mucus, hyaline, smooth, sandy, cylindrical, taper towards both ends that are obtusely rounded,
5–6-septate, prominently constricted at septa, median cells
6–10 × 2–4 µm long, terminal cells 6–10 × 2–4 µm.
Culture characteristics: Colonies on PDA, 37–42 mm
diam. after 3 weeks, colonies from above: medium dense,
flat, slightly raised, rough surface with irregular edges, fluffy
not smooth, leaden grey to light grey at the margin, black to
ollvnceous black in the centre; reverse: grey to light grey at
the margin, grey-brown to grey in the centre.
Material examined: China, Guangdong Province, Guangzhou City, Haizhu District, Zhongkai University of Agriculture Engineering, 23° 6′ 32″ N, 113° 16′ 37″ E, alt. 20 m, on
the dead leaves of Dimocarpus longan Lour. (Sapindaceae),
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74
Fig. 52 Phylogram generated from parsimony analysis based on combined ITS, LSU and tef1 sequence data of Helicoma. ML and MP
bootstrap support values ≥ 70% are indicated above the nodes, and
branches with Bayesian posterior probabilities ≥ 0.95 are given above
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Fungal Diversity (2022) 117:1–272
the nodes. The ex-types (reference strains) are in bold; the new isolates are in blue bold. The tree is rooted with Kamalomyces thailandicus (MFLUCC 13–0233) and Kamalomyces thailandicus (MFLUCC
11–0158)
Fungal Diversity (2022) 117:1–272
75
Fig. 53 Wiesneriomyces
laurinus (ZHKU 22–0008, new
host record) a–c Appearance of
sporodochia on host. d Squash
mount of sporodochium. e, f
Conidiophores with setae. g–i
Conidia. j Colonies on PDA.
Scale bars: d = 100 µm, e,
f = 20 µm, g, i = 25 µm
23 July 2021, YH. Yang & CF. Liao (ZHKU 22-0008); living cultures ZHKUCC 22-0008, ZHKUCC 22-0009.
Known hosts: Carissa carandas (Apocynaceae), Clusia rosea (Clusiaceae), Hibiscus elatus (Malvaceae), Ficus
ampelas (Moraceae), Gyranthera caribensis (Malvaceae),
leaf litter of Caesalpinia echinata (Fabaceae), Laurus nobilis (Lauraceae), Ocotea leucoxylon (Lauraceae), Pandanus
urophyllus (Pandanaceae), Psidium guajava (Myrtaceae),
Roystonea regia (Arecaceae), Talipariti elatum, Theobroma
cacao (Malvaceae), Thuja occidentalis (Cupressaceae),
Dimocarpus longan (Sapindaceae)
Distribution: Brazil (da Silva and Grandi 2008), China
(this study), Cuba (Delgado-Rodriguez et al. 2002), Hong
Kong (Lu et al. 2000; Zhuang 2001), Mexico (Begerow et al.
2018), Myanmar (Thaung 2008), Russia (Melnik and Popushoi 1992), United Kingdom (Dennis 1986), Venezuela
(Castaneda-Ruiz et al. 2003).
GenBank numbers: ZHKUCC 22-0008—OM780294
(LSU), OM780298 (SSU), OM780284(ITS)
ZHKUCC 22-0009—OM780295 (LSU), OM780306
(SSU), OM780286 (ITS)
Notes: Our two strains (ZHKUCC 22-008 and ZHKUCC
22-009) share similar characters with Wiesneriomyces
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76
Fig. 54 The best scoring RAxML tree for Wiesneriomycetaceae, for
combined dataset of LSU, SSU and ITS sequence data. The tree is
rooted with Bezerromyces brasiliensis (URM7411) and Bezerromy-
13
Fungal Diversity (2022) 117:1–272
ces pernambucoensis (URM7412). Ex-type strains are in bold and
newly generated sequences are in red. Bootstrap support values for
ML ≥ 50% and BYPP ≥ 0.95 are given above the nodes
Fungal Diversity (2022) 117:1–272
77
Fig. 55 Asterina brigadeirensis (VIC 44217, holotype) a
Colony with open thyriothecia
and surface mycelium. b Ascomata opened by a central starshaped fissure. c Cross section
of the ascomata. d Globose to
pyriform unicelular appressoria.
e Immature ascus with pseudoparaphyses. f Mature ascus. g
Immature ascospores. h Brown
and verruculose ascospore.
Scale bars: a = 200 µm,
b–c = 50 µm, d–e–f = 20 µm,
d = 10 µm, g–h = 10 µm
laurinus (Heredia et al. 2000; Rajashekhar and Kaveriappa 2000; Suetrong et al. 2014; Pratibha et al. 2015; Tennakoon et al. 2021). The phylogeny also showed that our
strains (ZHKUCC 22-008 and ZHKUCC 22-009) clustered with other Wiesneriomyces laurinus strains (closer to
BCC18609), with moderate statistical support (Fig. 54). This
is the first report of Wiesneriomyces laurinus on Dimocarpus longan.
Dothideomycetes orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Notes: The order was introduced by Hawksworth and
Eriksson (1986), based on the type species of the order,
Asterina melastomatis Léveillé, which had its DNA
extracted, sequenced and studied phylogenetically for the
first time by Guatimosim et al. (2015), demonstrating that
the Asterinales is polyphyletic. Based on molecular data
of the type species, Asterinales stricto sensu includes two
13
78
families, namely: Asterinaceae and Parmulariaceae (Guatimosim et al. 2015; Giraldo et al. 2017; Phookamsak et al.
2019; Johnston and Park 2019; Hongsanan et al. 2020a; Le
Renard et al. 2020; Firmino and Pereira 2021).
Asterinaceae Hansford, Mycol. Pap. 15: 188 (1946)
Notes: Asterinaceae was proposed by Hansford (1946),
undergoing several modifications over time in relation to the
genera belonging to the family. Asterinaceae is polyphyletic
and the Asterinaceae stricto sensu comprises the species
grouping with the type species, Asterina melastomatis.
Asterina Léveillé, Annls Sci. Nat., Bot., sér. 3 3: 59 (1845)
Notes: The genus was described by Léveillé (1845),
having Asterina melastomatis as the type species. The
genus is characterized by having circular shaped ascomata, opening by a star-shaped fissure, with absence of
hypostroma, adhering to the host by superficial hyphae
with lateral appressoria (hyphopodia), bitunicate asci disposed as an upright palisade layer, and 2–celled brownish
ascospores.
Asterina brigadeirensis A.L. Firmino & O.L. Pereira, sp.
nov.
Index Fungorum number: IF900066; Facesoffungi number: FoF13383; Fig. 55
Etymology: Name refers to the mountain range, where the
fungus was collected, Serra do Brigadeiro.
Holotype: VIC 44217
Sexual morph: Colonies amphigenous, circular to irregular, single to confluent, dark brown, black, 0.1–5 mm diam.
Hyphae straight to slightly flexuous, branching irregularly,
brown, septate, hyphal cells cylindrical, 4.5–7.5 μm diam.,
smooth. Appressoria numerous, entire to lobate, sessile, lateral, alternate to unilateral, never opposed, globose to pyriform, unicellular, straight to angular, 7.5–10 × 6–9.5 μm,
brown, penetration peg central on the appressorial cell. Ascomata superficial, thyriothecia, scutiform, on top of mycelial
mat, circular to ellipsoid, single to confluent, fringed at
margins, randomly distributed in the colony, 135–262.5 μm
diam., opening by a central star-shaped fissure to an irregular fissure, dark brown to black; wall of textura radiata to
irregulate on cells isodiametric to cylindrical. Pseudoparaphyses cylindrical, filiform, septate, unbranched, hyaline, up
to 2.5 μm wide. Asci bitunicate in structure, fissitunicate,
disposed as an upright palisade layer, ovoid to cylindrical,
8-spored, hyaline, 65–95 × 35–45 μm. Ascospores cylindrical to oblong, ends rounded, straight or slightly arched,
1-septate, constricted at the median septum, hyaline, becoming brown at maturity, verruculose, 30–35 × 12.5–16 μm.
Asexual morph: Not observed.
Material examined: Brazil, Minas Gerais, Araponga,
on living leaves of Miconia cinnamomifolia Naudin
13
Fungal Diversity (2022) 117:1–272
(Melastomataceae), 10 September 2014, A.L. Firmino (VIC
44217, holotype).
GenBank numbers: MZ475298 (LSU)
Notes: Asterina brigadeirensis differs from the species
previously reported on Melastomataceae (Léveillé 1845;
Hennings 1904, 1909; Theissen 1912, 1913; Sydow and
Sydow 1916; Yates 1917; Maublanc 1920; Ryan 1924, 1928;
Sydow 1927, 1930; Sydow and Petrak 1929; Chardón and
Toro 1930; Orejuela 1944; Petrak 1950; Hansford 1954;
Yamamoto 1957; Hosagoudar and Abraham 2000). It is closest to A. venezuelana, which has smaller and ovoid to conoid
appressoria, smaller and ovoid to clavate asci, and smaller
and dark brown ascospores. Asterina brigadeirensis is easily
separated from A. amadelpha, A. belluciae, A. centroniae, A.
chrysophylli, A. confertissima, A. hypophyla, A. maublancii,
A. melanotes, A. melastomatis, A. melastomatis-candidi, A.
memecylonicae, A. pulla, A. schlechteriana, A. sinsuieiensis,
A. uribei in having verruculose ascospores. Asterina antioquensis is distinct from the new species in having ovoid to
ellipsoid appressoria, smaller asci, and smaller ascospores
with an upper third septum. Asterina astroniae has narrow
hyphae, smaller and ovoid appressoria, smaller and subglobose to ovoid asci, and finally smaller ascospores. Asterina denigrata differs from Asterina brigadeirensis in the
hypophyllous colonies, smaller ascomata and asci with 2–6
spores, and smaller and dark brown ascospores. Asterina
hughesii differs in the narrow appressoria, smaller and spatulate asci, lacking pseudoparaphyses, and smaller ascospores.
Asterina melastomatacearum differs from Asterina brigadeirensis in the smaller asci, and smaller and ellipsoid-ovoidal
ascospores. Asterina madikeriensis differs in the colonies
epiphyllous, entire appressoria and smaller ascospores with
tuberculate wall. Asterina melastomaticola differs in the narrow hyphae, lacking pseudoparaphyses, and much smaller
ascospores with echinulate ornamentations. Asterina miconiae differs from Asterina brigadeirensis in the smaller and
cylindrical to subglobose appressoria, smaller and ellipticalclavate asci, and smaller ascospores. Asterina miconiicola
differs in the much smaller ascospores with an upper third
septum. Asterina theissenii differs in having narrow and
dark brown hyphae, cylindrical to hemispherical appressoria, lacking pseudoparaphyses, and much smaller asci and
ascospores. Asterina transiens differs from new species in
the cylindrical appressoria, smaller and elliptical asci, and
much smaller ascospores. Asterina venezuelana differs in
the narrow and dark brown hyphae, smaller and ovoid to
clavate asci, and smaller and dark brown ascospores. Finally,
Asterina lopi, described below differs from the new species in having smaller and cylindrical to globose appressoria, smaller and ovoid to subclavate asci, and much smaller
ascospores. (Léveillé 1845; Hennings 1904, 1909a; Theissen
1912, 1913; Sydow and Sydow 1916; Yates 1917; Maublanc 1920; Ryan 1924, 1928; Sydow 1927, 1930; Sydow
Fungal Diversity (2022) 117:1–272
79
Fig. 56 The phylogenetic tree was obtained by Bayesian inference
methods using the sequences of the LSU region. The posterior probability values are indicated at the nodes. Strain numbers are indicated
after species names. New sequence data is in bold and blue. The anal-
yses included 33 strains including representative genera of Asterinales stricto sensu and Asterinales lato sensu. The tree is rooted with
Venturia populina (CBS 256.38) and V. inaequalis (CBS 815.69)
(Pleosporales) as outgroup
and Petrak 1929; Chardón and Toro 1930; Orejuela 1944;
Petrak 1950; Hansford 1954; Yamamoto 1957; Hosagoudar
and Abraham 2000; Hosagoudar 2006). Based on our phylogenetic analyses (Fig. 56) and morphological analyses,
herein we introduce a new species, A. brigadeirensis. Asterina brigadeirensis is the tenth species of Asterina reported
on hosts belonging to Melastomataceae in Brazil, and the
twelfth on Miconia.
Sexual morph: Colonies epiphyllous, circular to irregular, single to confluent, black, 1–6 mm diam. Hyphae
straight to flexuous, branching unilaterally or irregularly, brown, septate, hyphal cells cylindrical, 4–4.5 μm
diam., smooth. Appressoria numerous, entire to irregularly lobate, sessile, lateral, alternate to unilateral, never
opposed, cylindrical to globose, unicellular, straight to
angular, 5–7.5 × 5–6 μm, brown, penetration peg central
on the appressorial cell. Ascomata superficial, thyriothecia, scutiform, on top of mycelial mat, circular to ellipsoid,
single to confluent, fringed at margins, randomly distributed in the colony, 112–212.5 μm diam., opening by a
central star-shaped fissure, dark brown; wall of textura
radiate to irregulata, cells isodiametric to cylindrical to
irregular. Pseudoparaphyses cylindrical, filiform, septate,
unbranched, hyaline, up to 2.5 μm wide. Asci bitunicate
Asterina lopi A.L. Firmino & O.L. Pereira, sp. nov.
Index Fungorum number: IF900067; Facesoffungi number: FoF13384; Fig. 57
Etymology: The name refers to the mountain range, where
the fungus was collected, Serra do Lopo.
Holotype: VIC 44219
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Fungal Diversity (2022) 117:1–272
Fig. 57 Asterina lopi (VIC
44219, holotype) a Colony
with open thyriothecia and
surface mycelium. b Ascoma
opened by a central star-shaped
fissure. c Cross section of
the ascoma. d Cylindrical to
globose unicelular appressoria.
e Immature ascus. f Mature
ascus. g Immature ascospores. h
Mature brown ascospores. Scale
bars: a = 200 µm, b = 50 µm,
c–f = 20 µm, d–e = 10 µm,
g–h = 5 µm
in structure, fissitunicate, disposed as an upright palisade layer, ovoid to subclavate, 8-spored, and hyaline,
35–55 × 20–25 μm. Ascospores cylindrical to oblong, ends
broadly rounded, straight, 1-septate, constricted at the
median septum, hyaline, becoming pale brown to brown at
maturity, verruculose, 12.5–15 × 6–7 μm. Asexual morph:
Not observed.
Material examined: Brazil, Minas Gerais, Extrema,
on living leaves of Miconia sp. (Melastomataceae), 24
13
December 2014, A.L. Firmino & S.R. Pacheco (VIC 44219,
holotype).
GenBank numbers: MZ475299 (LSU)
Notes: Twenty-seven species of Asterina have been
reported previously in association with living leaves of
melastomataceous hosts. Asterina guianensis on Miconia
guianensis from Costa Rica and Miconia mirabilis from
Puerto Rico and Virgin Islands, A. racemosae on Miconia
racemose from Puerto Rico, and A. tetrazygiae on Tetrazygia
Fungal Diversity (2022) 117:1–272
81
Table 4 Host, country and source details of Asterina species in Melastomataceae
Species on
Host
Source and country
A. amadelpha Syd
A. antioquensis (Toro) Garcés
Conostegia oerstediana
Miconia ciliate. M. milleflora and M. dodonaea
Astronia sp.
Bellucia sp.
Miconia cinnamomifolia
Sydow & Petrak (1929), Costa Rica
Chardón & Toro (1930), Colombia, Venezuela
Centronia excelsa
Arthrostemma campanulare
Miconia acinodendron, M. laevigata, M.
prasina and Henriettea succosa
Blakea sp.
Miconia racemosa
Melastomataceae member
Miconia sp.
Memecylon sp.
Melastomataceae member
Blakea sp. and Miconia granulosa
Melastoma sp.
Melastoma malabathricum
Miconia sp.
Melastoma candidum
Memecylon edule
Miconia sp. and M. rubiginosa
Miconia racemosa
Melastomataceae member
Clidemia dentata
Barthea formosana
Miconia sp.
Miconia candolleana
Miconia toroi and M. ciliate
Clidemia sp., C. bonplandii and C. hirta
Petrak (1950), Ecuador
Syd. & P. Syd. (1916), Brazil
Guatimosim et al. (2015), Brazil, Trinidad and
Tobago. Puerto Rico and the Virgin Islands
Sydow & Petrak (1929), Costa Rica
Hosagoudar & Abraham (2000), Puerto Rico
Berkeley ex Theissen (1912), Guatemala
This study, Brazil
Hosagoudar (2006), India
Maublanc (1920), Brazil
Sydow & Petrak (1929), Costa Rica, Colombia
Theissen (1913), Brazil
Hansford (1954), Indonesia
Léveillé (1845), Brazil
Yamamoto (1957), China
Ryan (1928), India
Theissen (1913), Brazil
Ryan (1924), Puerto Rico, Dominican Republic
Léveillé (1845), Bolivia
Sydow (1927), Costa Rica
Yamamoto (1957), China
Ryan (1924), Puerto Rico
Theissen (1913), Brazil
Chardón & Toro (1930), Colombia, Venezuela
Sydow (1930), Venezuela
A. astroniae H.S. Yates
A. belluciae Henn
A. brigadeirensis A.L. Firmino & O.L.
Pereira
A. centroniae Petr
A. confertissima Speg
A. chrysophylli Henn
A. denigrata Petr
A. hughesii Hosag. & T.K. Abraham
A. hypophyla Berk. ex Theiss
A. lopi A.L. Firmino & O.L. Pereira
A. madikeriensis Hosag
A. maublancii (G. Arnaud) Maubl
A. melanotes Syd
A. melastomatacearum (Henn.) Theiss
A. melastomaticola Hansf
A. melastomatis Lév
A. melastomatis-candidi W. Yamam
A. memecylonicae R.W. Ryan
A. miconiae Theiss
A. miconiicola R.W. Ryan
A. pulla Lév
A. schlechteriana Syd
A. sinsuieiensis W. Yamam
A. theissenii R.W. Ryan
A. transiens Theiss
A. uribei Toro
A. venezuelana Syd
sp., and Tetrazygia elaeagnoides from Puerto Rico and Virgin Islands, were described by Ryan (1924) and not used in
the comparisons for having an ostiolar opening, a characteristic that does not belong to Asterina.
Asterina lopi differs from the species previously reported
on Melastomataceae (Léveillé 1845; Hennings 1904, 1909;
Theissen 1912, 1913; Sydow and Sydow 1916; Yates 1917;
Maublanc 1920; Ryan 1924, 1928; Sydow 1927, 1930;
Sydow and Petrak 1929; Chardón and Toro 1930; Orejuela
1944; Petrak 1950; Hansford 1954; Yamamoto 1957; Hosagoudar and Abraham 2000; Hosagoudar 2006) in morphology and on host association (Table 4). It is morphologically
closer to A. melastomatacearum, which has larger and ellipsoid-ovoidal ascospores. Asterina lopi is easily separated
from A. amadelpha, A. belluciae, A. centroniae, A. chrysophylli, A. confertissima, A. hypophyla, A. maublancii, A.
melanotes, A. melastomatis, A. melastomatis-candidi, A.
H.S. Yates (1917), Philippines
Hennings (1904), Brazil
This study, Brazil
memecylonicae, A. pulla, A. schlechteriana, A. sinsuieiensis,
and A. uribei in having verruculose ascospores; A. antioquensis, A. madikeriensis, A. miconiae, A. transiens and A.
venezuelana in having lobate appressoria. Asterina astroniae
is distinct from the new species in having ovoid appressoria, ascomata with irregular fissure, wider asci, and larger
ascospores. Asterina denigrata has hypophyllous colonies,
wider and dark brown hyphae, dark brown and subglobose
to conoid appressoria, wider and ovoid to ellipsoid asci with
2–6 spores, and larger and dark brown ascospores. Asterina
hughesii differs from Asterina lopi in the wider hyphae,
larger appressoria, narrow and spatulate asci, lacking pseudoparaphyses, and larger ascospores. Asterina melastomaticola differs in the pulvinate to conoid appressoria, globose to
ovoid asci, lacking pseudoparaphyses, and larger ascospores.
Asterina miconiicola differs from Asterina lopi in the wider
hyphae, larger appressoria, and narrow ascospores with
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an upper third septum. Asterina theissenii differs in having narrow and dark brown hyphae, globose to ovoid asci,
lacking pseudoparaphyses, and larger ascospores (Léveillé
1845; Hennings 1904, 1909; Theissen 1912, 1913; Sydow
and Sydow 1916; Yates 1917; Maublanc 1920; Ryan 1924,
1928; Sydow 1927, 1930; Sydow and Petrak 1929; Chardón
and Toro 1930; Orejuela 1944; Petrak 1950; Hansford 1954;
Yamamoto 1957; Hosagoudar and Abraham 2000). Based on
our phylogenetic analyses (Fig. 56) and morphology, herein
we introduce a new species, A. lopi. Asterina lopi is the ninth
species of Asterina reported on hosts belonging to Melastomataceae in Brazil, and the eleventh on Miconia.
Botryosphaeriales C.L. Schoch, Crous & Shoemaker.
Notes: Botryosphaeriales was introduced to accommodate a single family Botryosphaeriaceae by Schoch et al.
(2006). Schoch et al. (2009a) accepted its position in the
Pleosporomycetidae. Later, Planistromellaceae was recognised as a distinct family within Botryosphaeriales (Minnis
et al. 2012) Phyllostictaceae was reinstated for Phyllosticta
(Wikee et al. 2013). Another three families were introduced
by Slippers et al. (2013), namely Saccharataceae for Saccharata, Aplosporellaceae for Aplosporella and Melanopsaceae for Melanops. Wyka and Broders (2016) introduced
Septorioideaceae for Septorioides. Yang et al. (2018) raised
Endomelanconiopsis and Pseudofusicoccum to familial status as Endomelanconiopsidaceae and Pseudofusicoccaceae,
respectively. Wijayawardene et al. (2018) accepted the above
nine families in the order. However, Phillips et al. (2019)
synonymised Endomelanconiopsidaceae with Botryosphaeriaceae, Pseudofusicoccaceae with Phyllostictaceae, and
Septorioideaceae with Saccharataceae based on morphology, phylogeny and evolutionary divergence times. Therefore, six families, i.e. Aplosporellaceae, Botryosphaeriaceae,
Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae, are phylogenetically proved in the order now
(Hongsanan et al. 2020a, b; Wijayawardene et al. 2020).
Aplosporellaceae Slippers, Boissin & Crous, Stud. Mycol.
76(1): 41 (2013)
Aplosporellaceae was introduced in Botryosphaeriales
with two genera Aplosporella and Bagnisiella (Slippers et al.
2013). Sharma et al. (2017) added Alanomyces as a new
genus in this family based on four loci phylogeny. Two genera are accepted in Aplosporellaceae viz. Alanomyces and
Aplosporella (Dissanayake et al. 2021a, b; Wijayawardene
et al. 2022).
Aplosporella Speg. Anal. Soc. cient. argent 10(4): 157
(1880).
Aplosporella was treated as a type genus of Aplosporellaceae in Botryosphaeriaceae (Crous et al. 2006; Schoch
et al. 2006; Damm et al. 2007; Liu et al. 2012). The genus
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was established by Spegazzini (1880) and typified by Aplosporella chlorostroma. Later, Aplosporellaceae was introduced as a new family by Slippers et al. (2013). Wijayawardene et al. (2014a, b, 2016, 2020, 2022) also confirmed
the phylogenetic placement and accepted Aplosporellaceae
in Botryosphaeriaceae. There are 342 epithets of Aplosporella listed in Index fungorum (2022a, b). Aplosporella
is characterized by muti-locular conidiomata, with hyaline
to brown, aseptate conidia and, filiform paraphyses (Damm
et al. 2007; Ekanayaka et al. 2016; Dissanayake et al. 2021a,
b; Hyde et al. 2021). Aplosporella species are not easy to
identify based on only morphology because of their wide
range of host and morphological similarities. Many species
were introduced based on their host occurrence and suggested that species in Aplosporella are not host-specific
(Damm et al. 2007; Fan et al. 2015; Ekanayaka et al. 2016;
Dou et al. 2017).
Aplosporella artocarpi Trakun., L. Lombard & Crous, in
Trakunyingcharoen et al., Persoonia 34: 91 (2014)
Index Fungorum number: IF810167; Facesoffungi number: FoF10747, Fig. 58
Saprobic on dead stems of Chromolaena odorata. Sexual morph: Not observed. Asexual morph: Conidiomata
(280–) 300–350 × 370–450 (–500) µm (x̄ = 300 × 415 µm,
n = 5), solitary, immersed to semi-immersed with 2–3
locules, globose to subglobose, black. Ostiole absent.
Peridium 45–50 (–60) µm wide, multi-layered, comprised
of dark brown cells of Textura angularis. Hamanthecium
3–5 µm wide, numerous, hyaline, aseptate, paraphyses.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 1–2 µm wide, holoblastic, cylindrical, hyaline.
Conidia 15–20 × 5–10 µm (x̄ = 17 × 7 µm, n = 15), aseptate,
rough-walled, granular appearance, hyaline to dark brown,
ellipsoidal, without appendages.
Culture characteristics: Conidia germinating on
PDA within 24 h, reaching 85 mm after 7 days at room
temperature, concentric, flat, irregular, rough surface,
greenish-grey.
Material examined: Thailand, Chiang Rai Province,
Doi Pui, on the dead stems of Chromolaena odorata (L.)
(Asteraceae), 10 July 2020, Zin Hnin Htet, SW23 (MFLU
22–0108), living culture MFLUCC 22-0010.
Known hosts and distribution: On asymptomatic twig of
Artocarpus heterophyllus (Moraceae) in Chiang Mai Province, Thailand (Trakunyingcharoen et al. 2015); on asymptomatic leaves of Stoechospermum marginatum (Dictyotaceae) and Caulerpa taxifolia (Caulerpaceae) in India (Sahoo
et al. 2021); on dead branches of Mangifera indica (Anacardiaceae) in China (Yang et al. 2022); on dead stems of
Chromolaena odorata (Asteraceae) in Chiang Rai Province,
Thailand (this study)
GenBank numbers: ON834371(LSU), ON823183(ITS)
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83
Fig. 58 Aplosporella artocarpi
(MFLU 22–0108, new host
record). a, b Appearance of
conidiomata on host substrate.
c Section through conidoma. d
Peridium. e Pseudoparaphyses.
f Conidia on the conidiogenous
cells. g-j Conidia. k Sporulation l–m Culture on PDA from
surface and reverse. Scale
bars. a = 1000 µm, b = 500 µm,
c = 100 µm, d = 30 µm,
e = 5 µm, f, l, m = 20 µm,
g,h,I,j,k = 10 µm
Notes: Aplosporella artocarpi was introduced by Trakunyingcharoen et al. (2015). The species has morphologically
similar to other Aplosporella species such as A. hesperidica,
A. prunicola, and A. thailandica in having dark-brown, multilocular conidiomata with aseptate, hyaline to dark-brown
conidia (Damm et al. 2007; Ekanayaka et al. 2016; Dissanayake et al. 2021a, b). In our phylogenetic analysis, Aplosporella artocarpi (MFLUCC 22-0010) is closely related
to Aplosporella artocarpi (CPC 22,791) with ML = 70%,
BYPP = 0.54. (Fig. 59). According to BLASTn result, the
closest match for the LSU sequence was Aplosporella
artocarpi (CPC 22791) with 99.83% similarity. The closest match for the ITS sequence was Aplosporella prunicola (CBS 121167) with 97.81% similarity. Furthermore,
comparisons of ITS region between our taxon, Aplosporella
artocarpi (MFLUCC 22-0010) and ex-type strain of A. artocarpi (CPC 22791) show one base pair difference (0.18%)
across 531 nucleotides. Aplosporella species can be found
on a wide range of hosts such as Anacardiaceae, Asteraceae,
Caulerpaceae, Cupressaceae, Dictyotaceae, Fabaceae, Gingkoaceae, Moraceae, Myrtaceae, Proteaceae, and Rosaceae
(Du et al. 2017; Mapook et al. 2020; Sahoo et al. 2021; Yang
et al. 2021). We collected Aplosporella artocarpi (MFLU
22-0108) from Thailand and reported here as a new host
record associated with Chromolaena odorata.
Botryosphaeriaceae Theiss. & Syd. [as 'Botryosphaeriacae'], Annls mycol. 16(1/2): 16 (1918)
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Fig. 59 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS, and tef1 sequence data for the Aplosporella. The combined dataset consists of 19 taxa from Aplosporella
and our taxon, Aplosporella artocarpi (MFLUCC 22–0010). Melanops tulasnei (CBS 116805 and CBS 116806) are used as outgroup.
The topology of the maximum likelihood analysis is similar to Bayesian analysis. Bootstrap support value for ML ≥ 60% and BYPP ≥ 0.95
are given above the branches. The ex-type strains are bold. The newly
generated sequence is indicated in bold and red
Notes: Botryosphaeriaceae was introduced by Theissen
and Sydow (1918) for three genera, Botryosphaeria, Phaeobotryon and Dibotryon. Initially, the family had been successively put into the orders Dothideales (Müller and von
Arx 1950) and Pleosporales (Luttrell 1955), until Schoch
et al. (2006) raised Botryosphaeriaceae to orderal status as
Botryosphaeriales. Over decades of taxonomic revisions
and updates based on morphology, the family has become
increasingly complex. Kirk et al. (2008) estimated that there
are 26 genera and 1517 species in the family, while Liu et al.
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Fungal Diversity (2022) 117:1–272
85
Fig. 60 Botryosphaeria dothidea (IFRD500–008, new geographic and habitat record)
a, b Appearance of ascomata
on host substrate. c Section of
ascomata. d Peridium. e Pseudoparaphyses and asci. f–i Asci.
j–l Ascospores. m–n Colony
on PDA (m from front, n from
reverse). Scale bars: a = 500 μm,
b = 200 μm, c, e = 100 μm,
d = 30 μm, f–i = 20 μm,
j–l = 10 μm
(2012) accepted 29 genera and approximately 1485 species.
Phillips et al. (2013) consider morphological characters
alone as inadequate to define genera or identify species, and
they detailed described 17 genera and 110 species which has
molecular data. Thereafter, Dissanayake et al. (2017) introduced six new genera and 85 new species/species combinations. So far, 22 genera are accepted in Botryosphaeriaceae
(Phillips et al. 2019; Hongsanan et al. 2020a).
Botryosphaeria Ces. & De Not., Comm. Soc. crittog. Ital.
1(fasc. 4): 211 (1863)
Notes: Botryosphaeria was introduced by Cesati and
De Notaris (1863) based on the type species B. dothidea.
However, as the type material was immature, B. dothidea was epitypified by Slippers et al. (2004) based on
morphology and phylogenetic data which combined ITS,
tef1 and tub2 genes. The sexual morphs are characterized
by brown to black, globose ascostromata, comprising a
botryose aggregate, or sometimes solitary, with a central
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86
ostiole, papillate or not, bitunicate, clavate asci, with a
short pedicellate and a small ocular chamber, intermixed
with hyphae-like, wide, septate pseudoparaphyses, and
hyaline, aseptate, fusoid to ovoid ascospores, with or without a mucilaginous sheath (Liu et al. 2012). The asexual
morphs of Botryosphaeria were reported as Dichomera,
Diplodia, and Fusicoccum (Crous and Palm 1999; Slippers et al. 2004; Crous et al. 2006). They are characterized
by uni- to multilocular pycnidial, frequently embedded in
stromatic tissue, holoblastic, hyaline, subcylindrical conidiogenous cells, with 1–2 percurrent proliferation, and
hyaline, aseptate, narrowly fusiform, or irregularly fusiform conidia, rarely forming a septum before germination,
smooth with granular contents (Slippers et al. 2004). So
far, 286 species names are recorded in Index Fungorum
(2022a, b), in which 30 species known from culture are
accepted in the genus.
Botryosphaeria dothidea (Moug.) Ces. & De Not., Comm.
Soc. crittog. Ital. 1(fasc. 4): 212 (1863).
See Species fungorum for synonyms.
Index Fungorum number: IF183247; Faces of fungi
number:FoF03512; Fig. 60
Saprobic on submerged wood of freshwater. Sexual
morph: Ascomata 300–500 μm diam., black, circular or
subglobose to globose, scattered, gregarious, uni- to multiloculate, immersed to erumpent on host tissue, with visible
black dots or papilla. Ostiole circular, central, papillate.
Peridium composed of two-layered locules, outer layer composed of dark brown or brown thick-walled cells of textura
angularis, inner layer composed of hyaline thin-walled cells
of textura angularis lining the locule. Pseudoparaphyses
2–4 μm wide, hyphae-like, septate. Asci 96–144 × 18–25 μm
( x̄ = 116 × 23 μm, n = 10), 8-spored, bitunicate, fissitunicate,
cylindrical to clavate, short pedicellatea, pically rounded
with an ocular chamber. Ascospores 21–32 × 10–14 μm
( x̄ = 26.3 × 11.7 μm, n = 40), biseriate, hyaline, aseptate,
fusoid to ovoid, sometimes with tapered ends, spindleshaped, thin-walled, smooth with granular contents. Asexual
morph: Not observed.
Cultural characteristics: Ascospore germinating on
PDA within 24 h. Colonies on PDA fast-growing, reaching
7–8 cm diameter in 50 days at 20–25 °C, with dense, hairy,
black mycelium on the surface, reverse black.
Material examined: China, Yunnan Province, a small
river of Puzhehei wetland, on dead submerged decaying
wood of unidentified plants, 23 June 2019, Hao Yang, p27
(IFRD500–008), living culture KUMCC 20–0186.
Known hosts and distribution: broad range of hosts and
wide geohraphical distribution (Farr and Rossman 2022)
GenBank numbers: MT559116 (LSU), MT559099 (ITS)
Notes: Botryosphaeria dothidea is the type species of
the genus, and was reported extensively from all around
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the world (Fries 1823; Arx and Müller 1954). It was epitypified by Slippers et al. (2004) based on morphology and
phylogeny. This taxon was collected from southwest China
and its ITS sequence data of our isolate are 100% identical
to the verified sequences of B. dothidea (MH992666 and
MH973592). Based on morphology and phylogenetic analyses (Fig. 61), we identify the strain as B. dothidea. It is a
new geographic record from China and a new habitat record
from freshwater.
Class Laboulbeniomycetes Engler
Laboulbeniales Lindau
Notes: This order includes more than 2000 species
described as obligate ectosymbionts on Arthropods. Wijayawardena et al. (2022) accepted three families viz. Ceratomycetaceae, Euceratomycetaceae and Laboulbeniaceae in
this order.
Laboulbeniaceae G. Winter [as 'Laboulbenieae'], Rabenh.
Krypt.-Fl., Edn 2 (Leipzig) 1.2: 918 (1886)
Notes: Goldman and Weir (2018) based on SSU rDNA
sequence data identified this family to be consist of essentially terrestrial and sexually reproducing taxa with simple or
compound endogenous antheridia. Santamaria and Pedersen
(2021) accepted 147 genera in this family.
Rhachomyces Thaxt., Proc. Amer. Acad. Arts & Sci. 30:
468 (1895) [1894]
Notes: Rhachomyces is quite numerous: with six species
described very recently, the number of accepted species in
the genus is now 91 (Rossi and Christian 2020; Rossi and
Leonardi 2020; Santamaria et al. 2020; Buyck et al. 2021).
This genus is characterized by a series of superposed, usually short cells forming an axis that remembers a spinal
column (hence the name, from Greek rachis = spine); these
cells produce laterally both sterile appendages of various
lengths and antheridial appendages, the latter ending with a
single antheridium consisting of a simple phialide. In mature
thalli the perithecia are usually found in an apical position
and more frequently are single, but can be two or even more
in a few species. The cells forming the outer wall of perithecia are arranged in four rows, each consisting of four unequal
cells. Most of the species of Rhachomyces occur on ground
beetles (Carabidae), but a few are associated with rove beetles (Staphylinidae) and two were found on small carrion
beetles (Leiodidae Cholevinae). A single sequence is available for species in this genus (Goldmann and Weir 2018).
Rhachomyces cruralis W. Rossi & M. Leonardi, sp. nov.
Index Fungorum number: IF559505; Facesoffungi number: FoF13385; Fig. 62
Etymology: From Latin crus = leg, because the thalli of
the new species are found only on the legs of the host insect.
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87
Fig. 61 Phylogram generated from maximum likelihood analysis
based on combined ITS, and tef1 sequence data for the Botryosphaeria. The combined dataset consists of 23 taxa from Botryosphaeria
with our strain. The tree is rooted with Macrophomina phaseolina
(CBS 227.33). The topology of the maximum likelihood analysis is
similar to Bayesian analysis. Bootstrap support value for ML ≥ 60%
and BYPP ≥ 0.90 are given above the branches. The ex-type strains
are in bold. The newly generated sequence is in red
Holotype: FI WR1997
Axis of the receptacle is straight or slightly curved, consisting of 10–15 brownish cells gradually increasing in size
and bearing a dark brown band in the lower portion. Appendages almost straight, more numerous in the upper portion
of the thallus, usually consisting of 5–6 dark brown cells
separated by back septa and slight constriction, the distal one
being distinctly longer and gradually paler, with the lower
portion slightly inflated and the tip hyaline or almost so.
Antheridial appendages very few, consisting of a short and
brownish lower cell followed by a straight hyaline antheridium. Perithecium reddish-brown, subsessile, symmetrically
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Fig. 62 Rhachomyces cruralis (FI WR1997, holotype). Scale
bar = 50 µm
Fungal Diversity (2022) 117:1–272
Fig. 64 Rhachomyces magrinii (FI WR4049, holotype). Scale
bar = 100 µm
of Pachydesus rufipes (Boheman) (Carabidae, Trechini) (FI
WR1997, holotype). Same data as the type, M. Zapparoli
legit (FI WR1987 and WR 2000, paratypes); same data as
the type, A. Vigna Taglianti legit, FI WR1998 and WR1999,
paratypes).
Notes: The species is more similar to Rhachomyces
cruralis and R. moreti W. Rossi et Proaño, parasitic on
Trechisibus calathiformis Deuve from Ecuador. The latter
fungus, however, has an oblong perithecium, the axis of
the receptacle consists of 16–18 cells, and the appendages
are paler and slenderer (Rossi and Proaño Castro 2009).
Fig. 63 Rhachomyces hyperommae (FI WR2331, holotype). Scale
bar = 50 µm
elliptical, about twice as long as it is broad, regularly tapering to the darker, subconical tip and almost hyaline, rounded
apex. Length from foot to perithecial apex 270–350 µm.
Perithecium, including basal cells 125–175 × 60–80 µm.
Longest appendages 100 µm.
Material examined: South Africa, E Transvaal, Mt.
Sheba Nat. Res., 14–15.II.1995, S. Zoia, on posterior legs
13
Rhachomyces hyperommae W. Rossi & M. Leonardi, sp.
nov.
Index Fungorum number: IF559509; Facesoffungi number: FoF13386; Fig. 63
Etymology: Named after the host insect genus.
Holotype: CAMB WR2331a
Axis of the receptacle composed of 12–17 cells very different in size, shape, and color: the basal and the suprabasal
are blackened, narrow, and elongate; the following 2 or 3 are
relatively small, dark and isodiametric; the others gradually
broader and paler from below upwards. Appendages short,
stiff, one-sided, projected obliquely upwards, consisting of
4 cells, of which the lower 3 are dark brown while the upper
has a paler and curved tip. Antheridial appendages similar
to the sterile ones but shorter and paler, each bearing distally a brown, elongate antheridium tapering in a curved and
truncate paler tip. Perithecium broadly fusiform, with the
posterior side more convex, nearly hyaline or tinged with
Fungal Diversity (2022) 117:1–272
89
pale yellow, brownish yellow near the base in older specimens, the hyaline and conical tip ending in a blunt apex.
Length from foot to perithecial apex 280–380 µm. Perithecium, including basal cells 110–145 × 35–42 µm. Antheridia
20–25 µm. Longest appendage 90 µm.
Material examined: Autralia, NSW Border Ranges N. P.,
Tweed Range Rd., 4.6 km SW of Bridle Ck. Rd., alt. 580 m,
4.II–9.IV.1993, M. Gray & G. Cassis, on a femur and the
abdomen of Hyperomma sp. (Staphylinidae, Paederinae,
Paederini, Cryptobiina), CAMB WR2331a, holotype; FI
WR2331b isotype).
Notes: The new species seems to be allied to Rhachomyces arbusculus Thaxt., described on an unidentified rovebeetle from Liberia, West Africa (Thaxter 1896). The two
parasites have similar appendages and are also somewhat
similar in the general habitat. However, R. abusculus is distinctly more slender and more elongate, with the cells forming the receptacle more numerous (20–25) and not strongly
different from each other, and with a much narrower perithecium suffused with brown at the apex.
Rhachomyces magrinii W. Rossi & M. Leonardi, sp. nov.
Index Fungorum number: IF559510; Facesoffungi number: FoF13387; Fig. 64
Etymology: Named after the entomologist Paolo Magrini,
who supplied us with the material utilized for the description
of the new species.
Holotype: FI WR4048
Axis of the receptacle from almost straight to distinctly
curved or slightly sigmoid, consisting of (14)18–29 cells;
basal and suprabasal cells slender, elongate and dark brown
colored, the others gradually larger and paler. Appendages
occurring in the lower part of the thallus are slender, consisting of 5–6 brownish, elongate cells; these appendages
are visible only on young thalli, but are broken off in very
early stages of development. The upper and more lasting
appendages are thicker, generally consisting of 5 relatively
short, subequal cells, the series ending with a 6th cell distinctly longer and much paler, with an almost hyaline tip.
Antheridial appendages consist of a single, pale brown cell
bearing apically a hyaline antheridium with a tapering and
distinctly curved efferent neck. Perithecium subsessile, pale
grayish brown, more or less bent as to the axis of the receptacle, symmetrical or slightly asymmetrical, about three
times longer than broad, slightly more inflated below, the
truncate-conical and paler tip ending in a broad and blunt
apex. Length from foot to perithecial apex 525–950 µm.
Perithecium 135–180(200) × 45–60(70) µm. Antheridial
appendages 40–45 µm. Longest sterile appendages 150 µm.
Material examined: Mexico, Querétaro, Pinal de Amoles,
Ojo de Agua, 9.III.2011, G. Trezzi, on Mexaphaenops elegans Barr (Carabidae, Trechini) (FI WR4048, holotype; FI
WR4049, WR4488, paratype).
Fig. 65 Rhachomyces platyprosophi (FI WR3973a, holotype). Scale
bar = 100 µm
Notes: The only previously described species in Rhachomyces on a cave-dwelling Trechini from Central America is R. quetzalcoatl Balazuc, occurring on Paratrechus
spp. from Mexico and Guatemala. The two species of
fungi share the absence of dark pigmentation on the perithecial tip and the structure of the upper appendages, but
differ greatly in other characteristics, as the much shorter
receptacle of R. quetzalcoatl, consisting of dozens of cells,
and its more inflated perithecium (Rossi and Cesari-Rossi
1977, Fig. 63).
Rhachomyces platyprosophi W. Rossi & M. Leonardi, sp.
nov.
Index Fungorum number: IF559511; Facesoffungi number: FoF13388; Fig. 65
Etymology: Named after the host insect genus.
Holotype: FI WR3973.
Axis of the receptacle from almost straight to variably
curved or sigmoid, consisting of 15–22 brownish cells
gradually enlarging upwards and separated by oblique septa.
Appendages spreading, nearly opaque except for the gradually paler and curved tip, longer and more numerous in the
upper portion of the receptacle, reaching and sometimes
exceeding the perithecial apex. Antheridial appendages
relatively numerous, chestnut brown colored, consisting
of a slender and elongate cell followed by a very slender
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antheridium with a paler and sigmoid tip. Perithecium long
and slender, subsessile, oblong, light brown colored, the tip
gradually tapering, hardly distinguished except for the darker
color, the apex hyaline and subtruncate. Length from foot
to perithecial apex 500–830 µm. Perithecium 190–290 ×
50 µm. Antheridial appendages 50–65 µm. Longest sterile
appendage 520 µm.
Material examined: Indonesia, Sumatra, Palembang,
s. d., s. c. (from the collection of A. Fauvel in the Institut
Royal des Sciences Naturelles, Bruxelles), on abdomen and
legs of Platyprosopus indicus Motschulski (Staphylinidae,
Staphylininae, Platyprosopini) (FI WR3973a, holotype; FI
WR3973b and WR3973c isotypes).
Notes: Due to the large dimensions and the oblong perithecium, the new species can be compared with Rhachomyces carbonii W. Rossi & M. Leonardi, recently described
on a rove beetle from Sierra Leone, which however bears
a slender and much darker receptacle, shorter and slender
appendages and has perithecium with spirally twisted wall
cells with an abruptly distinguished tip (Rossi and Leonardi
2018).
Sexual morph: Thallus crustose, covering areas up to
20 cm diam., marginal 3 mm continuous, remaining parts
rather regularly cracked with areoles flat, c. 0.5 mm diam.,
not corticate, dull, almost pure white, very regularly c.
0.3 mm thick, not surrounded by a prothallus, but margin
thinner though at least 0.1 mm thick. Isidia and soredia
absent. Ascomata numerous, singly or aggregated, partly in
concentrical zones (especially the outer rim of apothecia at
3 mm from the margin), immersed, flush with the thallus,
rather uniformly 0.3–0.5 mm diam., sparse, solitary, disc
flat, grey, margin barely raised, c. 0.05 mm thick, white,
both disc and margin densely white pruinose. Excipulum
uniformly dark brown in section. Epihymenium a layer of
c. 15 µm high of brown paraphysal tips overlain with an c.
25 µm high epipsamma composed of many grey crystals that
do not dissolve in K (calcium oxalate). Hymenium 70–90 µm
high, hyaline, not inspersed, amyloid, paraphyses 1–1.5 µm
wide, with dark brown upper. Hypothecium dark brown, in
the central part up to 75 µm high, towards the margin tapering to c. 25 µm high. Ascospores 8/ascus, brown, 1-septate,
ellipsoid, 12–14 × 6–7 µm. Asexual morph: not observed.
Class Lecanoromycetes O.E. Erikss. & Winka
Subclass Lecanoromycetidae P.M. Kirk et al.
Caliciales Bessey.
Notes: Caliciales is an order of lichenized fungi, including
mostly crustose, but also foliose forms (Lücking et al. 2016)
Chemistry: Thallus and apothecia UV + patchily salmonorange, especially in the submarginal zone, C–, P–, K–.
TLC: nil.
Ecology and distribution: On exposed limestone in Atlantic rain forest biome; only known from Brazil.
Material examined: Brazil, Mato Grosso do Sul: Serra
da Bodoquena, Bodoquena, Dente de Cão, summit, alt.
450 m, 20° 47ʹ 05″ S, 56° 45ʹ 03″ W, on exposed limestone
in Atlantic rain forest biome, 7 November 2018, Aptroot
77815 (holotype, CGMC).
GenBank number: MW322683 (ITS).
Notes: This species is not keyed out in Malme (1912).
It shows similarities with species of the Buellia subalbula
(Nyl.) Müll. Arg. aggregate (Bungartz et al. 2011) and seems
most similar to Buellia amabilis de Lesd. (see Bungartz and
Nash 2004), which differs by the ornamented ascospores and
the apothecia that become more convex, and the absence of
UV luminescence. Phylogenetically, it clusters deep inside
Buellia De Not. in the current sense (Fig. 66). Sequences of
other species of the B. subalbula aggregate are not available.
As part of a continuous effort to explore lichenologically relatively unidentified regions in Brazil, we investigated the microlichens on an isolated limestone outcrop
in a tropical south-western inland region, viz. the Dente
de Cão in the Parque Nacional da Serra da Bodoquena in
the state of Mato Grosso do Sul, close to the borders with
Paraguay and Bolivia. This range of hills is the only larger
forested area still in existence in this state. Many microlichens were recently reported from the area (Aptroot and
Spielmann 2020). This area is close to the venue of IAL9,
the nearby town of Bonito.
Caliciaceae Chevall. [as 'Calicineae'], Fl. gén. env. Paris
(Paris) 1: 314 (1826)
Notes: Caliciaceae is a family of lichenized fungi, including mostly crustose, but also foliose forms (Lücking et al.
2016)
Buellia De Not., G. bot. ital. 2(1.1): 195 (1846)
Notes: Buellia is a large, heterogeneous and probably a
polyphyletic genus. Some species groups are relatively welldefined and may warrant formal recognition. This includes
the Buellia subalbula (Nyl.) Müll. Arg. aggregate (Bungartz
et al. 2011). Below we describe a new species that belongs
to this group.
Buellia pruinocalcarea Aptroot, M.F. Souza & Spielmann,
sp. nov.
Index Fungorum number: IF900068; Facesoffungi number: FOF13389; Fig. 66
Saxicolous Buellia on limestone with thallus thick, white,
apothecia flush with the thallus, densely white pruinose, and
ascospores 12–14 × 6–7 µm.
Holotype: Aptroot 77815.
Etymology: Named after the white pruina and the calcareous habitat.
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91
Fig. 66 Buellia pruinocalcarea
(Aptroot 77815 holotype) a.
habitat. b. Upper and lower left
in daylight. c. lower right in
365 nm UV light. Scale bars:
a-c = 20 mm
The natural vegetation of the Serra da Bodoquena is
the Atlantic rain forest, a biome that stretches all along
the coast from north-eastern to south-eastern Brazil. In
fact, it is the most western patch of Atlantic rainforest in
existence, and as such unique. Exposed limestone outcrops
occur in various places in the tropics, but their extent is
often limited and they tend to be soon grown over by vegetation. The Dente de Cão is formed of white Precambrian
(neoproterozoic, see Boggiani 1997) limestone known as
the Bocaiana formation, and reaches 450 m alt. It weathers into karst and the name of the rock outcrop is after the
sharp tooth-like rock points. Somewhat to our surprise,
the exposed limestone outcrop, although only scarcely
colonized by fanerogams, was not completely covered by
lichens. Cyanobacteria were more abundant. The most
common lichens were the species of Collemataceae and
Lichinaceae (Fig. 67).
Lecanorales Nannf.
Notes: Lecanorales is the largest order of lichenized fungi,
one of the largest containing crustose lichens and the largest
containing foliose lichens (Lücking et al. 2016).
Lecanoraceae Körb. [as 'Lecanoreae'], Syst. lich. germ.
(Breslau): 104 (1855).
Notes: Lecanoraceae is a family of lichenized fungi, one
of the largest containing crustose lichens (Lücking et al.
2016).
Lecanora Ach., in Luyken, Tent. Hist. Lich.: 90 (1809).
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Fungal Diversity (2022) 117:1–272
Fig. 67 Phylogram generated from maximum likelihood analysis based on ITS.
Bootstrap support values for
ML ≥ 80% and Bayesian posterior probabilities ≥ 0.95 are
given near nodes respectively.
The tree is rooted in Diplotomma rivas-martinezii (13365
BA). Ex-type strains are in bold.
The newly generated sequences
are indicated in bold blue
Fig. 68 Lecanora immersocalcarea (Aptroot 77822, holotype). Habitat. Scale bar = 23 mm
Notes: Lecanora is a large, heterogeneous and possibly
paraphyletic genus. Some species of this group are welldefined. Below we describe a new species in the somewhat
aberrant Lecanora marginata (Schaer.) Hertel & Rambold
group (Rambold 1989).
Lecanora immersocalcarea Aptroot, M.F. Souza & Spielmann, sp. nov.
13
I n d ex Fu n g o r u m n u m b e r: I F 9 0 0 0 6 9 ;
Facesoffungi:FOF13390; Fig. 68
Saxicolous Lecanora on limestone with thallus 0.5–0.8(2.0) mm thick, very pale ochraceous white, apothecia
immersed in the thallus, black, immature.
Etymology: Named for the immersed apothecia and the
calcareous habitat.
Holotype: Aptroot 77822.
Thallus crustose, superficial, covering areas up to 10 cm
diam., dull, cretaceous, very pale ochraceous white, regularly cracked with the areoles angular, slightly convex and
c. 0.5–0.9 mm diam., c. 0.5–0.8 mm thick but in some
places with lobate superficial outgrowths of up to 2 mm
thick, not surrounded by a prothallus, but margin thick
and raised, sharply delimited, not lobed or fissured. Isidia
and soredia absent. Ascomata immature apothecia, several
per areole, flush with the thallus, rounded, 0.2–0.4 mm
diam.; disc black, margin thin, black. Epihymenium grey,
without crystals. Excipulum without crystals. Hypothecium hyaline. Mature ascospores not observed. Pycnidia
not observed.
Chemistry: Thallus UV–, C–, P + yellow, K + yellow.
TLC: atranorin.
Distribution: On exposed limestone outcrop in Atlantic
rain forest biome; only known from Brazil.
Material examined: Brazil. Mato Grosso do Sul: Serra da
Bodoquena, Bodoquena, Dente de Cão, summit, alt. 450 m,
20° 47ʹ 05″ S, 56° 45ʹ 03″ W, on exposed limestone in an
Fungal Diversity (2022) 117:1–272
93
Fig. 69 Phylogram generated
from maximum likelihood analysis based on ITS. Bootstrap
support values for ML ≥ 80%
and Bayesian posterior probabilities ≥ 0.95 are given near nodes
respectively. The tree is rooted
in Tephromela atra (L1489).
Ex-type strains are in bold. The
newly generated sequences are
indicated in bold blue
Atlantic rain forest biome, 7 November 2018, Aptroot 77822
(holotype, CGMC).
GenBank number: MW322682 (ITS)
Notes: This species is locally abundant. Fully fertile material was not found, but the type specimen was sequenced and
clustered inside Lecanora Ach. in the current sense. Morphologically it would belong to the Lecanora marginata
(Schaer.) Hertel & Rambold group, in which indeed calciferous species or at least specimens are known. Only one
described species seems close, however, viz. Lecanora oreinodes (Körb.) Hertel & Rambold (Rambold 1989), which
differs by the flatter, flush areoles and the somewhat fractured/lobate thallus margin, and which is not known to occur
on pure limestone. Phylogenetically, it clusters deep inside
Lecanora, where it clusters with several species with usnic
acid instead of atranorin, but with low support (Fig. 69).
Sequences of other species of the L. marginata group are
not available. This species is described from the same locality as Buellia pruinocalcarea (for details see under Buellia
pruinocalcarea).
Teloschistales D. Hawksw. & O.E. Erikss.
Notes: Teloschistales is a large order of lichenized fungi,
one of the largest containing crustose lichens (Arup et al.
2013).
Teloschistaceae Zahlbr. [as 'Theloschistaceae'], in Engler,
Syllabus, Edn 2 (Berlin): 45 (1898).
Fig. 70 Wetmoreana blastidiocalcarea (Aptroot 77,806, holotype).
Habitat. Scale bar = 17 mm
Notes: Teloschistaceae is a family of lichenized fungi,
one of the largest containing crustose lichens. Most species
are generally still treated under the genus Caloplaca Th. Fr.
(Schumm and Aptroot 2019).
Wetmoreana Arup, Søchting & Frödén, in Arup, Søchting
& Frödén, Nordic Jl Bot. 31(1): 66 (2013)
Notes: Wetmoreana is a recently split genus in Teloschistaceae (Arup et al. 2013). At the moment, three to five
species are recognized in this genus, depending on whether
or not Fulgogassparrea S.Y. Kondr., N.-H. Jeong, Kärnefelt,
Elix, A. Thell & Hur (Kondratyuk et al. 2013) is recognized
as a separate genus.
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Fungal Diversity (2022) 117:1–272
Fig. 71 Phylogram generated
from maximum likelihood analysis based on ITS. Bootstrap
support values for ML ≥ 80%
and Bayesian posterior probabilities ≥ 0.95 are given near
nodes respectively. The tree is
rooted in Xanthoria parietina
(pop4_26). Ex-type strains are
in bold. The newly generated
sequences are indicated in bold
blue
Wetmoreana blastidiocalcarea Aptroot, M.F. Souza &
Spielmann sp. nov.
Index Fungorum number: IF900070; Facesoffungi number: FOF13391; Fig. 70
Saxicolous Wetmoreana on limestone with thallus thick,
yellow-orange, radially lobate, central part of the thallus
areolate, covered by c. 0.05 mm diam, semiglobose corticate
bulbs of thallus color that might serve as blastidia.
Etymology: Named after the calcareous and the blastidious habitat.
Holotype: Aptroot 77806.
Thallus placodioid, up to 2 cm diam., corticate, dull,
yellow-orange, white pruinose on the marginal lobes, up
to 0.3 mm thick. The central part of the thallus areolate,
covered by c. 0.05 mm diam, semiglobose corticate bulbs
of thallus colour that might serve as (and would in former
species of Fulgensia usually being called) blastidia, sometimes partly dissected into secondary lobes resembling the
marginal lobes. Marginal lobes much radially divided into
less than 0.1 mm wide lobuli, gradually thinning towards the
margin. Isidia and soredia absent. Ascomata and pycnidia
not observed.
Chemistry: Thallus and apothecia UV + red, C−, P−,
K+ crimson. TLC: anthraquinones.
Distribution: On exposed limestone in Atlantic rain forest
biome; only known from Brazil.
13
Material examined: Brazil. Mato Grosso do Sul: Serra
da Bodoquena, Bodoquena, Dente de Cão, summit, alt.
450 m, 20° 47ʹ 05″ S, 56° 45ʹ 03″ W, on exposed limestone
in Atlantic rain forest biome, 7 November 2018, Aptroot
77806 (holotype; CGMC).
GenBank number: MW322681 (ITS).
Notes: Morphologically, this species shows similarities
with several genera in the Teloschistaceae. The corticated
granules in the central part of the thallus resemble the blastidia known from several species of Fulgensia A. Massal.
& De Not. In this sense, it was used for most of the past
century. Sequencing of the type showed that this species
belongs to Wetmoreana. None of the species in this genus
are blastidiate. Phylogenetically, it clusters deep inside Wetmoreana (Fig. 71). Note that Caloplaca brouardii de Lesd.
is phylogenetically also a Wetmoreana (Wilk et al. in prep).
This species is described from the same locality as Buellia
pruinocalcarea (for details see under that species).
Class Leotiomycetes O.E. Erikss. & Winka
Phacidiales C.E. Bessey
Phacidiales was placed in Leotiomycetes by Bessey
(1907). Quijada et al. (2018) included three families in
Phacidiale (viz. Helicogoniaceae, Phacidiaceae, Tympanidaceae) and one informal taxonomic lineage with 29 genera. Wijayawardene et al. (2022) accepted two families viz.
Fungal Diversity (2022) 117:1–272
95
introduce a new species of Phacidium from decaying wood
in terrestrial habitats in China.
Fig. 72 Phacidium chinense (KUN-HKAS 112899, holotype). a
Herbarium. b Conidiomata on the host. c Vertical sections of conidioma. d Sections of peridium. e Conidiogenous cells and developing
conidia. f–k Conidia. l Germinating conidium. m, n Culture on PDA.
Scale bars: c = 100 µm, d = 50 µm, e, k = 10 µm, f–j = 5 µm, l = 20 µm
Helicogoniaceae and Phacidiaceae in this order and accepted
Tympanidaceae in Leotiales.
Phacidiaceae Fr. [as 'Phacidiacei'], Summa veg. Scand.,
Sectio Post. (Stockholm): 367 (1849).
Phacidiaceae was introduced by Fries (1849) and typified
by Phacidium (Crous et al. 2014). Six genera (Allantophomopsiella, Allantophomopsis, Bulgaria, Darkera, Phacidium
and Potebniamyces) were included in Phacidiaceae based
on DNA sequence data (Crous et al. 2014; Li et al. 2020).
Wijayawardene et al. (2022) accepted nine genera in this
family.
Phacidium Fr., Observ. mycol. (Havniae) 1: 167 (1815).
The generic name Phacidium was introduced by Fries
(1815) with Phacidium lacerum as the type species. Phacidium species are widely distributed throughout the globe
and has been reported as pathogens on dead leaf tips and as
saprobic on dead leaves of several host families (Crous et al.
2014; Li et al. 2020). Both asexual morph and sexual morph
of this genus are known (Li et al. 2020). There are 43 taxa
are listed in Species Fungorum (2022a, b) (http://www.speci
esfungorum.org/Names/Names.asp). However, it is a poorly
studied genus due to the lack of molecular data. Here we
Phacidium chinense G.C. Ren & K.D. Hyde sp. nov.
Index Fungorum number: IF559693; Facesoffungi number: FoF10836, Fig. 72
Etymology: The species epithet reflects the country where
the species was collected.
Holotype: KUN-HKAS 112899
Saprobic on dead wood of Rosa sp. Sexual morph:
Not observed. Asexual morph: Conidiomata 170–200 μm
high, 150–240 μm diam. ( x̄ = 180 × 200 μm, n = 5), black,
pseudostromatic, solitary or gregarious, semi-immersed to
superficial, multi-locular, with 3–10 locules embedded in
the pseudostroma. Ostioles 60–85 × 45–70 ( x̄ = 73.5 × 58.5,
n = 5) μm, centrally located, circular. Conidiomata wall
15–35 µm thick, 3–5 layered, comprising brown cells of
textura angularis, thick-walled at basal, thin-walled at side.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3.5–5.5 × 1.2–2.0 µm ( x̄ = 4.8 × 1.5, n = 10), hyaline, enteroblastic, phialidic, discrete, cylindrical, smoothwalled, arising from stratum. Conidia 4.5–6 × 2.0–2.4 µm
( x̄ = 5.2 × 2.1 µm, n = 30), hyaline, oblong, unicellular, thickand smooth-walled.
Culture Characters: Colonies on PDA, reaching
80–90 mm diam., after four weeks at 20–25 °C, medium
dense, circular, rough, fluffy, cotton, gray, with white papillate on the surface, reverse dark-gray.
Material examined: China, Yunnan Province, Diqing
Autonomous Prefecture, Xianggelila (27.28′ 8° N, 99.50′
45° E), 2958 m, on dead wood of Rosa sp. (Rosaceae), 30
August 2020, Guang-Cong Ren, W06 (KUN-HKAS 112899,
holotype), ex-type living culture KUMCC 20-0168.
GenBank numbers: ON490924 (LSU), ON490925(ITS),
ON506923(tef1), ON506922 (rpb2)
Notes: Phacidium chinense is introduced as a new species based on its distinct morphology, which is supported
by phylogenetic analyses. In the phylogenetic analyses, P.
chinense is distinct from extant species in Phacidium and
formed a sister clade to Phacidium calderae, however, there
is no bootstrap support (Fig. 73). Phacidium chinense is different to P. calderae in having oblong conidia and phialidic,
cylindrical conidiogenous cells, while P. calderae in having
subcylindrical conidia with apical mucoid appendage and
proliferating with periclinal thickening conidiogenous cells
(Crous et al. 2014).
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan., Maharachch. &
K.D. Hyde
Diaporthales Nannf
Diaporthales containing numerous important endophytic,
saprobic and phytopathogenic ascomycetous families. Even
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96
Fig. 73 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS, tef1, and rpb2 sequence data. Fortyeight strains are included in the combined analyses which comprised
3378 characters (8200 characters for LSU, 546 characters for ITS,
879 characters for tef1, 1133 characters for rpb2) after alignment.
The tree topology of the maximum likelihood analysis is similar to
13
Fungal Diversity (2022) 117:1–272
the Bayesian analysis. The evolutionary model SYM + G applied to
ITS sequence data, while SYM + I + G applied to LSU, GTR + G tef1
and rpb2 gene regions. Bootstrap support values for ML ≥ 80% and
BYPP ≥ 0.95 are given near nodes respectively. The tree is rooted in
Phlyctema vincetoxici (CBS 123,726). Ex-type strains are in bold.
The newly generated sequences are indicated in bold red
Fungal Diversity (2022) 117:1–272
97
Fig. 74 Diaporthe foeniculina
(JZB320201, new host record)
a. Appearance of conidiomata
on the host. b. Section through
the conidiomata. c. Mature
conidia attached to the conidiophore and the conidioma cell
wall. d. Alpha and beta conidia.
e. Mature conidia. Scale bars: b,
c = 20 µm, d, e = 5 µm
though, families, and genera in this order showed high distinct morphological diversity, taxonomic placements are still
problematic (Senanayake et al. 2017).
Diaporthales introduced by Nannfeldt (1932), to accommodate Höhnel’s Eu-Diaportheen and Valseen taxa (Senanayake et al. 2018) and currently 30 families are in this
order named: Apiosporopsidaceae, Apoharknessiaceae,
Asterosporiaceae, Auratiopycnidiellaceae, Coryneaceae,
Cryphonectriaceae, Cytosporaceae, Diaporthaceae, Diaporthosporellaceae, Diaporthostomataceae, Dwiroopaceae,
Erythrogloeaceae, Gnomoniaceae, Harknessiaceae, Juglanconidaceae, Lamproconiaceae, Macrohilaceae, Melanconidaceae, Melanconiellaceae, Neomelanconiellaceae,
Phaeoappendicosporaceae, Phaeochorellaceae, Prosopidicolaceae, Pseudomelanconidaceae, Pseudoplagiostomataceae, Schizoparmaceae, Stilbosporaceae, Sydowiellaceae,
Synnemasporellaceae and Tubakiaceae (Hyde et al. 2020a,
b, c; Wijayawardena et al. 2022).
Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638 (1926)
Diaporthaceae was introduced and placed in Diaporthales
by von Höhnel (1917). The members of Diaporthaceae are
known to be endophytic, pathogenic and saprobic. Species in
Diaporthaceae mostly inhabit in terrestrial hosts and rarely
on aquatic hosts (Udayanga et al. 2011; Dissanayake et al.
2017; Senanayake et al. 2017). There are 15 genera accepted
in Diaporthaceae, viz. Apioporthella, Apiosphaeria, Chaetoconis, Chiangraiomyces, Diaporthe, Hyaliappendispora,
Leucodiaporthe, Massariothea, Mazzantia, Ophiodiaporthe,
Paradiaporthe, Phaeocytostroma, Phaeodiaporthe, Pustulomyces, and Stenocarpella (Hyde et al. 2020a, b, c).
Diaporthe Nitschke, Fungi rhenani exsic., suppl., fasc. 5:
no. 1988 (1867)
Diaporthe is the type genus of Diaporthaceae, and it was
established by Nitschke (1867). Diaporthe species have been
recorded as endophytes or saprobes on a wide range of host
plants in different geographical areas (Udayanga et al. 2011;
Dissanayake et al. 2017; Abeywickrama et al. 2020). Many
economically significant crops are infected by pathogenic
Diaporthe species leading to severe crop losses (Manawasinghe et al. 2019; Abeywickrama et al. 2020), with blights,
fruit and root rots, cankers, diebacks, wilts and leaf spots
(Manawasinghe et al. 2019; Abeywickrama et al. 2020). The
genus contains 1152 epithets in Index Fungorum (assessed
in 29.08.2022; Index Fungorum 2022a, b).
Diaporthe foeniculina (Sacc.) Udayanga & Castl., in Udayanga et al., Persoonia 32: 95 (2014).
Index Fungorum number: IF803929; Facesoffungi number: FoF02183; Fig. 74
Saprobic on dead aerial branch of Ficus carica. Sexual
morph: See Udayanga et al. (2014). Asexual morph:
coelomycetous. Conidiomata observed as small black
dots on the host, semi-immersed to immersed, pycnidial,
pyriform, scattered, ostiolate, 150–300 µm diam. Conidiomata wall consisting of 3–4 layers of pale brown, thickwalled cells of textura angularis. Conidiophores hyaline,
smooth, unbranched. Alpha conidia hyaline, smooth-walled,
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98
Fig. 75 Phylogram generated
from maximum likelihood
analysis based on ITS, cal,
his, tef1 and tub2 sequenced
data of given Diaporthe species. Related sequences were
obtained from GenBank, and
223 strains are included in the
sequence analyses, with 2674
columns, 1972 distinct patterns
1439 parsimony-informative,
340 singleton sites, 894 constant
sites. Diaporthella corylina
(CBS121124) is used as the
outgroup taxon. Bootstrap
support values for ML ≥ 65%,
BYPP ≥ 0.90 are given near the
nodes. Type strains are in bold.
Newly generated strains are in
red bold
13
Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
99
Fig. 75 (continued)
bi- to multi-guttulate, ovate to ellipsoidal, base sub-truncate,
5–7.5 × 1.5–3 µm (n = 20). Beta conidia aseptate, hyaline,
smooth, apex and base bluntly rounded, slightly curved,
15–25 × 0.5–2 µm (n = 10).
Culture characteristics: Colonies on PDA entirely white
both on surface and reverse. Aerial mycelium cottony, colonies reaching 60 mm diam. after 7 days in room temperature.
Material examined: Italy, Province of Forlì-Cesena [FC],
near Pianetto—Galeata, on dead and aerial branch of Ficus
carica L. (Moraceae), 21 December 2018, E. Camporesi, IT
4192 (JZBH 320201), living culture JZB 320201.
Hosts: Wide host range, including Achillea, Ailanthus, Amorpha, Angelica, Arctium, Asparagus, Camellia,
Castanea, Chenopodium, Citrus, Cupressus, Diospyros,
Eucalyptus, Ficus carica Hemerocallis, Lunaria, Melilotus,
Microcitrus, Persea, Platanus, Prunus, Rosa, Rubus, Vicia
and Wisteria (Farr and Rossman 2022; this study).
Distribution: Wide geographical range, including in
Chile, Greece, Iran, Italy, Malta, New Zealand, Portugal,
Serbia, South Africa, Spain, Thailand, Turkey, Uruguay, US
(Farr and Rossman 2022; this study).
GenBank numbers: OP002068 (ITS), OP837431(his),
OP837429 (tub2)
Notes: Diaporthe foeniculina (JZB 320201) was recovered from a dead aerial branch of Ficus carica in Italy. Our
strain shared similar morphology with the type strain of D.
13
100
Fungal Diversity (2022) 117:1–272
difference in HIS gene (94.23% similarity). We were unable
to obtain cal and tef1 sequence data for our strain, and we
could not compare the base pair difference for them. Thus,
based on the multi-gene phylogeny and morphology; this
study presents the first report of D. foeniculina from a Ficus
carica from Italy.
Fig. 76 Micromorphological features of D. longicolla (CPDl21,
new host record) a. Cowpea stem affected by Diaporthe longicolla.
b. Stereo view of infected region showing pycnidial structures. c–d.
Stereo view showing cirri of spores erupted from pycnidia. e. SEM
image of pycnidium. f. Conidia of D. longicolla observed in SEM.
Scale bars: b = 10 mm; c– d = 2 mm; e – f = 10 µm
foeniculina (CBS 111553) which was introduced by Udayanga et al. (2014), with minor dimensional differences.
Conidiomata of our strain (JZB 320201) are comparatively smaller than those of D. foeniculina (CBS 111553)
(150–300 μm diam. vs 400– 700 µm diam.). Further we
have observed smaller alpha conidia in our strain than
CBS 111553 (5–7.5 × 1.5–3 µm vs 8.8 ± 0.3 × 2.4 ± 0.1 μm)
(Udayanga et al. 2014). These morphological differences
probably due to environmental factors and host variations.
Phylogenetic analyses using combined ITS, cal, his, tef1,
tub2 sequence data confirmed that our strain is D. foeniculina and it is clade with the strain MFLUCC 20-0151
with high statistical support (92/1.00) (Fig. 75). Comparisons of base pair differences for ITS, tub2 and his genes
between our strain (JZB 320201) and the ex-epitype strain
of D. foeniculina (CBS 111553) reveal less than 1% base
pair differences in ITS and tub2 gene regions (ITS = 0.57%,
tub2 = 0.97%). However, we observe 5.77% base pair
13
Diaporthe longicolla (Hobbs) J.M. Santos, Vrandečić &
A.J.L. Phillips, in Santos, Vrandečić, Čosić, Duvnjak &
Phillips, Persoonia 27: 13 (2011)
Index Fungorum number: IF164797; Faceoffungi number: FOF11682; Figs. 76, 77
Pathogenic and associated with stem of Vigna unguiculata. Sexual morph: Not observed. Asexual morph:
Conidiomata 80–130 µm high, 230–320 µm diam.
x̄ = 118 × 290 μm, n = 20, pycnidial, pyriform, initially
immersed, erumpent at maturity, globose to pyriform, black,
elongated neck, often with light yellowish white conidial
cirrus extruding from ostiole. Pycnidial wall parenchymatous consisting of 4–7 layers of pale brown, thick-walled
cells of textura angularis, Pycnidia globose locules and
prominent beaks, which immersed in medium, black, solitary, discoid or irregular. Conidiophores 4–7 × 4.1–7.3 μm
( x̄ = 4.4 × 6.3 μm, n = 30), ampulliform, straight to sinuous, unbranched, hyaline, smooth. Conidiogenous cells
7.8–13.8 × 1.4–2.7 μm ( x̄ = 10.9 × 2.1 μm, n = 30), phialidic, terminal, cylindrical, slightly tapering towards the
apex. Alpha-conidia 5.1–7.5 × 1.2–3.4 μm ( x̄ = 6.1 × 2.6 μm,
n = 20), aseptate, hyaline, smooth, ovate to ellipsoidal, guttulate. Beta-conidia 5.8–7.5 × 2.5–3.5 μm ( x̄ = 6.4 × 2.8 μm,
n = 10), hyaline, filiform, hamate.
Cultural characteristics: On potato dextrose agar, the
fungus initially produced white fluffy aerial hyphae, forming relatively dense concentric pattern colony, which subsequently exhibited light yellow pigmentation.
Material examined: India, Karnataka, Mysuru Doddamaragowdanahally, on infected stem of cowpea plants
as pathogen. July, 2020, S. Mahadevakumar, Y.S. Deepika (UOM-IOE 20/25), living cultures CPDl21, CPDl22,
MKSVu012.
Hosts: Wide host range, including Abutilon, Acer, Actinidia, Ambrosia, Arachis, Chamaesyce, Cucumis, Euphorbia, Glycine, Helianthus, Ipomoea, Kalanchoe, Phaseolus,
Pisum, Pyrus, Rumex, Solanum, Trichilia, Vigna and Xanthium (Farr and Rossman 2022; this study).
Distribution: Wide geographical range, including in
Argentina, Australia, Brazil, China, Croatia, Greece,
India, Italy, Malaysia, Missouri, South Korea, US (Farr
and Rossman 2022; this study).
GenBank Numbers: CPDl21—MW737797 (ITS),
OM934823 (tub2), OM934820 (tef1)
CPDl22–MW737798 (ITS), OM934824 (tub2),
OM934821(tef1)
Fungal Diversity (2022) 117:1–272
101
Fig. 77 Micromorphological
features of D. longicolla: a–d.
Conidial mass with spores
observed under compound
microscope. e–g Beta conidia of
Diaporthe longicolla recorded
on cowpea plants. Scale bar:
a–g = 20 µm
MKSVu012– KT819767 (ITS), OM934825(tub2),
OM934822 (tef1)
Notes: The symptoms were observed on stems of cowpea. Initial symptoms appeared as small lesions, more
or less circular, later elongated, blackish-brown lesions,
eventually pycnidia developed (Fig. 76). Stem girdling
occurs and the shoot above the infected area wilts and
dries up. Pathogenicity tests were conducted and proved to
be pathogenic on healthy cowpea plants. Morphologically
our strain shares similar morphology with the ex-type
strain of D. longicolla (Fig. 77). In the multigene phylogenetic analyses, our strain clusters with D. longicolla with
a high bootstrap support (Fig. 75). Previously, Phomopsis
longicolla is known to be associated with cowpea seeds.
However, no reports are available on the association of D.
longicolla of cowpea in India. This is the first report of D.
longicolla associated with cowpea from India.
Diaporthe phaseolorum (Cooke & Ellis) Sacc., Syll. Fung.
(Abellini) 1: 692 (1882).
Index Fungorum number: IF164797; Faceoffungi number: FoF10638; Figs: 78, 79.
Pathogenic and associated with stem of Vigna unguiculata. Sexual morph: Not observed. Asexual morph: Conidiomata 105–192 µm high, 165–285 µm diam. x̄ = 122 × 255
μm, n = 30, pycnidial, pyriform, initially immersed,
erumpent at maturity, globose to pyriform, black, elongated
neck, often with light yellowish white conidial cirrus extruding from ostiole. Pycnidial wall parenchymatous consisting
of 3–6 layers of pale brown, thick-walled cells of textura
angularis. Pycnidia globose locules and prominent beaks,
which immersed in medium, black, solitary, discoid or
irregular. Conidiophores 3.8–7.5 × 3.8–7.5 μm ( x̄ = 4.2 × 5.8
μm, n = 30), ampulliform, straight to sinuous, unbranched,
hyaline, smooth. Conidiogenous cells 8.2–12.8 × 1.6–2.5
μm ( x̄ = 9.8 × 2.2 μm, n = 30), phialidic, terminal, cylindrical, slightly tapering towards the apex. Alpha-conidia
5.3–7.7 × 1.5–4.6 μm ( x̄ = 6.5 × 2.8 μm, n = 30), aseptate,
hyaline, smooth, ovate to ellipsoidal, guttulate. Beta-conidia
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Fungal Diversity (2022) 117:1–272
Fig. 78 Diagnostic features
of cowpea stem blight and
pod blight disease caused by
Diaporthe phaseolorum and
D. longicolla: a–b. field view
of cowpea plants affected with
Diaporthe stem blight and pod
blight disease. c, d, g. Stem
blight disease caused by D.
longicolla. e,f,h,i. pod blight
disease symptoms caused by D.
phaseolorum
10.2–17.5 × 1.2–2.3 μm ( x̄ = 12.6 × 1.2 μm, n = 30), hyaline,
filiform, hamate.
Culture characteristics: On PDA, colonies with white,
floccose, aerial mycelium were recorded after 7 days of incubation. Pure cultures obtained from the colonies expressed
from infected pod and stem samples.
Material examined: India, Karnataka, Mysuru Doddamaragowdanahally, on infected stem of cowpea plants as
pathogen, July, 2020, S. Mahadevakumar, Y.S. Deepika, N.
13
Lakshmidevi (UOM-IOE 20/26), living cultures CPDp1,
CPDp2.
Hosts: Wide host range, including Acer, Actinidia,
Aeschynomene, Arctium, Aspalathus, Aster, Calopogonium, Cannabis, Caperonia, Capsicum, Centrosema, Clitoria, Cyphomandra, Desmanthus, Desmodium, Eriobotrya,
Euphorbia, Glycine, Helianthus, Hylocereus, Ipomoea, Jatropha, Lablab, Lupinus, Lycopersicon, Macroptilium, Macrotyloma, Maytenus, Ocimum, Olearia, Panicum, Phalaris,
Fungal Diversity (2022) 117:1–272
103
Fig. 79 Cultural and micromorphological characteristics of Diaporthe phaseolorum isolated from Cowpea: a–c. Pure cultures of D. phaseolorum on PDA (15 days old). d–f. Alpha conidia of D. phaseolorum. Scale bar: d–f = 10 µm
Phaseolus, Pyrus, Stokesia, Vigna, Vitis and Zea (Farr and
Rossman 2022; this study).
Distribution: Wide geographical range, including in Australia, Barbados, Brazil, Brunei Darussalam, Cameroon,
Canada, China, Colombia, Cook Islands, Croatia, Cuba,
Dominican Republic, Fiji, India, Italy, Jamaica, Korea,
Maryland, Mauritius, Missouri, New Zealand, Papua New
Guinea, Spain, South Africa, Switzerland, Thailand, Tonga,
Trinidad and Tobago, United Kingdom, United States, Uruguay, Venezuela and West Indies (Farr and Rossman 2022;
this study).
GenBank Numbers: CPDp1– MW737799 (ITS),
OM934818 (tub2), OM934816 (tef1)
CPDp2– MW737800 (ITS), OM934819 (tub2),
OM934817 (tef1)
Notes: Morphologically our strain is similar to the extype strain of D. phaseolorum (Fig. 78). In the multigene
phylogenetic analyses, our strain clustered with the ex-type
strain of D. phaseolorum (Fig. 75). Pathogenicity tests were
conducted and proved to be pathogenic on healthy cowpea
plants. Diaporthe longicolla causes stem blight of cowpea
while D. phaseolorum causes pod blight of the same host
(Fig. 79). This is the first report of D. phaseolorum associated with cowpea from India and worldwide.
Melanconiellaceae Senan., Maharachch. & K.D. Hyde, in
Senanayake et al., Stud. Mycol. 86: 275 (2017).
Melanconiellaceae was invalidly introduced (Locquin
1984) for Melanconiella and it was validated by Senanayake
et al. (2017). The type species of Melanconiella is M. spodiaea (Tul. & C. Tul.) Sacc. (Fan et al. 2018). Fan et al. (2018)
accepted Melanconiella, Microascospora and Sheathospora
and Senanayake et al. (2018) accepted Greeneria, Melanconiella and Microascospora in the family. Hyde et al (2020c)
accepted five genera; Greeneria, Melanconiella, Microascospora, Septomelanconiella and Sheathospora in the
family.
Melanconiella Sacc., Syll. fung. (Abellini) 1: 740 (1882)
Melanconiella was established by Saccardo (1882) for
the type species M. spodiaea Tul. & C. Tul. and second
species M. chrysostroma (Fr.) Tul. & C. Tul. Melanconiella species are found in overwintered plants as saprobes
or as mild canker causing agents (Voglmayr et al. 2012;
Senanayake et al. 2018). There are 40 species epithets
recorded in Index Fungorum (January 2022).
Melanconiella meridionalisVoglmayr & Jaklitsch, in Voglmayr et al., Fungal Diversity 57(1):33 (2012).
Index Fungorum number: IF800123; Facesoffungi number: FoF10701, Fig. 80
Saprobic on dead twigs of Fagus sylvatica. Sexual
morph: Pseudostromata indistinct, less commonly distinct and circularoutline, causing minute bumps in the
bark. Ectostromatic disc flat, 2–2.3 mm long, well-defined,
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104
Fungal Diversity (2022) 117:1–272
Fig. 80 Melanconiella meridionalis (MFLU 15–2604, new
host and geographical record)
a, b Appearance of pseudostromata on dead branch of Fagus
sylvatica c Pseudostroma in
transverse section d, e Pseudostroma in vertical sections
f Peridium. g Paraphyses.h–j
Asci. k–n Ascospores. Scale
bars: d = 50 μm, e = 80 μm,
f = 10 μm, h–j = 25 μm,
k–n = 10 μm
with circular or elliptic outline, cream, light or pale yellow. Central column dark brown. Entostroma yellowish
hyphae. Ostiole central. Perithecia 200–600 μm, subglobose, immersed, coriaceous, brown to black, Perithecia
wall 8–10 µm wide, comprising brown cells of textura
angularis of inner layer and 15–18 µm wide, unequally
thick, comprising irregular dark brown cells of textura
prismatica of outer layer. Hamathecium comprising
1–1.5 µm septate, unbranched, cellular pseudoparaphyses. Asci (85–)95–100(–115) × (8–)10–12(–15) μm
( x̄ = 97 × 11 µm, n = 20), broadly cylindrical to fusoid,
13
8-spored, distinct apical ring with short pedicel or sessile.
Ascospores uni- or irregularly biseriate, hyaline, fusoid,
constricted at the septum, (20–)23–25(–28) × (4–) 5–6(–7)
μm, ( x̄ = 24 × 5.2 µm, n = 30); ends rounded, upper cell
mostly larger and some guttules. Asexual morph: Not
observed.
Material examined: Italy, Province of Forlì-Cesena, Santa
Sofia, on twigs of Fagus sylvatica L. (Fagaceae), 3 August
2015, E. Camporesi, IT2575 (MFLU 15-2604).
Host and distribution: Ostrya carpinifolia in China
(Voglmayr et al. 2012; Fan et al. 2018), Europe and North
Fungal Diversity (2022) 117:1–272
105
Fig. 81 Phylogram generated from the maximum likelihood analysis
based on combined LSU, ITS, rpb2 and tef1 sequence data representing family Melanconiellaceae. Related sequences are taken from Fan
et al. (2018) and Senanayake et al. (2018). Melanconis stilbostoma
(CFCC 50,480) and M. stilbostoma (MS) are used as the outgroup
taxa. Twenty-eight strains are included in the combined gene analyses
comprising 3380 characters after alignment (880 characters for LSU,
500 characters for ML ≥ 75% and BYPP ≥ 0.95 are given above the
nodes. Ex-type strains are in bold and new strain is indicated in blue
America (Voglmayr et al. 2012), Fagus sylvatica in Italy
(this study).
GenBank numbers: OM403250 (ITS), OM403249 (LSU)
Notes: The new strain shares a close phylogenetic affinity
to Melanconiella meridionalis in our combined LSU, ITS,
rpb2 and tef1 sequence data analyses (Fig. 81). This species was previously recorded from dead corticated twigs and
branches of Ostrya carpinifolia (Betulaceae) from different
localities i.e. Australia, Croatia, Greece, Italy and Slovenia
(Voglmayr et al. 2012). Melanconiella meridionalis has not
been reported from Fagaceae and here we provide the first
association of sexual morph of species with Fagus sylvatica.
crown age of 50.08 Mya and stem age of 101.46 Mya (Hyde
et al. 2020a, b, c). The data of divergence time estimates
for Pararamichloridiales is line with recommendations for
ranking families.
Pararamichloridiales Crous
Notes: Pararamichloridiales was introduced by Crous
et al. (2017) to accommodate a monotypic family including two genera, namely Pararamichloridium and Woswasia. However, Woswasia (Woswasiaceae) was treated in
Diaporthomycetidae families incertae sedis by Zhang et al.
(2017) based on morphological and phylogenetical analyses. This result was supported by Hyde et al. (2020a, b,
c). Divergence time estimates for Pararamichloridiales are
Pararamichloridiaceae Crous, in Crous et al., Persoonia
39: 357 (2017)
Notes: Pararamichloridiaceae was introduced as a monotypic family by Crous et al. (2017) for Pararamichloridium
Crous. However, Zhang et al. (2017) established a new
family Woswasiaceae to accommodate Cyanoannulus and
Xylomelasma while Woswasia was placed in Diaporthomycetidae families incertae sedis based on its close phylogenetic affinity, and this was conferred in later studies (Hyde
et al. 2020a, b, c; Wijayawardene et al. 2020). Members of
Pararamichloridiaceae are pathogenic on plant leaves (Crous
et al. 2017, 2018).
Pararamichloridium Crous, in Crous et al., Persoonia 39:
357 (2017)
Notes: Pararamichloridium, the type genus of Pararamichloridiaceae, was established by Crous et al. (2017) to
13
106
Fungal Diversity (2022) 117:1–272
Fig. 82 Pararamichloridium
aquisubtropicum (GZAAS
21–0382, holotype) a–b Colony
on decaying wood c–e Conidiophores with attached conidia
d–g Conidiogenous cells and
conidia h Conidia i Germinating
condium j–k Colony on PDA
from above and below. Scale
bars: c–e = 20 μm, f–i = 10 μm
accommodate two species, P. livistonae Crous (as the type
species) and P. verrucosum (V. Rao & de Hoog) Crous.
Crous et al. (2018) introduced the third species of Pararamichloridium based on blast search and phylogenetic analysis.
In this study, morphological characteristics and multi-gene
phylogenetic analysis of a combined LSU and ITS sequence
13
data reveals the fourth new species of Pararamichloridium from dead wood collected in freshwater from China
(Figs. 82, 83).
Pararamichloridium aquisubtropicum J.Y. Zhang, Y.Z. Lu
& K.D. Hyde, sp. nov.
Fungal Diversity (2022) 117:1–272
107
Fig. 83 Phylogram generated
from maximum likelihood
analysis based on combined
LSU and ITS sequence data.
Twenty–two taxa were included
in the combined analyses, which
comprised 1524 characters
(LSU: 885, ITS: 639) after
alignment. Bootstrap support values for ML ≥ 50% and
BYPP ≥ 0.95 are given above
the nodes. The tree is rooted
with Diaporthe passiforae (CBS
132,527) and D. obtusifoliae
(CPC 32,226). The ex–type
strains are indicated in bold.
The newly generated sequence
is indicated in blue
Index Fungorum number: IF559507; Facesoffungi number: FoF10677; Fig. 82
Etymology: Referring to the aquatic habitat and collecting
site in subtropical country, China
Holotype: GZAAS 21–0382
Saprobic on submerged decaying wood. Asexual
morph: Colonies on natural substrate superficial, brown,
smooth, solitary. Mycelium partly immersed, consisting of branched, smooth, brown hyphae. Conidiophores
119–202 × 2.6–4.7 μm ( x̄ = 161.4 × 3.6 μm, n = 24),
macronematous, mononematous, sucylindrical, straight,
unbranched, brown, smooth, 6–9-septate. Conidiogenous
cells 27.3–37 × 2.6–3.8 μm ( x̄ = 27.6 × 3.3 μm, n = 15),
holoblastic, polyblastic, integrated, terminal, subhyaline
to pale brown, subcylindrical. Conidia 4–8 × 3–4.7 μm
( x̄ = 5.2 × 3.7 μm, n = 25), acrogenous, solitary, aseptate,
pale brown, globose to subglobose. Sexual morph: Not
observed.
Culture Characters: Colonies on PDA reaching 33 mm
in 44 days at 25 °C, flat, curler, near–round or round, gray,
smooth, middle erumpent; In reverse, yellow at the center,
lightly brown or gray at the margin.
Material examined: China, Guizhou Province, Xishui
County, on decaying wood submerged in a freshwater stream, 16 September 2020, Jian Ma, XY2(GZAAS
21–0382, holotype); ex–type living culture, GZCC 21–0668.
GenBank numbers: OM339437 (ITS), OM339434 (LSU).
Notes: The phylogenetic analysis (Fig. 83) revealed that
Pararamichloridium aquisubtropicum forms distinct lineage belonging to Pararamichloridium, where it is sister
to P. caricicola with 100% BS, 1.00 BYPP, high support.
Pararamichloridium caricicola is the closest species based
on BLASTn result of LSU and ITS region with 99.55%
and 96.11% similarity, respectively. Pararamichloridium
caricicola is found in culture, while our new collection was
saprobic on submerged decaying wood. Pararamichloridium aquisubtropicum shares the same morphology with P.
caricicola in having brown, septate, smooth conidiophores
integrated, terminal, subcylindrical conidiogenous cells
and ellipsoid, pale brown aseptate, conidia. However, P.
13
108
aquisubtropicum differs from P. caricicola by it darker and
longer (119–202 × 2.5–4.7 μm vs 35–100 × 2.5–3 μm) conidiophores. Thus, P. aquisubtropicum is introduced here as
a distinct novel species based on its distinct morphological
features and phylogenetic placement.
Distoseptisporales Z.L. Luo, K.D. Hyde & H.Y. Su Fungal
Diversity 99: 482 (2019).
Distoseptisporaceae was established by Su et al. (2016)
with a single genus, Distoseptispora. Based on morphology
and phylogenetic analysis of combined LSU, SSU, rpb2, and
tef1 sequence data, Luo et al. (2019) established a new order,
Distoseptisporales. Previously, the order Distoseptisporales
was classified under the class Sordariomycetes, subclass
Diaporthomycetidae. Aquapteridospora was described by
Yang et al. (2015) and assigned to the Diaporthomycetidae
genus incertae sedis. Later, Dong et al. (2021) performed
a molecular phylogeny study using combined LSU, ITS,
tef1, and rpb2 sequence data and established a new family, Aquapteridosporaceae K.D. Hyde & Hongsanan, to
accommodate a single genus, Aquapteridospora, and placed
Aquapteridosporaceae in the order Distoseptisporales. Distoseptisporales currently consists of two families (Aquapteridosporaceae and Distoseptisporaceae), with Distoseptisporaceae serving as the type family.
Distoseptisporaceae K.D. Hyde & McKenzie, Fungal
Diversity 80: 402 (2016)
Su et al. (2016) identified two Sporidesmium-like taxa
with distinct morphology and phylogenetic relationships.
They can be distinguished from Sporidesmiaceae based on
strong molecular evidence and morphological investigation.
As a result, the family Distoseptisporaceae was established
to include Sporidesmium-like species under the type genus
Distoseptispora. In the previous study, there was no sexual
morph known for this family (Su et al. 2016; Yang et al.
2018; Hyde et al. 2019, 2020a, b, c, 2021; Luo et al. 2019;
Sun et al. 2020a, b). Recently, Yang et al. (2021) proposed
a sexual morph of Distoseptispora and this is the first time
that the sexual morph of Distoseptispora has been found.
Distoseptispora K.D. Hyde, McKenzie, Maharachch. Fungal
Diversity 80:375–409 (2016)
Su et al. (2016) defined Distoseptispora as a genus with
Distoseptispora fluminicola as the type species. Yang et al.
(2018) updated the Distoseptispora genus description. In
addition, the sexual morph of this species is unidentified in
the previous study (Su et al. 2016; Yang et al. 2018; Hyde
et al. 2019, 2020a, b, c; Luo et al. 2019; Sun et al. 2020a,
b). There is only one species of sexual morph described
in Distoseptispora (Yang et al. 2021). The sexual morph
is characterized by being solitary or gregarious, immersed
to semi-immersed, perithecial, subglobose to ellipsoidal,
13
Fungal Diversity (2022) 117:1–272
ostiolate, dark brown ascomata with a short neck and hyaline, 0–3-septate, ascospores with a mucilaginous sheath
(Yang et al. 2021). The asexual morph is distinguished
by hyphomycetous, macronematous conidiophores, percurrent, elongate conidiogenous cells, olivaceous, brown,
yellowish, or reddish brown, euseptate or distoseptate
conidia, and rarely muriform conidia. (Su et al. 2016; Xia
et al. 2017; Luo et al. 2018a, b; Tibpromma et al. 2018;
Yang et al. 2018; Hyde et al. 2020a, b, c). The genus now
has 46 recognized species, 14 of which are from terrestrial
habitats and 32 from freshwater habitats (Su et al. 2016;
Hyde et al. 2016, 2019, 2020, 2021; Xia et al. 2017; Yang
et al. 2015, 2018; Luo et al. 2018a, b, 2019; Monkai et al.
2020; Song et al. 2020; Sun et al. 2020a, b; Li et al. 2021a,
b; Yang et al. 2021, Hyde et al. 2021, Index Fungorum
2022a, b). Distoseptispora hyalina J. Yang and K.D. Hyde
is the first sexual morph reported in the genus based on
molecular DNA data (Yang et al. 2021).
Distoseptispora bambusicola X. Tang, Jayaward, J.C Kang
& K.D. Hyde sp. nov.
Index Fungorum number: IF558533; Facesoffungi number: FoF09940; Fig. 84
Etymology: Named after the host bamboo from which the
holotype was found.
Holotypus: GZAAS21-0379.
Saprobic on decaying stems of bamboo submerged in a
freshwater stream habitat. Sexual morph: Not observed.
Asexual morph: Colonies effuse, scattered, hairy, brown
to dark brown. Mycelium mostly immersed, composed of
branched, septate, brown, smooth hyphae. Conidiophores
macronematous, mononematous, pale brown to brown, solitary, 4–7-septate, erect, straight or flexuous, unbranched,
slightly constricted at septa, smooth, cylindrical,
64–116 μm × 4–7 μm ( x̄ = 90.5 × 5.5 μm, n = 30), truncate at
the apex. Conidiogenous cells holoblastic, monoblastic, integrated, terminal, determinate, brown, cylindrical. Conidia
72–193 μm × 7.5–14.5 μm ( x̄ = 126 × 14 μm, n = 30) acrogenous, solitary, obclavate or lanceolate, rostrate, straight or
slightly curved, multi-distoseptate, up to 16-distoseptate,
guttulate, pale brown, tapering towards the rounded apex,
truncate at the base, slightly constricted at septa, smoothwalled, rounded at apex, with a truncate base and faintly to
heavily pigmented scar.
Culture characters: Colony grown on PDA in room
temperature, circular, fluffy, white, dense in the center, but
sparse, slightly olivaceous in the outside. In reverse, deeply
olivaceous in the center, slightly yellow at the entire margin.
Refer to the pigment produced on PDA and OA, purpure.
Material examined: China, Guizhou Province, Zunyi
City, on decaying stems of bamboo submerged in a freshwater stream, 21 February 2021, Xia Tang, K1 (GZAAS210379, holotype), ex-type living culture, GZCC21-0667.
Fungal Diversity (2022) 117:1–272
109
Fig. 84 Distoseptispora bambusicola (GZAAS21-0379,
holotype). a, b Colonies on
dead wood c–e Conidiophores
f Cnidiogenous cell g–n
Conidia o Germinated conidia
p, q Colony on PDA (up-front,
down-reverse). Scale bars:
c–j = 20 μm, k–o = 50 μm
GenBank numbers: MZ474873 (ITS), MZ474872 (LSU),
MZ474866 (SSU), OM272845 (tef1).
Notes: Distoseptispora bambusicola clustered with D.
hydei, D. obpyriformis, and D. rostrata with a 100%ML,
99%MP and 1.00 BYPP support. According to the morphological comparisons (Fig. 85), our novel species can be
distinguished from closely related species. Distoseptispora
bambusicola differs from D. rostrata by its smaller conidiophores (64–116 μm × 4–7 μm vs 82–126 μm × 5–7 μm),
larger conidia (72–193 μm × 7.5–14.5 μm vs
115–155 μm × 9–11 μm) and less distosepta (16-distosepta vs 23-distosepta). Distoseptispora bambusicola
13
110
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Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
◂ Fig. 85 Phylogram generated from parsimony analysis based on
combined ITS, LSU, SSU, rpb2 and tef1 sequence data of Distoseptispora. The ML and MP bootstrap support values ≥ 70% are and
branches with BYPP ≥ 0.95 are given above the nodes. The ex-types
(reference strains) are in bold; the new isolates are in blue bold. The
tree is rooted with Aquapteridospora fusiformis (MFLU 18–1601),
Aquapteridospora lignicola (MFLUCC 15–0377) and Aquapteridospora aquatica MFLUCC 17–2371)
shares similar morphological characteristics with D.
obpyriformis in the size, shape, color of conidiophores,
but differ in conidial size (72–193 μm × 7.5–14.5 μm vs
53–71 μm × 12–16 μm). Distoseptispora bambusicola differs from D. obpyriformis by having obclavate or lanceolate,
111
longer conidia. Distoseptispora bambusicola can be distinct from Distoseptispora hydei by conidal shape and
size (72–193 μm × 7.5–14.5 μm vs 32–58 μm × 10–15 μm;
obclavate or lanceolate vs obpyriform to fusiform). Distoseptispora hydei is characterized with a gelatinous sheath
around the tip of conidia, while D. bambusicola lacks this
character. According to the comparisons of our novel species with other phylogenetically related taxa based on a
pairwise nucleotide comparison of ITS (Jeewon and Hyde
2016), D. bambusicola differs from D. rostrata in 13/ 517 bp
(2.5%), D. hydei in 8/ 395 bp (2.0%) and differs from D.
obpyriformis in 25/571 bp (4.0%). Pairwise nucleotide comparison of LSU and tef1 also showed that D. bambusicola
Fig. 86 Colletotrichum
aeschynomenes (MFLU
22–0148, new host record)
a Shorea siamensis with leaf
spots, which the fungus was
isolated from b Front colony on
MEA c Reverse colony on MEA
d Orange coloured spore masses
on MEA e–j Conidiogenesis
k Conidia fused by conidial
anastomosis tubes l–o Conidia.
Scale bar. d = 1000 μm,
e–o = 10 μm
13
112
differs from D. rostrata in 10/ 826 bp (1.2%) for LSU
and 42/ 857 bp (4.9%) for tef1, differs from D. hydei in 7/
848 bp (0.8%) for LSU, and differs from D. obpyriformis in
6/ 790 bp (0.7%) for LSU and 40/ 838 bp (4.7%) for tef1.
Thus, we consider D. bambusicola as a novel species in
Distoseptispora.
Glomerellales Chadef. ex Réblová et al.
Réblová et al. (2011) validated Glomerellales to accommodate the families Australiascaceae, Glomerellaceae and
Reticulascaceae. Maharachchikumbura et al. (2016) and
Tibpromma et al. (2018) added Plectosphaerellaceae and
Malaysiascaceae to this order.
Glomerellaceae Locq. ex Seifert & W. Gams, in Zhang
et al., Mycologia 98(6): 1083 (2007) [2006].
The monotypic family is characterised by a Colletotrichum asexual morph and a Glomerella sexual morph
(Hyde et al. 2020a, b, c).
Colletotrichum Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze
Deutschl.) 3(12): 41 (1831).
Colletotrichum comprises important plant pathogens,
endophytes and saprobes as well as human and animal
pathogens (Cannon et al. 2012; Jayawardena et al. 2021a;
Talhinhas and Baroncelli 2021). Bhunjun et al. (2021)
demonstrated that coalescent approaches and multi-locus
phylogeny are vital in establishing species boundaries in
Colletotrichum. Liu et al. (2022) accepted 280 species in
16 species complexes and 15 singleton species and established a genome tree comprising 94 species. Further 13
species were added by Alizadeh et al. (2022), Hassan et al.
(2022) and Zheng et al. (2022).
Colletotrichum aeschynomenes B.S. Weir & P.R. Johnst.,
in Weir, Johnston & Damm, Stud. Mycol. 73: 135 (2012).
Index Fungorum number: IF563590; Facesofungi number: FoF11441; Fig. 86
Associated with leaf spots of Shorea siamensis. Sexual
morph: Not observed. Asexual morph: Vegetative hyphae
1.5–4.5 µm diam. ( x̄ = 3.4, n = 15), hyaline to pale brown,
smooth-walled, septate, branched. Pycnidia forming on
MEA, 500–1000 μm diam. solitary or aggregated, globose to irregular, releasing conidia in milk-orange, slimy,
glistening masses. Setae not observed. Conidiophores
8.0–53.0 × 1.5–4.5 μm ( x̄ = 25.1 × 2.8 μm, n = 20), hyaline
to light brown, cylindrical to clavate, smooth-walled, septate, sometimes branched or reduced to conidiogenous cells.
Conidiogenous cells 1.5–16 × 1.5–4 μm ( x̄ = 6.1 × 3.0 μm,
n = 15), hyaline to pale brown, solitary or aggregated,
cylindrical, ovoid or ampulliform, smooth-walled. Conidia
17.0–24.5 × 5–7.5 μm ( x̄ = 20.2 × 6.0 μm, n = 60), aseptate,
hyaline, cylindrical, clavate or ellipsoidal, with slightly
13
Fungal Diversity (2022) 117:1–272
curved basal end and mostly rounded apices, smooth-walled
or slightly verruculose, guttulate, forming conidial anastomosis tubes. Conidial anastomosis tubes 2–10.5 × 2–3 μm
( x̄ = 4.6 × 2.4 μm, n = 15), hyaline to brown, smooth-walled,
aseptate. Appressoria not observed.
Culture characteristics: Colonies on MEA flat or effuse
with entire margin. Greyish white, reverse olivatious, grey
towards the edge, reaching approximately 70 mm diam. in
5 days at 25 °C, with 20-days for sporulation. Aerial mycelium dense.
Material examined: Thailand, Chiang Mai Province,
Omkoi, on leaf spots of Shorea siamensis (Dipterocarpaceae), 15 October 2019, D. Gomdola, MFLU22-0148;
living culture MFLUCC 22-0086.
Known host and distribution: Aeschynomene virginica
(USA), Manihot esculenta (Thailand), Myrciaria dubia
(Brazil), Platostoma palustre (Taiwan), Shorea siamensis
(Thailand, this study), Theobroma cacao (Brazil) (Farr and
Rossman 2022).
GenBank numbers: OP278978 (ITS), OQ053325 (gapdh)
Notes: Weir et al. (2012) introduced C. aeschynomenes
from a stem lesion of Aeschynomene virginica. Colletotrichum aeschynomenes is a common phytopathogen
known to cause anthracnose in leaves. Listed chronologically, the pathogen has been found to infect cassava
in central Thailand (Sangpueak et al. 2018), and cause
anthracnose on Theobroma cacao in Brazil (Nascimento
et al. 2019), Myrciaria dubia in Brazil (Matos et al. 2020)
and Ixora coccinea in China (Li et al. 2021a, b). Herein,
C. aeschynomenes is associated with leaf spots of Shorea siamensis. To delineate C. aeschynomenes from C.
fructicola, tub2 or gapdh gene regions are required (Weir
et al. 2012). Phylogenetic analyses based on the concatenated ITS and gapdh sequence data depict our isolate as
C. aeschynomenes. The latter is located in the C. gloeosporioides complex and clusters with the ex-type strain
(ICMP 17673) with 71% MP, 84% ML and 0.91 BYPP
(Fig. 87). We report our collection as a new host of C.
aeschynomenes on living leaves of Shorea siamensis.
Colletotrichum flexuosum Damm, sp. nov.
Index Fungorum number: IF558527; Facesoffungi number: FoF10680; Fig. 88
Etymology: The species epithet is derived from the
shape of the setae that are often flexuous.
Holotype: CBS H-21899
Associated with leaf spots of Xanthophyllum sylvestre. Asexual morph on synthetic nutrient-poor agar
medium (SNA): Vegetative hyphae 1.5–8 µm diam.,
hyaline to pale brown, smooth-walled, septate, branched.
Chlamydospores not observed. Conidiomata absent, conidiophores and setae formed directly on hyphae. Setae
medium brown, smooth-walled, verrucous (warts 1–2 mm
Fungal Diversity (2022) 117:1–272
Fig. 87 One of two most parsimonious trees obtained with PAUP
v. 4.0b10 (Swofford 2003) from a heuristic search of the combined
sequence alignment of ITS, gapdh, chs-1, act and tub2 of the Colletotrichum gleosporioides species complex, rooted with C. truncatum
113
(CBS 151.35). Bootstrap support values of MP and ML ≥ 70% and
BYPP values ≥ 0.90 are shown at the nodes. Ex-type and reference
strains are in bold. New sequence data are in red
13
114
Fungal Diversity (2022) 117:1–272
Fig. 88 Colletotrichum flexuosum (CBS 134419, ex-type living culture) a–b. Conidiomata. c, h. Tips of setae. d, i. Bases of setae. e–g,
j–k. Conidiophores. m–r. Appressoria. s–t. Conidia. a, c–g, s. from
Anthriscus stem. b, h–r, t. from SNA. a–b. Dissecting microscope
(DM). c–t. Differential interference contrast illumination (DIC). Scale
bars: a = 100 µm, e = 10 µm. Scale bar of a applies to a–b. Scale bar
of e applies to c–t
diam.) close to the tip, 80–140 µm long, 3–4-septate,
base inflated, 6–10 µm diam., tip ± acute to round, often
strongly bent. Conidiophores pale brown, smooth-walled,
simple or septate, to 20 µm long. Conidiogenous cells
pale brown, smooth-walled, cylindrical to subglobose,
9–20(–26) × 5–9 µm, opening 2–2.5 µm diam., collarette
1 µm long, periclinal thickening conspicuous. Conidia
hyaline, smooth-walled, aseptate (few septate conidia
observed), straight, cylindrical, with both ends rounded,
(14.5–)16.5–24(–21) × (6.5–)7–8.5(–9.5) µm, mean ± S
D = 18.9 ± 2.3 × 7.8 ± 0.8 µm, L/W ratio = 2.4 (n = 30).
Appressoria single, pale brown, smooth-walled, with a
navicular to clavate outline and a lobate to crenate margin,
(9–)12.5–18(–24) × (4–)5.5–8.5(–10) µm, mean ± SD = 1
5.1 ± 2.8 × 7.1 ± 1.4 µm, L/W ratio = 2.1 (n = 30). Sexual
morph: not observed.
Asexual morph on Anthriscus stem: Conidiomata, conidiophores and setae formed on medium brown, smoothwalled, roundish to angular cells, 5–11 µm diam. Setae
medium to dark brown, smooth-walled, 100–180 µm long,
3–4-septate, straight to ± flexuous, base cylindrical or
restricted, 4–8 µm diam., tip ± round to acute. Conidiophores
pale to medium brown, smooth-walled, septate, branched,
to 90 µm long. Conidiogenous cells pale to medium brown,
smooth-walled, cylindrical to clavate, 15–32 × 5–8 µm,
opening 1.5–2 µm diam., collarette 0.5–1 µm long, periclinal thickening distinct. Conidia hyaline, smooth-walled,
aseptate, cylindrical, with both ends rounded, sometimes a
distinct membranous appendage remains at the base, (21–)
23.5–28.5(–33.5) × (5–)6–7.5(–8) µm, mean ± SD = 25.9 ±
2.6 × 6.9 ± 0.8 µm, L/W ratio = 3.8 (n = 30). Sexual morph:
not observed.
Culture characteristics: Colonies on SNA flat with entire
margin, hyaline to honey, agar medium, filter paper and
Anthriscus stem partly covered with grey acervuli and short
floccose white aerial mycelium, reverse hyaline to honey,
agar medium, filter paper and Anthriscus stem partly pale
grey; growth 13.5–18 mm in 7 days at 20 °C under near
UV light with 12 h photoperiod (21–28 mm in 10 days).
Colonies on OA flat with entire margin; buff, grey olivaceous to olivaceous grey, partly covered with short floccose
white aerial mycelium and dark grey to black spots, reverse
13
Fungal Diversity (2022) 117:1–272
Fig. 89 One of two most parsimonious trees obtained with PAUP
v. 4.0b10 (Swofford 2003) from a heuristic search of the combined
sequence alignment (gene boundaries of ITS: 1–550, tub2: 551–1240,
gapdh: 1241–1510, act: 1511–1756, chs-1: 1757–2007, his3: 2008–
2385) of the Colletotrichum gigasporum species complex, rooted
with C. gloeosporioides CBS 112999 (sequences from Damm et al.
2012; Rakotoniriana et al. 2013; Liu et al. 2014; Silva et al. 2018;
Zhou et al. 2019; Liu et al. 2022). Bootstrap support values (BS)
above 70% (bold) and Bayesian posterior probability (BYPP) values
above 0.90 are shown at the nodes. Bootstrap support values have
been calculated based on 10 000 replicates, and a Markov Chain
Monte Carlo algorithm was used to generate phylogenetic trees with
Bayesian probabilities using MrBayes v. 3.2.6 (Ronquist et al. 2012).
Numbers of ex-type strains are in bold
olivaceous-grey, growth 20–21.5 mm in 7 days (30.5–32 mm
in 10 days). Conidia in mass whitish.
Material examined: Vietnam, Ninh Bình Province, Cúc
Phương National Park, rain forest, from leaf spots of Xanthophyllum sylvestre (Polygalaceae), 6 December 2012,
Ulrike Damm (CBS H-21899, holotype); ex-holotype living culture, CBS 134419. Ninh Bình Province, Cúc Phương
National Park, rain forest, from leaf spots of Xanthophyllum
sylvestre (Polygalaceae), 6 December 2012, Ulrike Damm,
living culture CBS 137338.
GenBank numbers: CBS 134419—MZ444580 (ITS),
MZ444582 (tub2), MZ444584 (gapdh), MZ444586 (act),
MZ444588 (chs-1), MZ444590 (his3).
CBS 137338—MZ444581 (ITS), MZ444583 (tub2),
MZ444585 (gapdh), MZ444587 (act), MZ444589 (chs-1),
MZ444591 (his3).
115
Notes: Colletotrichum flexuosum was isolated from leaves
of Xanthophyllum sylvestre, a tree species native to Laos,
Thailand and Vietnam (http://www.plantsoftheworldonline.
org). Few fungi were previously reported from Xanthophyllum, including an unidentified Colletotrichum species on X.
octandrum in Australia (Simmonds 1966; Farr and Rossman
2022). No Colletotrichum species was previously described
or reported from X. sylvestre.
Based on blastn searches and sequence comparisons on
NCBI GenBank, the closest neighbour of the strains from
Xanthophyllum is C. vietnamense, belonging to the C. gigasporum complex. The ITS, tub2, gapdh, act, chs-1 and his3
sequences of this species were 80, 97, 91, 98, 98 and 96%
identical (105, 23, 9, 4, 4and 15 nucleotides difference)
with those of the ex-holotype strain of C. vietnamense (Liu
et al. 2014). No strain was more than 82% identical with its
ITS sequences. The closest match with the act sequences
was Colletotrichum sp. gnqczg15 (KC293585, F. Huang,
unpubl. study) with two nucleotides difference. In a phylogeny inferred from concatenated ITS, tub2, gapdh, act, chs-1
and his3 sequences of the C. gigasporum species complex
(Fig. 89), the two strains formed a well-supported (BS 100%,
BYPP 1) sister clade to C. vietnamense and a clade formed
by two recently described species, C. subvariabile and C.
variabile (Liu et al. 2022). Colletotrichum flexuosum can be
identified with all loci included.
The species from Xanthophyllum is morphologically different from all previously described species of the C. gigasporum complex. Its conidia are shorter than those of C.
gigasporum (22–32 × 6–9 μm on PDA), C. magnisporum
(av. 34.3 × 9.7 μm on SNA, av. 33.8 × 9.9 μm on Anthriscus
stem), C. serranegrense (24–37 × 7–9 μm on MEA) and C.
vietnamense (av. 31.2 × 9.6 μm on SNA, av. 32.3 × 9.5 μm on
Anthriscus stem) and larger than those of C. jishouense (av.
10.8 × 3.7 μm on PDA). In contrast to C. arxii, C. jishouense,
C. pseudomajus, C. radicis and C. vietnamense, neither
curved nor clavate conidia were formed, while conidia with
membranous appendages and distinctly flexuous setae as
formed by C. flexuosum were previously not observed in
the C. gigasporum complex (Rakotoniriana et al. 2013; Liu
et al. 2014, 2022; Silva et al. 2018; Zhou et al. 2019). Conidiogenous cells with a distinct periclinal thickening are typical for species of the C. boninense complex; within the C.
gigasporum complex this was previously only observed with
C. zhaoqingense (Liu et al. 2022). However, in contrast to C.
zhaoqingense, conidial bases of C. flexuosum sometimes end
with a membranous appendage, while most species of the C.
boninense and the C. dracaenophilum complexes develop a
prominent scar (Damm et al. 2012, 2019; Liu et al. 2022).
Colletotrichum pandanicola Tibpromma & K.D. Hyde, in
Tibpromma et al., MycoKeys 33: 47 (2018).
13
116
Fungal Diversity (2022) 117:1–272
Fig. 90 Colletotrichum pandanicola (MFLU 18–1852, new
host record) a Dead leaf of
Mangifera indica. b Germinating spore. c Reverse view of the
7d old culture on PDA. d Upper
view of the 7d old culture on
PDA. e Setae. f Vegetative
hyphae. g Conidiogenous cell. h
Conidia. i–j Appressoria. Scale
bars: e = 50 μm, b = 10 μm;
scale bar of b applies to f–j
Index Fungorum number: IF823841; Facesoffungi number: FoF05832; Figs. 90, 91
Saprobic on Mangifera indica and pathogenic on Alstonia
scholaris. Sexual morph: Not observed. Asexual morph:
On PDA vegetative hyphae greyish white, dense, cottony.
Conidiomata acervuli, black, circular to oval, submerged,
solitary or aggregated. Sporulation abundant. Setae scattered, straight or ± bent, dark brown up to the tip, opaque,
2- to 4-septate, 50–80 µm long, smooth-walled, base cylindrical, 2–5 µm diam., tip acute. Conidiophores hyaline to
light brown, cylindrical to clavate, smooth-walled, simple,
occurring in densely arranged clusters. Conidiogenous
cells enteroblastic, hyaline, smooth-walled, cylindrical to
slightly inflated, periclinal thickening not visible. Conidia
7–12 × 2–6 μm ( x̄ = 7 × 4 μm, n = 40), hyaline, smoothwalled, aseptate, ovoid, cylindrical or clavate with rounded
apices, guttulate. Appressoria 9–16 × 5–6 μm ( x̄ = 13 × 5 μm,
n = 10), solitary to aggregated, in small groups or short
chains, medium to dark brown, smooth-walled, irregular,
rarely lobed.
13
Culture characteristics: Colonies grown from single
conidia on PDA 50–70 mm diam. in 7 days, at first white
becoming dark grey, reverse pale yellow.
Material Examined: Thailand, Chiang Rai Province, on
leaf of Mangifera indica L. (Anacardiaceae), 19 May 2018,
Ruvishika S. Jayawardena (MFLU 18-1852), living culture
MFLUCC 18-1182. India, Karnataka, Mysuru on infected
leaves of Alstonia scholaris (Apocyanaceae) as pathogen,
May 2019, S. Mahadevakumar (UOM-IOE 19/15), living
cultures (AsCs1, AsCs4).
GenBank numbers: MFLUCC 18-1182-MK629453 (ITS),
MK639363 (gapdh), MK639357 (chs-1), MK639359 (act)
and MK639361 (tub2)
AsCs1–OM912803 (ITS), OM934812 (gapdh),
OM934814 (tub2)
AsCs4–OM912804 (ITS), OM934813(gapdh),
OM934815 (tub2)
Notes: The species of the C. gloeosporioides species
complex are mainly known as plant pathogens (Weir et al.
2012; Jayawardena et al. 2016, 2018, 2021a, b; Bhunjun
et al. 2021), some species are also as endophytes or as
Fungal Diversity (2022) 117:1–272
117
Fig. 91 a–b Leaf blight/early
symptoms of anthracnose
disease on Alstonia scholaris.
c Complete death of infected
branch due to anthracnose
disease. d–e anthracnose symptoms appearing at the tip of the
leaves caused necrosis. f a close
view of anthracnose symptoms.
g pure cultures of 10 days old
Colletotrichum siamense on
PDA. i–k Conidiogenous cells
and conidia of C. siamense
observed under a compound
microscope (Scale bar: 10 µm)
saprobes (Jayawardena et al. 2021a, b). Strain MFLUCC
18-1182 belongs to the Colletotrichum gloeosporioides species complex and clusters with the ex-type of C. pandanicola
(Fig. 87). However, strain MFLUCC 18–1182 differs from
C. pandanicola by forming setae and appressoria, which
were not observed in the ex-type strain of C. pandanicola
(Tibpromma et al. 2018). This represents the first report of
C. pandanicola on Mango from Thailand.
Colletotrichum pandanicola was isolated from leaves
of Alstonia scholaris, is an evergreen tropical tree and an
important medicinal plant distributed throughout the Indian
peninsula including Western Ghats regions of Karnataka.
Colletotrichum gleosporioides has been reported as a pathogen on A. scholaris (Chandra 1974; Mathur 1979; Sarbhoy
and Agarwal 1990; Sarbhoy et al. 1971). Pathogenicity
tests were performed on healthy A. scholaris plants. After
10–12 days of pot inoculation, the initial necrotic spots
developed on leaves were smaller and later coalesced to
form larger necrotized lesions. The lesions developed on
inoculated plants were fusiform and grayish brown. Phylogeny supports (Fig. 87) our strain to be C. pandanicola.
This is a new report of C. pandanicola on Alstonia scholaris
from India.
Colletotrichum thasutense Armand, K.D. Hyde and Jayaward., sp. nov.
Index Fungorum number: IF900127; Faceoffungi number: FoF13361; Fig. 92
Etymology: Referring to the sub-district where the specimen was collected.
Holotype: MFLU 22-0206
Associated with leaf spot on Syngonium sp. Sexual
morph: not observed. Asexual morph: On host: Acervuli
present, setose, creamy to greyish white, subepidermal,
produced solitary. Conidiophores rarely observed, hyaline, barrel shaped, 6.5–9 × 3–4 μm; Conidiogenous cells
hyaline, cylindrical; 10.5–23 × 2.5–4 μm (x̄ = 16 × 3 μm,
n = 20). Conidia aseptate, hyaline, smooth-walled, cylindrical, straight to slightly curved, mostly rounded ends, rarely
obtuse at one end, 13.5–17.5 × 3.5–5 μm (x̄ = 15.5 × 4.5 μm,
n = 20), Appressoria not observed.
On PDA: Acervuli, Setae and Sclerotia absent.
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Fungal Diversity (2022) 117:1–272
Fig. 92 Colletotrichum
thasutense (MFLU 22–0206,
holotype) a Host plant. b Host
leaf with symptom. c Acervuli
on the host. d setae. e–g Conidiogenous cells and conidia. h–j
Conidia. Scale bars: d = 10 µm,
e–g = 5 µm, h-j = 10 µm
Culture characteristics: Colonies on PDA white to pale
yellow, reverse same color, reaching 70 mm diam. in 7 days
at 28 °C. Colonies cottony, circular, slightly raised and
depressed in the center, Aerial mycelia white and medium
in dense.
Material examined: Thailand, Chiang Rai, Mueang,
ThaSut, from Syngonium sp. (Araceae), associated with leaf
spot; 16 October 2021, A. Armand, A2 (MFLU 22-0206,
holotype), living ex-type living culture MFLUCC 22-0173.
GenBank numbers: OP821902 (ITS), OP831280 (act),
OP831281 (chs-1),: OP831282 (gapdh), OP831283 (tub2).
Notes: Based on the phylogenetic analyses, Colletotrichum thasutense is closely related to C. xishuangbannaense
(Fig. 87). Morphologically, C. thasutense differs from C.
xishuangbannaense by producing longer and thicker conidia.
Colletotrichum xishuangbannaense has 9–12 × 3–4 µm
conidia (de Silva et al. 2021), whereas C. thasutense has
13.5–17.5 × 3.5–5 μm conidia. Colletotrichum thasutense
can be differentiated from C. xishuangbannaense in having slightly curved conidia and obtuse at one end. However, C. xishuangbannaense produces straight conidia with
rounded ends. Moreover, C. thasutense bears longer and
13
thicker conidiogenous cells (10.5–23 × 2.5–4 μm) than C.
xishuangbannaense (15–18 × 1.5–2 µm).
Hypocreales Lindau
Hypocreales includes 15 families, which are considered
subtropical and tropical, namely Bionectriaceae, Calcarisporiaceae, Clavicipitaceae, Cocoonihabitaceae, Cordycipitaceae, Cylindriaceae, Flammocladiellaceae, Hypocreaceae,
Myrotheciomycetaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae, Sarocladiaceae, Stachybotryaceae, and
Tilachlidiaceae, based on the basis of molecular evidence
(Hyde et al. 2020a, b, c; Wijayawardene et al. 2022). These
families differ in the presence of fleshy and colorful perithecial ascomata, ostiolate perithecia, varied ascospores, and
pigment production. It has been estimated that Hypocreales
diverged 229 million years ago (Hyde et al. 2020a, b, c).
Nectriaceae Tul. & C. Tul. [as 'Nectriei'], Select. fung. carpol. (Paris) 3: 3 (1865)
Nectriaceae was introduced by Tulasne and Tulasne
(1865) and re-evaluated by Lombard et al. (2015) based
on morphology and ten genes (acl1, act, cmdA, his3, ITS,
Fungal Diversity (2022) 117:1–272
119
Fig. 93 Cultural and morphological features of Fusarium
brachygibbosum: a–c pure
cultures of F. brachygibbosum isolated on PDA medium
(12 days old) (a–b Front view,
c– reverse view); d–h Microscopic view of F. brachyggibosum conidia structures observed
under compound microscope.
Scale bar: d–h = 20 µm
LSU, rpb1, rpb2, tef1 and tub2). This family includes 74
genera that are recognized by different perithecial pigments (Wijayawardene 2022; Lombard et al. 2015; Crous
et al. 2021). The family contains a wide range of species,
including plant pathogens, human pathogens, and industrial
and commercial species (Rossman 1996; Luo and Zhuang
2008; Chaverri et al. 2011; Lechat et al. 2015; Lombard
et al. 2015).
Fusarium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–
2): 10 (1809).
Fusarium is famous for its difficulty in identification
and pathogenicity. It has been extensively discussed in
recent decades as one of the most taxonomically confusing genus within Nectriaceae (Gräfenhan et al. 2011;
O'Donnell et al. 2013; Crous et al. 2021; Geiser et al.
2021). Crous et al. (2021) and Wang et al. (2022) have
updated the taxonomy of this genus to include 19 species
complexes. In addition, a variety of species within this
genus contain virulent crop pathogens, such as F. graminearum and F. oxysporun, two of the top ten most economically damaging fungal pathogens (Dean et al. 2012; Leslie
and Summerell 2006).
Fusarium brachygibbosum Padwick, Mycological Papers
12: 11 (1945).
Index Fungorum number: IF286508; Faceoffungi number: FoF11683; Figs. 93, 94
Pathogenic on roots of Vigna unguiculata. Sexual
morph: Not observed. Asexual morph: Conidiophores
27–58 µm long, carried on aerial mycelium, unbranched or
irregularly and/or sympodially branched bearing a terminal phialide. Conidiogenous cells 8–22 × 2–4 µm, polyphialide, subulate to subcylindical, smooth. Macroconidia
15.2–22 × 2–3 µm, hyaline, slightly curved with five distinct septa, wide central cells, slightly sharp apexes, basal
cells with foot like shape. Microconidia rarely observed.
Chlamydospores 6–24 µm diam. abundant, spherical o
globose, smooth, slightly verrucose, formed terminally or
intercalary in chains of two or three, wall 1–1.5 µm.
Culture characteristics: Colonies on PDA reaching
90 mm at 28 °C after 14 d in 12/12 dark, colonies appeared
white to pink with abundant aerial mycelium.
Materials examined: India, Karnataka, Mysuru, Doddamaragowdanahally, diseased root of cowpea (Vigna
unguiculata (L.) Walp. Fabaceae), May 2019, S. Mahadevakumar & Y.S. Deepika (UOM-IOE 19/16), living cultures
CPFb1, CPFb2, CPFb3, CPFb4.
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Fungal Diversity (2022) 117:1–272
Fig. 94 Morphological features
of Fusarium brachygibbosum:
a–b Conidial morphology of
F. brachygibbosum under compound microscope; c–d hyphal
structures and chlamydospores
of F. brachygibbosum; e–f a
single macroconidium enlarged.
Scale bar: a–d 50 µm; e–20 µm;
f– g 10 µm
Hosts: Wide host range, including Allium, Beta, Cannabis, Citrullus, Citrus, Euphorbia, Glycine, Gossypium, Helianthus, Nerium, Nicotiana, Phoenix, Plasmopara, Prunus,
Sansevieria, Sorghum, Triticum, Vigna and Zea (Farr and
Rossman 2022; this study).
Distribution: Wide geographical range, including in
Australia, Azerbaijan, China, India, Iran, Malaysia, Oman,
Qatar, Soudi Arabia, Tunisia, Turkey and United States (Farr
and Rossman 2022; this study).
GenBank numbers: CPFb1– MT804589 (ITS),
OM938019 (tef1)
CPFb2– MT804590 (ITS), OM938020 (tef1)
CPFb3– MT804591 (ITS), OM938021(tef1)
CPFb4– MT804592 (ITS), OM938022 (tef1)
Notes: Fusarium brachygibbosum is known to associated
with 19 host plants of which two records are represented
from India (Sorghum vulgare, Plasmopara viticola) (Farr
and Rossman 2022). This is the first record of F. brachygibbosum recorded on Cowpea (Fabaceae) from India (new host
record) (Fig. 95).
Fusarium purpurea S.L. Han, M. Raza, W.J. Duan & L.
Cai, sp. nov.
13
Index Fungorum number: IF555883; Facesoffungi number: FoF10818; Fig. 96
Etymology: Refers to the pigment produced on PDA and
OA, purpure.
Holotype: HMAS 351947.
Asexual morph: Hyphae 1.7–3.5 μm diam, hyaline,
smooth-walled, septate, branched. Conidiophores arises
on aerial mycelium, unbranched or irregularly branched,
11–29.5 × 2–4.5 μm ( x̄ = 19.9 × 3.0 μm, n = 35). Phialides
mono- and polyphialide, subulate to subcylindrical, smoothand thin-walled, 4.8–17.4 × 1.6–3.9 μm, periclinal thickening inconspicuous or absent. Microconidia hyaline, smoothand thin-walled, two types, clavate with truncate base
conidia (aseptate): 4.5–9.5 × 1.5–3.5 μm ( x̄ = 6.3 × 2.5 μm,
n = 50); globose conidia (aseptate): 6.5–11 × 7–11.5 μm
( x̄ = 9.7 × 9.4 μm, n = 50). Sporodochia and chlamydospores
not observed. Sexual morph: Not observed.
Culture characteristics: Colonies on PDA slow growing,
reaching 52–57 mm diam in 7 d after incubation at 25 °C in
the dark, colony flat, medium, filamentous, felted to velvety,
rhizoid; colony from above; raised, dull, wrinkled folded,
surface orchid purple (14C8) in the center, white (–A1) at
the margin; reverse beetroot purple (13D8) in the center,
Fungal Diversity (2022) 117:1–272
121
Fig. 95 RAxML phylogenetic tree generated from tef1 sequence data
of Fusarium sambucinum species complex. Maximum likelihood
bootstrap support ≥ 75% (in blue) and BYPP ≥ 0.95 (in green) are
indicated on the branches. Ex-type and epi-type cultures are indicated
in bold with ‘T’ and ‘ET’
white (–A1) at the margin; odour absent, not producing pigment in PDA media. On OA reaching 41–49 mm in 7 d after
incubation 25 °C in the dark; raised, felted to dusty, with
abundant aerial mycelium, margin entire; surface amethyst
(15C6) in the center, white (–A1) at the margin; reverse oak
brown (5D6); odour absent.
Material examined: Kazakhstan, intercepted at
Alashankou Port, isolated at Ningbo Customs, from seeds
of Triticum aestivum imported to China, Jan. 2019, W.J.
Duan & W.Z. Li (HMAS 351947, holotype), ex-type living
culture, CGMCC 3.23515 = LC15871. ibid., LC15872; ibid.,
LC15873; ibid., LC15874.
GenBank numbers: CGMCC 3.23515 – ON365812
(CaM), ON365816 (rpb1), ON365820 (rpb2), ON365828
(tef1), ON365824 (tub2)
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Fungal Diversity (2022) 117:1–272
Fig. 96 Fusarium purpurea (CGMCC 3.23515, ex-type living culture). a Colony on PDA. b Colony on OA. c-d Aerial conidiophores and phialides. e–f Microconidia. Scale bars = 10 μm
Fig. 97 RAxML phylogenetic
tree generated from combined
CaM, rpb1, rpb2, tef1 and tub2
sequence data of Fusarium
fujikuroi species complex.
Maximum likelihood bootstrap
support values greater than 75%
(in blue) and Bayesian posterior
probabilities > 0.95 (in green)
are indicated on the branches.
Ex-type and epi-type cultures
are indicated in bold with ‘T’
and ‘ET’
LC15872 – ON365813 (CaM), ON365817 (rpb1),
ON365821 (rpb2), ON365829 (tef1), ON365825 (tub2)
13
LC15873 – ON365814 (CaM), ON365818 (rpb1),
ON365822 (rpb2), ON365830 (tef1), ON365826 (tub2)
Fungal Diversity (2022) 117:1–272
LC15874 – ON365815 (CaM), ON365819 (rpb1),
ON365823 (rpb2), ON365831 (tef1), ON365827 (tub2)
Notes: Fusarium purpurea formed a well-supported sister clade to F. globosum with 100% ML and 1.00 Bayesian
posterior probabilities (BYPP) support (Fig. 96). Fusarium
purpurea differs by 3 bp in the CaM gene, 16 bp in the
rpb2 gene, 6 bp in the tef1 gene, and 1 bp in the tub2 gene
compared to F. globosum (Proctor et al. 2013; Yilmaz et al.
2021). Morphologically, F. purpurea differs in the types of
microconida production and its number of septation. For
example, F. purpurea produces two types of microconidia:
clavate with a truncate base (aseptate) and globose (aseptate) without papilla, while three types of microconidia were
found in F. globosum: clavate with a truncate base (0- to
3-septate), napiform/pyriform, and globose (0- to 1-septate)
which often have a distinct papilla (Rheeder et al. 1996;
Leslie and Summerell. 2006) (Fig. 97).
123
Microascales composed of seven families namely Ceratocystidaceae, Chadefaudiellaceae, Gondwanamycetaceae,
Graphiaceae, Halosphaeriaceae, Microascaceae and Triadelphiaceae (Maharachchikumbura et al. 2016; Hyde et al.
2020a, b, c; Wijayawardene et al. 2022). A total of 109 genera were accepted under Microascales which were distributed in seven families (Wijayawardene et al. 2022).
Microascaceae Luttr. ex Malloch, Mycologia 62(4): 734
(1970)
Microascaceae was confined by Luttrell (1951) in Microascales and authenticated by Malloch (1970) (Maharachchikumbura et al. 2016). It includes saprobic, plant and opportunistic human pathogenic fungal genera (de Hoog et al.
2000; Sandoval-Denis et al. 2016; Maharachchikumbura
et al. 2016). Currently this family is composed of 23 genera
(Wijayawardene et al.2022).
Microascales Luttr. ex Benny & R.K. Benj. (1980).
Scedosporium Sacc. ex Castell. & Chalm., Manual of Tropical Medicine: 1122 (1919)
Fig. 98 Scedosporium apiospermum (MFLU 22-0160, new record)
a. Contaminated oyster mushroom grow substrate. b,c. Colony on
PDA. d. Sporulation of the colony on PDA. e-i. Conidial attachments
and conidiogenous cells. j,k. Synnema with conidia. l. Conidia. Scale
bars: a = 5 cm, b,c = 30 mm, d = 200 μm, e,f,l = 4 μm, g = 20 μm,
h = 35 μm, i = 25 μm, j = 65 μm, k = 40 μm
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Fungal Diversity (2022) 117:1–272
◂ Fig. 99 Phylogram generated from maximum likelihood analysis
based on combined ITS and tub2 sequence data representing the
species of Scedosporium and related genera. Related sequences are
taken from Zhang et al. (2021). Sixty-two taxa are included in the
combined analyses. Microascus longirostris (CBS 196.61), Scopulariopsis brevicaulis (MUCL 40,726), Wardomyces anomalus (CBS
299.61) and Wardomyces giganteus (CBS 746.69) are used as the outgroup taxa. Bootstrap support values for ML ≥ 70% and BYPP ≥ 0.95
are given near the nodes. The newly generated sequence is in blue.
The type strains are indicated in black bold
Scedosporium is a ubiquitous filamentous fungus with
a worldwide distribution. Pseudallescharia boydii is the
sexual morph of Scedosporium apiospermum, which was
first discovered in 1889 as a causative agent of human otitis
(Siebenmann et al. 1899). Later Monosporium apiospermum
was discovered as an anamorphic state of P. bodydii from a
patient with human mycetoma in 1919 (Shear 1922).
This genus is typified by a sexual morph (Pseudallescheria boydii) characterized by closed ascomata (cleistothecia),
a peridium (ascomata wall) of ‘textura epidermoidea’, asci
that are broadly clavate or spherical, and ascospores that
are ellipsoidal or fusiform, which are symmetrical or nearly
so (von Arx et al. 1988; De Hoog et al. 2000). The asexual
morph includes: Scedosporium, characterized by hyaline,
cylindrical conidiogenous cells arising from undifferentiated hyphae that produce obovoidal, hyaline, sticky conidia.
Synnemata are characterized by large, erect bundles of
hyphae terminating in a dense aggregate of conidiogenous
cells. It produces conidia from a short extension of the conidiogenous cells with annellidic development (Gueho 1991;
Lackner et al. 2014; Ramirez-Garcia et al. 2018).
Scedosporium species have been reported commonly
from natural substrates such as soil, water anthropogenic
influenced habitats, cattle dung and sewage (De Hoog et al.
2000; Ramirez-Garcia et al. 2018). Seventeen species are
accepted under Scedosporium and these include: S. americanum, S. angustum, S. apiospermum, S. aurantiacum, S.
boydii, S. cereisporum, S. deficiens, S. dehoogii, S. desertorum, S. fusoideum, S. haikouense, S. hainanense, S. magalhaesii, S. minutisporum, S. multisporum, S. rarisporum and
S. sclerotiale (http://www.indexfungorum.org). The disease
caused by Scedosporium species is termed as Scedosporosis.
The occurrence of Scedosporium apiospermum associated
with mushroom cultivation and Scedosporium marina and
S. dehoogii from the marine environment are reported here.
Scedosporium apiospermum Sacc. ex Castell. & Chalm.,
Manual of tropical medicine (London): 1122 (1919).
Index Fungorum number: IF432048; Facesoffungi number: FoF11704; Fig. 98
Growing on Oyster mushroom grow substrate. Sexual
morph: Not observed. Asexual morph: on host, Mycelium
black powdery mass. On PDA, hyphomycetous, Hyphae
125
1.5–3.5 μm ( x̄ = 2.2 μm) wide, branched, septate, hyaline.
Conidiophores solitary or synnematous, solitary conidiophores 75–128 × 1.8–3 μm ( x̄ = 102.5 × 2.6 μm, n = 10)
hyaline, branched, forms 1–3 conidiogenous cells at the
end. synnematous conidiophores 132–352 × 8.2–17.5 μm
( x̄ = 288.4 × 12.7 μm, n = 15) cylindrical stipe, erect, forms
conidia at the end. Conidiogenous cells 4.8–25 × 1.2–2.5 μm
( x̄ = 15.2 × 1.3 μm, n = 10) lateral or terminal formation on
solitary conidiophores, hyaline, cylindrical. Conidia hyaline, guttulate, conidia arising from conidiogenous cells
6.2–8.2 × 2–4.5 μm ( x̄ = 7.4 × 3.3 μm, n = 30) obvoid to
ellipsoidal, conidia arising from synnematous conidiophores
6.4–9.2 × 1.7–3.5 μm ( x̄ = 7.5 × 2.8 μm, n = 35) cylindrical or
claviform with a truncate base, conidia from undifferentiated
hyphae 6–7.5 × 5.2–6.6 μm ( x̄ = 6.4 × 5.2 μm, n = 20) sessile,
globose to subglobose.
Culture characteristics: colonies on PDA attaining
65–75 mm diameter after 14 days at 25 °C, cottony, dense,
lanose, light gray to dark grey, margins fimbriate and irregular; reverse dark grey to brown, grey to white margins.
Material examined: Thailand, Phayao Province, growing on the oyster mushroom growing substrate, 06 June
2020, AJ. Gajanayake, AJ 072 (inactive dry culture, MFLU
22-0160, new host record), living culture, MFLUCC
22-0192.
GenBank numbers: ON714510 (ITS), ON714511(LSU),
ON730889 (tub2).
Notes: Our isolate MFLUCC 22-0192, clusters within
Scedosporium apiospermum with a 90% ML and 0.97 BYPP
support (Fig. 99). Furthermore, the morphology of our isolate MFLUCC 22-0192 resembles the original description
and illustrations for Scedosporium apiospermum by Saccarado in Castellani and Chalmers (1919) and the description
and morphological illustrations by Gilgado et al. (2010).
However, there are size differences of conidiophores, conidiogenous cells and conidia, when we compare our strain
with the strains described in Castellani and Chalmers (1919)
and Gilgado et al. (2010). The reason for this may be the
differences in the media in which the colonies were grown.
Scedosporium apiospermum has been mainly identified
as an opportunistic clinical pathogen relevant to many infections which commonly occur in the bones, central nervous
system, lungs, paranasal sinuses, skin and soft tissues (Shinohara and George (2009); Goldman et al. 2016). Seephueak
et al. (2017) isolated 21 fungal species from spent mushroom substrate of Pleurotus sp. collected from mushroom
farms in southern Thailand. Among those 21 fungal species
there were, Alternaria spp., Aspergillus spp., Chaetomium
sp., Cunninghamella spp., Fusarium spp., Lasiodiplodia
sp., Neurospora sp., Penicillium spp., Rhizoctonia sp. and
Trichoderma spp. (Seephueak et al. 2017). Suada et al.
(2015) have reported Aspergillus spp., Fusarium spp., Gliocladium sp., Mucor spp., Neurospora spp., Paecilomyces sp.,
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Fungal Diversity (2022) 117:1–272
Fig. 100 Scedosporium marina
on a twig (AMH-9946, holotype). a. Colonies on decaying
woody stem of Suaeda monoica.
b − c Synnemata. d − e, h Head
and conidiophores. f Apical part
of synnema producing conidia.
g Conidiogenous cells with
annellidic conidia. i Subcylindrical conidia. k − l Claviform conidia. m − o Obovoid
conidia. Scale bars: b = 50 µm,
c = 100 µm d − o = 10 µm
Penicillium spp., Pythium sp., Stachybotrys spp. and Trichoderma spp. as contaminants from oyster mushroom growing
substrate. According to best our knowledge this is the first
report of Scedosporium apiospermum as a contaminant of
oyster mushroom growing substrate.
Scedosporium marina Devadatha & V.V Sarma, sp. nov.
Index Fungorum number: IF558432, Facesoffungi number: FoF05035; Figs. 100, 101
Etymology: The specific epithet is in reference to the
marine environment in which the fungus was collected.
13
Holotype: AMH-9946.
Saprobic on decaying woody stem of the halophyte
Suaeda monoica. Asexual morph: Colonies effuse,
light brown. Mycelium immersed, composed of septate, branched, smooth, pale brown to hyaline hyphae.
Synnemata solitary to gregarious, erect, dark brown,
170–1110 µm tall with a cylindrical stipe, 10–25 µm wide
( x̄ = 642 × 16 µm, n = 10), dark gray, smooth-walled, terminate into a slimy head of conidia, slimy head 60–140 µm
long and 60–165 µm wide ( x̄ = 86 × 111 µm, n = 10).
Hyphae interwoven at the base, unbranched in the stipe,
Fungal Diversity (2022) 117:1–272
127
Fig. 101 Scedosporium marina
in culture (NFCCI-4273).
a Synnemata on PDA b − c
Synnemata. g − h Culture on
PDA after 14 days. d − e Slimy
head and Conidiophores. f
Conidiogenous cells with annellidic conidia. l Apical part of
synnema producing conidia. m
Subcylindrical conidia. n − r,
s Sessile obovoid conidia o − q
Claviform Conidia. Scale bars:
b − c = 100 µm d = 50 µm, e − f,
i − s = 10 µm
branching at the apex to form conidiophores. Conidiophores synnematous, solitary, branched, often reduced to
conidiogenous cells, growing laterally bearing a single verticil conidiogenous cell. Conidiogenous cells percurrent,
terminal or lateral, hyaline, smooth-walled, cylindrical to
slightly flask-shaped, 25–55 × 2–2.5 µm ( x̄ = 35 × 2.25 µm,
n = 10). Conidiogenous cells arising from undifferentiated hyphae are cylindrical to slightly flask-shaped, producing slimy heads of one-celled, smooth-walled, sub
hyaline, obovoid or sub-cylindrical conidia. Three types
of conidia are produced: (i) those produced on solitary
conidiophores sub hyaline, smooth-walled, obovoid, or sub
cylindrical 7–13 × 2.5–5.5 µm ( x̄ = 10 × 4 µm, n = 20); (ii)
those produced on synnemata predominantly cylindrical
or claviform, 5–13 × 1–3 µm ( x̄ = 9 × 2 µm, n = 20) with a
wide truncate base; (iii) those developed mainly from the
undifferentiated hyphae of the substrate, sessile or on short
13
128
protrusions, solitary, lateral, brown, smooth, and thickwalled, mostly obovoid 2.5–10 × 2–2.5 µm ( x̄ = 9 × 2.3 µm,
n = 20). Sexual morph: Not observed.
Culture characteristics: Conidia germinating on sea
water agar within 24 h. Germ tubes produced from the
conidial base. Colonies on PDA attaining 30–35 mm
diameter after 14 days at 25 °C, circular, raised, light
grayish at center and smoke gray at margins, cottony; yellow in reverse with entire margin. The optimum growth
temperature was from 25 to 37 °C, and did not grow at
45 °C. Hyphae hyaline to pale brown, branched and septate, 3–7.5 µm wide. Synnemata developed after 30 days
of incubation at 25 °C; synnemata solitary to gregarious, erect, dark brown, 190–1400 µm tall with a cylindrical stipe from 10 to 25 µm wide ( x̄ = 647 × 16.9 µm,
n = 10), dark gray, smooth walled, terminated into a slimy
head of conidia, 25–35 µm long and 35–50 µm wide
( x̄ = 30 × 36 µm, n = 10). Conidiophores solitary, branched,
often reduced to conidiogenous cells growing laterally
bearing single verticil of conidiogenous cells. Conidiogenous cells terminal or lateral, hyaline, smooth-walled,
cylindrical to slightly flask-shaped, 25–50 × 2–3 µm
( x̄ = 35 × 16 µm, n = 10). Conidiogenous cells arising
from undifferentiated hyphae, cylindrical to slightly flaskshaped, producing slimy heads of one-celled, smoothwalled, sub-hyaline, obovoid or sub-cylindrical conidia.
There were three types of conidia: (i) those produced on
solitary conidiophores subhyaline, smooth-walled, obovoid or subcylindrical 5–10 × 1.5–3 µm ( x̄ = 6.8 × 2.3 µm,
n = 20); (ii) those produced on synnemata predominantly cylindrical or claviform, 7.5–10 × 2–3 µm
( x̄ = 8.7 × 2.4 µm, n = 20) with a wide truncate base; (iii)
those developing mainly from the undifferentiated hyphae
of the substrate, sessile or on short protrusions, solitary,
lateral, brown, smooth, and thick-walled, mostly obovoid
5–10 × 2–2.5 µm ( x̄ = 9 × 2.4 µm, n = 20). Sexual state not
observed after incubation for 2 months at 25 °C.
Material examined: India, Tamil Nadu, Tiruvarur, Muthupet mangroves (10.4° N 79.5° E), on decaying woody stem
of the halophyte Suaeda monoica Forssk. ex J.F.Gmel.
(Amaranthaceae) 28 November 2015, B. Devadatha (AMH9946, holotype), ex-type living culture NFCCI-4273.
GenBank numbers: MF182397 (ITS), KY863508 (LSU),
MH571780 (SSU), MF687078 (tub2), MF182399 (tef1)
Notes: Our present collection of Scedosporium marina
(NFCCI-4273) has been assigned to Scedosporium based
on its similar morphological characteristics in having
Graphium-like synnemata and scattered, poorly differentiated, percurrent conidiogenous cells (Gueho 1991). The
present taxon, Scedosporium marina (NFCCI-4273) and
S. aurantiacum (FMR8630) share similar morphological characteristics in producing three different types of
conidia like the obovoid, or sub cylindrical, cylindrical
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or claviform, sessile obovoid conidia with overlapping
conidial dimensions and yellow diffusible pigments
on PDA. However, combined multigene phylogenetic
anlayses of combined datasets of ITS and tub2 revealed
that Scedosporium marina (NFCCI-4273) formed distinct lineage sharing a sister relation with Scedosporium
aurantiacum with significant statistical support 100%
ML, 97% MP (105). Morphologically S. marina (NFCCI4273) is distinct from S. aurantiacum (FMR8630) in
having long and wide synemmata (170–1110 × 10–25 vs
330–750 × 7.5–17.5), smaller heads (60–140 × 60–165
vs 60–70 × 140). The condiogenous cells of S. marina
(NFCCI-4273) are percurrent and long in contrast to S.
aurantiacum (FMR8630) (25–55 × 2–2.5 vs 10–37 × 1.5
2.5).
The optimum growth temperature of S. marina (NFCCI4273) was from 25 to 37 °C and did not grow at 45 °C
whereas S. aurantiacum (FMR8630) was from 37 to 40 °C
and growth was also found at 45 °C but the fungus did not
grow at 50 °C (Gilgado et al. 2005). The ITS sequence comparison revealed more than 30 nucleotide base pair differences between two taxa, which supports the establishment
of a new species for our new taxon. Furthermore, S. marina
is a saprobe on decaying woody stem of the halophyte
Suaeda monoica from a marine habitat unlike other species
of Scedosporium which are known to be human pathogens.
Scedosporium aurantiacum is clearly distinguished from S.
marina as an opportunistic human pathogen that is known
to cause various infections in lungs, ears, respiratory sinuses
and subcutaneous abscess in patients of diabetes and malignant lymphoma (Kondo et. al 2018). Hence, based on the
above mentioned morphological, cultural, molecular and
habitat differences we introduce a new species, S. marina,
in Scedosporium.
Scedosporium dehoogii Gilgado, Cano, Gene´ Guarro in
Journal of Clinical Microbiology 46: 2 (2008).
Index Fungorum number: IF538388; Facesoffungi number: FoF04829; Fig. 102, 103
Saprobic on decaying woody of Avicennia marina. Asexual morph: Colonies effuse, light brown. Mycelium mostly
immersed, composed of septate, branched, smooth, pale
brown to hyaline hyphae. Synnemata solitary to gregarious,
erect, dark brown, 190–390 µm tall with a cylindrical stipe
from 10 to 30 µm wide ( x̄ = 242 × 16 µm, n = 10), dark gray,
smooth-walled, terminated into a slimy head of conidia,
slimy head 45–75 µm long, 55–80 µm wide ( x̄ = 60 × 64 µm,
n = 10). Hyphae interwoven at the base, unbranched in the
stipe, branched at the apex to form conidiophores. Conidiophores synnematous, solitary, branched, often reduced to
conidiogenous cells which were subhyaline. Conidiogenous
cells terminal or lateral, hyaline, smooth-walled, cylindrical
to slightly flask-shaped, 25–40 × 2–3 µm ( x̄ = 31 × 2.5 µm,
Fungal Diversity (2022) 117:1–272
129
Fig. 102 Scedosporium
dehoogii (AMH 9945, new
record) a Colonies on decaying woody stem of Suaeda
monoica. b − c Synnemata. d − f
Slimy head and conidiophores.
g − h Conidiogenous cells with
annellidic conidia. j Cylindrical and obovoid conidia. Scale
bars: b = 50 µm c = 100 µm,
d − j = 10 µm
n = 10). Conidiogenous cells arise from undifferentiated
hyphae, cylindrical to slightly flask-shaped, producing slimy
heads of one-celled, smooth-walled, sub-hyaline, obovoid or
sub-cylindrical conidia. Conidia: two types of conidia: (i)
those produced on synnemata and solitary conidiophores
were predominantly cylindrical or claviform, hyaline,
5–10 × 2–3 µm ( x̄ = 7.3 × 2.7 µm, n = 20) with a wide truncate base; (ii) those developed mainly from the undifferentiated hyphae of the substrate were sessile or on short protrusions, solitary, lateral, brown, smooth, and thick-walled,
mostly obovoid 2.5–7.5 × 2–3 µm ( x̄ = 4.3 × 2.4 µm, n = 20).
Sexual morph: Not observed.
Culture characteristics: Conidia germinating on Sea
Water agar within 24 h. Germ tubes produced from the
conidial base. Colonies on PDA fast growing, attaining
40–45 mm diameter after 14 days at 25 °C, circular, raised,
with white to grey at centre and white to cream at margins,
cottony; pale yellow in reverse with entire margin. The optimum growth temperature was from 25 to 37 °C. The fungus did not grow at 45 °C. Hyphae hyaline to pale brown,
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Fungal Diversity (2022) 117:1–272
Fig. 103 Scedosporium
dehoogii (NFCCI- 4274) a − b
Cultures on PDA after 14 days.
c − d Synnemata. e Slimy head
and Conidiophores. f sessile
obovoid conidia. g Conidiogenous cells with annellidic
conidia. h. sub cylindrical
conidia i − n Cylindrical and
obovoid conidia and Scale bars:
c = 50 µm d − n = 10 µm
branched and septate, 1–3 µm wide. Conidiophores synemmatous; synnemata solitary to gregarious, erect, dark brown,
180–255 µm tall with a cylindrical stipe from 10–30 µm
wide ( x̄ = 223 × 17 µm, n = 10), dark gray, smooth walled,
terminating into a slimy head of conidia, 45–85 µm long and
55–90 µm wide ( x̄ = 63 × 66 µm, n = 10). Hyphae interwoven
at the base, unbranched in the stipe, branching at the apex to
form conidiophores. Conidiophores solitary, branched often
reduced to conidiogenous cells which were sub-hyaline.
Conidiogenous cells terminal or lateral, hyaline smoothwalled, cylindrical to slightly flask-shaped, 30–45 × 2–3 µm
( x̄ = 35 × 2.4 µm, n = 10). Conidiogenous cells arising from
13
undifferentiated hyphae, cylindrical to slightly flask-shaped,
producing slimy heads of one-celled, smooth-walled, sub
hyaline, obovoid or sub-cylindrical conidia. Conidia: two
types: (i) those produced on synnemata and solitary conidiophores, predominantly cylindrical or claviform, hyaline,
5–15 × 2–5 µm ( x̄ = 10 × 3 µm, n = 20) with a wide truncate
base; (ii) those developed mainly from the undifferentiated
hyphae of the substrate were sessile or on short protrusions,
solitary, lateral, brown, smooth, and thick-walled, mostly
obovoid 2.5–9 × 2–5 µm ( x̄ = 5 × 3.5 µm, n = 20).
Material examined: India, Tamil Nadu, Tiruvarur,
Muthupet mangroves (10.4° N 79.5° E), on decaying wood
Fungal Diversity (2022) 117:1–272
Fig. 104 Phylogram based on the RAxML analysis of a combined
ITSrDNA and tub2 sequence dataset. Bootstrap support values for
ML and MP higher than 70% values are given above each branch
respectively. The new isolates are represented in blue. The tree is
rooted to Parascedosporium sanyaense EM65901. Thirty-eight
sequences are included in the phylogenetic analyses with 1159 char-
131
acters including gaps. The maximum parsimonious dataset consisted
of 855 characters were constant, 166 parsimony-informative and 138
parsimony-uninformative. The parsimony analysis of the data matrix
resulted in Two hundred and seventy equally parsimonious trees with
a length of 486 steps (CI = 0.759, RI = 0.888, RC = 0.674, HI = 0.241)
13
132
of Avicennia marina (Forssk.) Vierh. (Acanthaceae), 28
November 2015, B. Devadatha (AMH 9945), living culture,
NFCCI- 4274.
Hosts: Agricultural areas, playgrounds, riverbanks, Soil,
Human infections (Rougeron et al. 2018)
Distribution: Australia, Austria, Chile, India, Netherlands, Spain, Thailand (Rougeron et al. 2018)
GenBank numbers: MH569493 (ITS), MH569492 (LSU),
MH571777 (SSU)
Notes: Multigene phylogenetic anlayses of combined
datasets of ITS sequence data revealed that Scedosporium
dehoogii (NFCCI-4274) clustered together with the type
and other existing strains of S. dehoogii with moderate
statistical 74% ML and 78% MP support (Fig. 103). Scedosporium dehoogii (NFCCI-4274) from decaying wood
of Suaeda monoica and Avicennia marina and colonies on
PDA also share similar morphological characteristics with
S. dehoogii (CBS-117406) reported from the soil (Gilgado
et al. 2008). Scedosporium dehoogii is a common environmental species occurs on locations with high human activities like soil, water, agricultural areas and not involved in
human infections (Kaltseis et al. 2009). However, this is
the first report of Scedosporium dehoogii (NFCCI- 4274)
from marine habitats. Earlier Medicopsis romeroi has been
reported from mangroves (Devadatha et al. 2020). Also,
Calabon et al. (2018) reported S. aurantiacum from Sponges
in mangroves. With the present two species from mangroves
totally three Scedosporium spp. are recorded from marine
environments (Figs. 103, 104).
Hypocreomycetidae incertae sedis (Rhexoacrodictys and
Dematipyriforma clade)
Notes: In the current study, Rhexoacrodictys and Dematipyriforma form a distinct clade with a high statistical
support (100/100/100 for ML/MP/BYPP, respectively)
that is phylogenetically related to the orders Pleurotheciales and Savoryellales and might represent a new lineage
(Fig. 105). The phylogenetic placement of Rhexoacrodictys is controversial in various studies. Shi et al. (2021)
placed Rhexoacrodictys in Pleurotheciales, while Boonmee et al. (2021) placed the genus in Savoryellales. Sun
et al. (2017) placed Dematipyriforma in Savoryellales
where it forms a basal branch to representatives of the
order.
Dematipyriforma L. Y. Sun, Hai-Yan Li, Xiang Sun & L.D.
Guo.
Notes: Dematipyriforma is a monotypic genus typified by
D. aquilaria L. Y. Sun, Hai-Yan Li, Xiang Sun & L.D. Guo
that was isolated as an endophyte from a trunk of Aquilaria
crassna in Laos (Sun et al. 2017). The genus is characterized by monoblastic, integrated, intercalary or terminal, pale
brown to brown, determinate, cylindrical conidiogenous
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Fungal Diversity (2022) 117:1–272
cells; solitary, pyriform conidia with transverse and often
oblique or longitudinal, usually with a single small basal
cell; produce variously shaped Chlamydospores in culture
(Sun et al. 2017). During an ongoing study of freshwater
fungi from River Nile in Egypt (e.g. Abdel-Aziz 2016a,
b, c, 2020), three new species of Dematipyriforma were
recorded on submerged wood and date palm rachis in the
River Nile, Egypt that are described in this article based
on morphology and their phylogenetic placement. Species
of Dematipyriforma are phylogenetically related to Rhexoacrodictys species, both genera have rhexolytic conidial
secession, however the later genus has macronematous conidiophores and percurrently extending conidiogenous cells
vs. absent or micronematous conidiophores and determinate
conidiogenous cells in Dematipyriforma. Boonmee et al.
(2021) described Rhexoacrodictys nigrospora Boonmee,
D.F. Bao & K.D. Hyde from decaying wood in Thailand
with micronematous or semi-macronematous conidiophores
and determinate conidiogenous cells. They did not include
the sequences of Dematipyriforma aquilariain their phylogenetic analyses and in our opinion R. nigrospora can be
placed in Dematipyriforma based on morphology of the
conidiophores (micronematous vs. macronematous with
bulbous base) and conidiogenesis (determinate vs. percurrent proliferation in Rhexoacrodictys erecta (Ellis & Everh.)
W.A. Baker & Morgan-Jones) and the phylogenetic analyses
(Fig. 105).
Dematipyriforma aquatica Abdel-Aziz &Abdel-Wahab sp.
nov.
Index Fungorum number: IF900081; Facesoffungi number: FoF13400; Fig. 106
Etymology: Named after the aquatic habitat, where this
fungus was collected.
Holotype: SUMCC H-12001 (Sohag University Microbial
Culture Collection)
Saprobic on submerged wood in the River Nile. Sexual morph: Not observed. Asexual morph: Mycelium
immersed and superficial, sub-hyaline to brown, septate,
branched, smooth, 1.5–4 μm wide. Conidiophores absent or
present, when present micronematous, mononematous, subhyaline to brown, flexuous, smooth, unicellular, 7–15 μm
long, 1.5–4 μm wide. Conidiogenous cells holoblastic, integrated, intercalary or terminal, sub-hyaline to brown, determinate, ampulliform, clavate, subglobose, smooth, 5–8 μm
long, 5–6 μm wide. Conidial secession rhexolytic. Conidia
solitary or aggregated, effuse and heavily covered the surface of the wood, intercalary or terminal, smooth, pyriform
or subglobose, rounded at the apex, black, with brown to
dark-brown basal cell, muriform, 4–10 cells, 3–5 transverse
septa and 0–2 longitudinal septa, not or slightly constricted
at the septa, 27–38 × 15–26 μm ( x̄ = 32.2 × 20.9 μm, n = 50),
apex and basal cells are singles. Chlamydospores intercalary
Fungal Diversity (2022) 117:1–272
133
Fig. 105 Phylogenetic relationship of Dematipyriforma with related
taxa based on the nucleotide sequences of the combined SSU and
LSU rDNA. The maximum likelihood (ML) tree was constructed in
MEGA X (Kumar et al. 2018). The maximum parsimonious data set
of the combined genes consisted of 30 taxa with three representa-
tives of Hypocreales used as outgroup. Phylogenetic trees obtained
from ML, MP and BYPP were similar in topology. Bootstrap support
on the nodes represents ML and MP ≥ 50%. Branches with a BYPP
of ≥ 95% are in bold. The three new Dematipyriforma species are in
blue
or terminal, catenated, straight, or curved, brown to dark
brown, smooth, granulate, phragmoseptate.
Culture characteristics: Colonies on PDA reaching
25 mm diam after 3 weeks, at 25 °C, dark-brown to black,
reverse dark-brown to black. Conidial dimensions and
shapes are similar to those found on natural wood.
Material examined: Egypt, Sohag City, the River Nile,
on submerged wood, 14 August 2012, F. A. Abdel-Aziz,
SUMCC H-12001, holotype, ex-type living culture,
SUMCC 12101.
GenBank numbers: MT522154 (SSU), MT522155 (LSU)
Notes: Combined phylogenetic analyses of SSU and LSU
rDNA placed the three new species of Dematipyriforma with
the type species D. aquilaria with high statistical support as
distinct new taxa. The three new species differ from D. aquilaria in having black conidia vs. pale grey olivaceous to pale
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Fig. 106 Dematipyriforma
aquatica (SUMCC H-12001,
holotype).a–j, l–pVariously
shaped conidia at different
stages of maturity. k Chlamydospores in culture. a–j Conidia
from natural wood. g–j, l–p
Conidia from pure culture.
Scale bars: a–p = 10 µm
brown in the latter species. Dematipyriforma aquilaria was
isolated as an endophyte from a trunk of Aquilaria crassna
in Laos (Sun et al. 2017), while the three new species are
freshwater taxa. Conidia of D. aquilaria are evenly pigmented, while the three new species have black conidia with
basal cells that are lighter. Chlamydospores of D. aquilaria
have thick walls with axial perforative canals, these canals
are absent in the three new species. A comparison of the 590
nucleotides of the D1/D2 region of the LSU rDNA of the
three new species of Dematipyriforma with D. aquilarias
hows 14 base pair differences (2.37%) which justifies the
erection of the three new species following the guidelines
of Jeewon and Hyde (2016).
Dematipyriforma globispora Abdel-Aziz &Abdel-Wahab
sp. nov.
Index Fungorum number: IF900083; Facesoffungi number: FoF13865; Fig. 107
Etymology: Named after the shape of the globose shape
of the conidia.
Holotype: SUMCC H-12002
Saprobic on submerged wood in the River Nile. Sexual morph: Not observed. Asexual morph: Mycelium
immersed and superficial, sub-hyaline to brown, septate,
13
branched, smooth, 2–4 μm wide. Conidiophores absent or
present, when present micronematous, mononematous, subhyaline to brown, flexuous, smooth, unicellular, 6–12.5 μm
long, 1.5–4.5 μm wide. Conidiogenous cells holoblastic,
integrated, intercalary or terminal, sub-hyaline to brown,
determinate, clavate, smooth, 4–6.5 μm long, 2.5–4.5 μm
wide. Conidial secession rhexolytic. Conidia solitary or
aggregated, effuse and heavily covered the surface of the
wood, intercalary or terminal, smooth, globose or subglobose, rounded at the apex, black, with brown to dark-brown
basal cell, muriform, with irregular transverse, longitudinal
and oblique septa and form mass of cells, not constricted
at the septa, sometimes surrounded by network of hyphae,
17–37 × 15–30 μm ( x̄ = 27.7 × 20.9 μm, n = 60), smaller buds
are produced from conidia that are yellow–brown to brown
in color, muriform, 11–19.2 × 7.1–14.5 μm. Chlamydospores
intercalary or terminal, catenated, straight, or curved, brown
to black, smooth, form large, black muriform masses similar
to conidia but much larger in size 37–120 × 17–30 μm.
Culture characteristics: Colonies on PDA reaching
20 mm diam after 3 weeks, at 25 °C, dark-brown to black,
reverse dark-brown to black. Conidial dimensions and
shapes are similar to those found on natural wood.
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135
Fig. 107 Dematipyriforma
globispora (SUMCC H-12002,
holotype).a–g, i–lVariously
shaped conidia at different
stages of maturity. h Chlamydospores in culture. a–e Conidia
from natural wood. f–g, i–l
Conidia from pure culture.
c–d Conidia are surrounded by
network of hyphae. e, l Conidia
with small buds. Scale bars:
a–l = 10 µm
Material examined: Egypt, Sohag City, the River Nile,
on submerged wood, 14 August 2012, F. A. Abdel-Aziz,
SUMCC H-12002, holotype, ex-type living culture,
SUMCC 12102.
GenBank numbers: MT522158 (SSU), MT522153 (LSU).
Notes: Dematipyriforma globispora differs from the other
three species in having smaller conidia, that are mostly globose, sometimes surrounded by networks of hyphae and produce buds that are pale-brown to brown. Dematipyriforma
globispora and D. aquatica are phylogenetically related,
however, they differ in their morphology. Conidial cells in
D. globispora are arranged irregularly and conidia are globose with small buds and surrounded by network of hyphae.
Conidia in D. aquatica are pyriform or subglobose with 3 to
5 continuous transverse septa. Rhexoacrodictys nigrospora
produce similar conidia with overlapping dimensions, however, conidia of D. globispora are surrounded by network
of hyphae with brown buds. Mycelium in R. nigrospora are
narrow (1–2 μm wide) with verruculose or finely echinulatewalled (Boonmee et al. 2021), while hyphae in D. globispora are 2–4 μm wide and smooth. A comparison of the
805 nucleotides of the LSU rDNA for D. globispora and
R. nigrosporashows 11 base pair differences (1.36%) that
confirm they are two different species.
Dematipyriforma nilotica Abdel-Aziz &Abdel-Wahab, sp.
nov.
Index Fungorum number: IF900084; Facesoffungi number: FoF13401; Fig. 108
Etymology: Named after the River Nile where the fungus
was collected.
Holotype: SUMCC H-12003
Saprobic on submerged date palm rachis in the River Nile.
Sexual morph: Not observed. Asexual morph: Mycelium
immersed and superficial, sub-hyaline to brown, septate,
branched, smooth, 1.5–3.5 μm wide. Conidiophores absent
or present, when present micronematous, mononematous,
sub-hyaline to reddish-brown, flexuous, smooth, unicellular,
7–13 μm long, 2.5–4.5 μm wide. Conidiogenous cells holoblastic, integrated, intercalary or terminal, sub-hyaline to
brown, determinate, ampulliform, clavate, subglobose, cylindrical, smooth, 1.5–8.5 μm long, 4.5–6.5 μm wide. Conidial
secession rhexolytic. Conidia solitary or aggregated, effuse
and heavily covered the surface of the wood, intercalary or
terminal, smooth, or surrounded by fine fibers, pyriform,
globose or subglobose, rounded at the apex, conidial cells
are filled gradually with black material, black when mature,
with brown to dark-brown basal cell, muriform, 6–9 cells,
3–5 transverse septa and 0–2 longitudinal or oblique septa,
not or slightly constricted at the septa, 31–45 × 21–37 μm
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Fig. 108 Dematipyriforma
nilotica (SUMCC H-12003,
holotype). a–l Variously shaped
conidia at different stages of
maturity. a–d Conidia from
natural date palm rachis. e–l
Conidia from pure culture. m
Chlamydospores in culture. b,d
Conidia surrounded by fine fibers. a–b, g–i Gradual filling of
the conidial rows by black material. Scale bars: a–m = 10 µm
( x̄ = 37 × 26.3 μm, n = 50), basal cells are singles. Chlamydospores intercalary or terminal, dark-brown to black, smooth,
form large, black muriform masses with irregular shapes.
Culture characteristics: Colonies on PDA reaching
30 mm diam after 3 weeks, at 25 °C, brown to reddish darkbrown, reverse brown to dark-brown. Conidial dimensions
and shapes are similar to those found on natural wood.
Material examined: Egypt, Sohag City, the River Nile, on
submerged date palm rachis, 14 August 2012, F. A. AbdelAziz, SUMCC H-12003, holotype, ex-type living culture,
SUMCC 12103.
GenBank numbers: MT522157 (SSU), MT522156 (LSU).
Notes: Dematipyriforma nilotica differs from the other
three species in having larger conidial dimensions that sometimes surrounded by fine fibres. Dematipyriforma nilotica is
phylogenetically related with D. aquilaria, however, the first
species have black and larger conidia (31–45 × 21–37 μm vs.
25–37 × 15–22 μm in D. nilotica and D. aquilaria respectively) and grow on decaying date palm rachis in freshwater habitat, while the latter species produce brown conidia
and live as an endophyte. Chlamdydospores in D. aquilaria
have perforative canal that is absent in D. nilotica. Both
13
D. aquatica and D. nilotica have 3–5 continuous transverse
septa with 0–2 longitudinal septa, however, conidia in the
latter species are surrounded by fine fibres and are larger
in size (27–38 × 15–26 μm vs. 31–45 × 21–37 μm for D.
aquatica and D. nilotica respectively). A comparison of the
800 nucleotides of the LSU rDNA for D. aquatica and D.
niloticashows 15 base pair differences (1.87%) that confirm
they are two different species.
Key to Dematipyriforma species
1. Endophytic species, produce brown conidia………
……………………………………………… D. aquilaria
1*.Saprobic species in freshwater habitat, produce black
conidia…………………………… ………………………2
2. Conidial cells arranged in rows with continuous transverse septa………………………………………………… 3
2*.Conidial cells arranged irregularly, globose
conidia with small buds and surrounded by network of
hyphae………………………………………D. globispora.
3. Conidial width less than 30 µm; smooth conidial
wall……………………………………………D. aquatica
3*.Conidial width more than 30 µm; conidial wall with
fine fibres ……………………………………….D. nilotica.
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137
Fig. 109 Coniochaeta caraganae (MFLU 17–2500, holotype). a, b. Appearance of conidiomata on host surface. c. Vertical section through the
conidioma. d. Peridium. e, f. Conidiogenesis. g–k. Conidia. Scale bars: a, b = 500 µm, c, d = 30 µm, e–k = 10 µm
Subclass Savoryellomycetidae Hongsanan, K.D. Hyde &
Maharachch.
Coniochaetales Huhndorf, A.N. Mill. & F.A. Fernández,
Mycologia 96(2): 378 (2004).
Coniochaetales was introduced by Huhndorf et al. (2004)
with Coniochaetaceae as the type family. Coniochaetales
comprises two families namely Coniochaetaceae and Cordanaceae. Cordanaceae was previously accommodated in
Cordanales. Hongsanan et al. (2017) treated Cordanales
as a synonym of Coniochaetales based on molecular clock
evidence. Hyde et al. (2020a, b, c) suggested the divergence
time for Coniochaetales to be around 131 MYA. Coniochaetales comprises two families and five genera (Hyde
et al. 2020a, b, c).
Coniochaetaceae Malloch and Cain (1971).
Coniochaetaceae was introduced by Malloch and Cain
(1971) to accommodate two genera Coniochaeta and
Coniochaetidium (Sacc.). Coniochaetaceae is typified by
Coniochaeta and accommodates two genera Barrina A.W.
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138
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Fungal Diversity (2022) 117:1–272
◂ Fig. 110 Phylogenetic tree generated from Maximum likelihood
analysis (RAxML) based on combined ITS and LSU sequence data
of Coniochaetaceae in the order Coniochaetales. Maximum likelihood bootstrap support values ≥ 70% (in blue) and Bayesian posterior probabilities ≥ 0.95 (in green) are indicated on the branches. The
new isolate is in blue. The tree is rooted with Chaetosphaeria jonesii
(MFLUCC 15–1015) and Chaetosphaeria garethjonesii (MFLUCC
15–1012)
Ramaley and Coniochaeta (Sacc.) Cooke (Wijayawardene
et al. 2020) based on multi gene phylogeny (Maharachchikumbura et al. 2015a, b; Samarakoon et al. 2018). Species
belonging to this family are saprobes and pathogenic on
various decaying wood and plants or animals, respectively
(Khan et al. 2013; Maharachchikumbura et al. 2015a, b,
2016; Wijayawardene et al. 2017a, b; Samarakoon et al.
2018).
Coniochaeta (Sacc.) Cooke, Grevillea 16(no. 77): 16
(1887)
Coniochaeta was introduced by (Sacc.) Cooke (1887a,
b) and is typified by Coniochaeta ligniaria (Grev.) Cooke.
Coniochaeta is characterized by solitary or aggregated,
typically setose, dark brown to black, pyriform to globose
ascomata, membranaceous to pseudoparenchymatous or
coriaceous peridium, paraphysate hamathecium, unitunicate
and thin-walled asci with a small non-amyloid apical ring
and one-celled, usually dark brown and often laterally compressed with a germ slit ascospores (Greville 1823–1824,
Cooke 1887a, b). The hyphomycetous asexual morph is
characterized by phialidic conidiogenous cells, previously
described in Lecythophora (Weber 2002; Khan et al. 2013).
In this study, we introduce a new species C. caraganae collected on dead twigs of Caragana frutex in Russia based on
morphology and phylogenetic evidence.
Coniochaeta caraganae D. Pem, Bulgakov & K.D. Hyde,
sp. nov.
Index Fungorum number: IF559528, Facesoffungi number: FoF08686; Fig. 109
Etymology: ″caraganae″ refers to the host plant from
which the fungus was isolated.
Holotype: MFLU 17-2500
Saprobic on dead branch of Caragana frutex. Sexual morph: Not observed. Asexual morph: Coelomycetous. Conidiomata 106–114 μm diam., 4–7 μm high
(x̄ = 110.7 × 6.2 μm, n = 20), small, pycnidial, solitary,
scattered, immersed, uniloculate, globose to subglobose,
thin-walled. Peridium 26–32 μm composed of two layers.
Inner layer consisting cells of textura prismatica, hyaline
to subhyaline, strongly compressed. Outer layer consisting
of densely packed, moderately thick-walled, brown cells of
textura angularis, tending to be darker and more isodiametric towards the outside. Conidiophores 7–10 × 2–3 μm
(x̄ = 6.5 × 2.3 μm, n = 20), hyaline, straight or irregularly
139
bent, reduced to conidiogeneous cells. Conidiogenous cells
2–3 × 4–6 μm (x̄ = 3.1 × 5.1 μm, n = 20), holoblastic, annellidic, simple, determinate, hyaline, doliiform to cylindrical,
smooth-walled. Conidia 3–4 × 5–6 μm (x̄ = 4.1 × 5.8 μm,
n = 20) oblong to ovoid, straight, rounded at both ends,
sometimes truncate at base, cylindrical, aseptate, smooth
and thick-walled, eguttulate.
Culture characteristics: colonies on MEA, reaching
25–35 mm diam. after 4 weeks at 25 °C, grey whitish, dense,
effuse, with white hyphal stands towards the edge, rough
surface towards centre, diffuse margin; reverse dark grey
with whitish edges, grayish orange at the center, radiating,
effuse and zonate.
Material examined: Russia, Donetsk People's Republic,
Shakhtersk district, regional landscape park «Donetsk ridge»
(Rus. «Donetsky kryazh»), steppe near Leontievsky forest,
on dead twigs of Caragana frutex (L.) K. Koch (Fabaceae),
19 May 2017, Timur S. Bulgakov (MFLU 17-2500, holotype); ex-type living culture MFLUCC 18-0780.
GenBank numbers: MT573224 (ITS), MT573223 (LSU).
Notes: Based on phylogenetic analyses, the new asexual
species Coniochaeta caraganae is closely related with C.
coluteae (Samarakoon et al. 2018; Fig. 110). The asexual
morph of C. coluteae is hyphomycetous and has been
obtained from culture. It is characterized by hyaline vegetative hyphae, hyphoide conidiophores, phialidic conidiogenous cells and one celled hyaline conidia. Our new
species C. caraganae has been obtained from dead branch
of Caragana frutex and is characterized by pycnidial conidiomata, hyaline conidiophores and oblong to ovoid conidia.
In a comparison of the 565 ITS (+ 5.8S) nucleotides of
these two strains C. coluteae MFLUCC 17-2299 to that of
C. caraganae reveals 11 (1.9%) nucleotide differences which
justifies these two isolates as two distinct taxa (Jeewon and
Hyde 2016). Therefore, C. caraganae is introduced herein
as a new species.
Pleurotheciales Réblová & Seifert
Notes: Pleurotheciales was introduced by Réblová et al.
(2016) with Pleurotheciaceae as the type family. Wijayawardene et al. (2020, 2022) listed Pleurotheciales as a
monotypic order and new taxa have been included in it
(Boonmee et al. 2021; Crous et al. 2021; Dong et al. 2021).
Pleurotheciaceae Réblová & Seifert, in Réblová, Seifert,
Fournier & Štěpánek, Persoonia 37: 63 (2015) [2016]
Notes: Wijayawardene et al. (2022) listed 14 genera
in Pleurotheciaceae. Recently, Saprodesmium was introduced in this family for a saprobic fungus found on decaying wood submerged in freshwater in China (Dong et al.
2021). Pleurotheciaceae was introduced by Réblová et al.
(2016) with Pleurothecium recurvatum as the type species.
Fourteen genera viz. Adelosphaeria, Anapleurothecium,
13
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Fungal Diversity (2022) 117:1–272
Fig. 111 Rhexoacrodictys
erecta (MFLU 21–0277, new
record). a, b Colonies on wood.
c–f Conidiophore with conidia.
g–i Conidia. j Germinating
conidia. k, l Culture on PDA
from surface and reverse. Scale
bars: c–f, j = 20 μm, g–i = 10 μm
Coleodictyospora, Dematipyriforma, Helicoascotaiwania,
Melanotrigonum, Monotosporella, Neomonodictys, Phaeoisaria, Phragmocephala, Pleurotheciella, Pleurothecium,
Saprodesmium, and Sterigmatobotrys are accepted in the
family (Hyde et al. 2020a, b, c; Boonmee et al. 2021; Dong
et al. 2021; Bao et al. 2022). Species in Pleurotheciaceae
are cosmopolitan with a wide range of hosts and substrates
from terrestrial and freshwater habitats (Hyde et al. 2020a,
b, c; Dong et al. 2021; Boonmee et al. 2021; Bao et al.
2022).
The sexual morphs of Pleurotheciaceae share perithecial, immersed to superficial, papillate ascomata, leathery
to fragile, carbonaceous peridial walls, unitunicate, cylindrical, 8-spored, asci with a distinct non-amyloid apical
annulus, abundant paraphyses and ellipsoidal to fusiform,
septate, hyaline or versicolorous ascospores (Réblová et al.
2016; Luo et al. 2018a, b; Hyde et al. 2020a, b, c). The
asexual morphs of Pleurotheciaceae have been reported as
hyphomycetes forming indeterminate synnemata or loose
fascicles. Conidiophores are macronematous or semimacronematous. Conidiogenous cells produce holoblastic conidia, with rhexolytic conidial secession on short
13
denticles or extending polyblastically on a sympodial
rachis. Conidia are hyaline to brown, varied in shape, septate or aseptate (Hyde et al. 2020a, b, c, Dong et al. 2021,
Boonmee et al. 2021, Bao et al. 2022).
Rhexoacrodictys W.A. Baker & Morgan-Jones, Mycotaxon
82: 98 (2002)
Rhexoacrodictys was introduced by Baker et al. (2002) to
accommodate species previously identified as Acorcdictys
(i.e., A. erecta, A. fimicola, A. fuliginosa and A. queenslandica) and wherein Rhexoacrodictys erecta was designated as the type. Two additional species R. martini and R.
broussonetiae were subsequently added to the genus based
on morphological characteristics (Delgado 2009; Xiao et al.
2018). While R. martini and R. queenslandica were transferred to Distoseptispora and Junewangia based on phylogenetic analysis (Xia et al. 2017). Currently, four species are
accepted in Rhexoacrodictys (R. broussonetiae, R. erecta, R.
fimicola, and R. fuliginosa).
Rhexoacrodictys erecta (Ellis & Everh.) W.A. Baker &
Morgan-Jones, in Baker, Partridge & Morgan-Jones, Mycotaxon 82: 99 (2002)
Fungal Diversity (2022) 117:1–272
Refer to Species Fungorum (2022a, b) for synonyms.
Index Fungorum number: IF381123; Facesoffungi number: FoF13392; Fig. 111
Saprobic on dead culms of bamboo. Sexual morph:
Not observed. Asexual morph: Colonies on the substratum superficial, hairy, effuse, blackish, shining.
Mycelium mostly immersed, cylindrical, brown to
dark brown hyphae. Conidiophores 28–57 × 4–5 μm
( x̄ = 42.5 × 4.5 μm, n = 15), macronematous, mononematous, erect, single, straight or somewhat flexuous,
cylindrical, smooth-walled, brown to dark brown, septate. Conidiogenous cells monoblastic, pale brown to
brown, integrated, terminal. Conidia 19–27.5 × 15–19 μm
( x̄ = 23.5 × 17 μm, n = 30), holoblastic, solitary, dry, broad
oval to subglobose, muriform, acrogenous, transversely
and longitudinally septate, dark brown to black, smoothwalled, narrowly truncate at the base.
Culture characteristics: Colonies on PDA, 30 mm diam
after two weeks at 28 °C, brown to blackish at the front and
reverse sides, mycelium sparse; reverse blackish.
Material examined: Thailand, Chiang Rai Province,
Muang District, on dead culms of Bambusa sp. (Poaceae),
11 November 2020, X. G. Tian U–2–3 (MFLU 21–0277),
living culture, MFLUCC 21–0157.
Habitat: on submerged wood, Arundo donax, Bambusa
multiplex, Bambusa sp., Fagus crenata, palm tree, Saccharum officinarum, Sorghum bicolor, Sporoschisma saccardoi
and Zea mays (Ellis 1961; Baker et al. 2002; Zhao et al.
2011; Xia et al. 2017; Shi et al. 2021).
Distribution: Known from China, India, Japan, Sierra
Leone, South Africa, Thailand, USA and Venezuela (Ellis
1961; Baker et al. 2002; Zhao et al. 2011; Xia et al. 2017;
Shi et al. 2021; This study).
GenBank numbers: OL606411 (ITS), OL606151 (LSU),
OL606015 (SSU).
Notes: This species was reported as Acrodictys erecta
on Arundo donax in Venezuela and on Zea mays in USA
by Ellis (1961). Baker et al. (2002) examined several type
specimens of synonyms of this species and erected Rhexoacrodictys based on morphological analysis. Our new isolate
(MFLUCC 21-0157) clustered with the strains of Rhexoacrodictys erecta (HMAS 245615, IFRD500–016 and HMAS
245616; Fig. 112). Morphologically, our new collection is
similar to those of Rhexoacrodictys erecta (Ellis 1961; Baker
et al. 2002).
Based on nucleotide comparisons of ITS, LSU and SSU,
our new strain (MFLUCC 21–0157) and three of Rhexoacrodictys erecta (HMAS 245615, IFRD500–016 and HMAS
245616) show no differences; however, the new strain
(MFLUCC 21–0157) is different from Monotosporella seteosa (HKUCC 3712) in 14/516 bp (2.71%) of the LSU (data
contains 6 gaps). Based on both phylogeny and morphology
141
showed that our strain (MFLUCC 21-0157) is identical to
Rhexoacrodictys erecta.
Phaeoisaria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 330 (1909).
Notes: Phaeoisaria is a genus of hyphomycetes (von
Höhnel 1909) having morphological features mainly characterized by ‘indeterminate synnemata with aseptate or septate ellipsoidal, obovoidal, fusiform-cylindrical to falcate,
hyaline conidia’ (Boonmee et al. 2021). We follow Crous
et al. (2021), Boonmee et al. (2021) and Wijayawardene
et al. (2022) for the classification and updated account of
Phaeoisaria species.
Phaeoisaria goiasensis H.M. Silva, A.D. Cavalcanti &
J.D.P. Bezerra, sp. nov.
MycoBank number: MB840294, Facesoffungi number:
FoF09975, Fig. 113
Etymology: The name refers to the Brazilian state, Goiás,
where the fungus was isolated.
Holotypus: UFG 71083.
Isolated from a Petri-dish with culture medium storage
in a fridge. Asexual morph: Hyphae hyaline to pale brown
with age, smooth wall, septate, 1.5–2.5(–3) µm wide. Synnemata erect, brown, smooth, indeterminate, composed of
compactly and parallels conidiophores and commonly with
conidiogenous cells in the above half, 93–147 × 3.5–4.5 µm.
Conidiophores straight or slightly curved, septate, reduced
to conidiogenous cell, cylindrical, hyaline to pale brown,
smooth wall, (10–)15–49(–72) × (1.5–)2–2.5(–3.5) µm. Conidiogenous cells polyblastic, integrated, terminal or intercalary, cylindrical, hyaline, smooth wall, (3–)3.5–9.5(–15.5)
× (1.5–)2–3(–3.5) µm, forming conidia on denticles, 1–2 µm
long, 0.5–1 µm wide, scattered or clustered in the apical
region. Conidia ellipsoidal to obovoid, straight or slightly
curved, rounded at the ends or occasionally tapering toward
the base, hyaline, aseptate, guttulate, smooth wall, (4.5–)
7.5–9(–10.5) × (2–)2.5–3(–4) µm. Chlamydospores terminal, globose, pyriform, first hyaline and becoming brown to
dark brown with age, (8–)8.5–10.5(–17) × (2–)7–8(–8.5) µm.
Sexual morph: Not observed.
Culture Charactersistics: Colonies grew in the dark for
7 days at 25 °C. On PDA, colonies elevated, aerial mycelium
absent, irregular, greyish to dark grey with edge whitish,
up to 15 mm. On MEA, colonies plane, aerial mycelium
absent, greyish to dark grey with edge brown to dark brown,
up to 10 mm. On oatmeal agar (OA), colonies growing up to
18 mm. At 36 °C, colonies are similar to at 25 °C, growing
up to 8 mm on PDA, 9 mm on MEA, and 6 mm on AO. No
growth at 10 °C.
Material examined: Brazil, Goiás state, Goiânia City,
Universidade Federal de Goiás (UFG), Instituto de Patologia
Tropical e Saúde Pública (IPTSP), Laboratório de Micologia
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Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
◂ Fig. 112 Combined phylogeny using ITS, LSU SSU, rpb2 and tef1 of
selected members of four orders of the Hypocreomycetidae. The dataset of combined ITS, LSU, SSU, rpb2 and tef1 sequence data comprise 79 strains with 4413 characters including gaps (rpb2: 1–1051
bp, tef1: 1052–2046 bp, ITS: 2047–2623 bp, LSU: 2624–3413 bp,
SSU: 3414–4413 bp). Leotia lubrica (AFTOL-ID1) and Microglossum rufum (AFTOL-ID 1292) were used as outgroup taxa. RAxML
and Bayesian analyses were conducted and resulted in generally congruent topologies. Bootstrap support values for ML ≥75% and BYPP
≥0.95 are given as ML/BYPPP above the nodes. Newly obtained
sequences are indicated in red and ex-type strains are in bold
(LabMicol), isolated from a Petri dish with culture medium
storage in a fridge, 19 November 2019, J.D.P. Bezerra &
H.M. Silva (UFG 71083, holotype); ex-type living culture
URM 8387 = FCCUFG 02; ibid. living culture FCCUFG 03.
GenBank numbers: URM 8387 = FCCUFG 02
–MT210320 (ITS), MT375865 (LSU), MT384422 (tub2),
MT384424 (tef1)
FCCUFG 03 – MT210321 (ITS), MT375866 (LSU),
MT384423 (tub2), MT384425 (tef1)
Notes: Phaeoisaria was described by von Höhnel (1909)
and has 32 records in the Index Fungorum and MycoBank
databases (8 June 2022). Based on our phylogenetic analysis
(Fig. 114), our isolates here are proposed for the new species P. goiasensis. Phaeoisaria goiasensis differs from P.
annesophieae, which was isolated from soil in The Netherlands, by the presence of synnemata in old cultures (after
30 days) and defined conidiophores in P. goiasensis, and by
the size of conidiogenous cells (12–39 × 1–3.5 µm), conidia
(4.5–9 × 2–3.5 µm), and chlamydospores (9–18 × 7–9.5 µm) of
P. annesophieae (Crous et al. 2017). The BLASTn searches (8
June 2022) using ITS sequences of P. goiasensis demonstrated
that they are identical to sequences deposited as Phaeoisaria
sp. INBio 4514E, which was isolated from substrate related
to Passalidae galleries in decayed trunks (Vargas-Asensio
et al. 2014), and to sequences obtained from submerged wood
and deposited as Ascomycota (Brown et al. 2016). The ITS
sequences from P. goiasensis also had highest similarity to P.
annesophieae (strain CBS 143235, GenBank MG022180.1;
Identities = 500/511 (98%), 0 gap (0%)). The LSU sequences
had high identity to Phaeoisaria sp. INBio 4514E and P.
annesophieae MFLU 19–0531, amongst other sequences
deposited as Phaeoisaria species/isolates. The tef1 sequences
had high similarity to P. filiformis MFLU 18-1462 and it was
also 96.59% identical to P. sedimenticola S-908. The tub2
sequences had low identity to Sordariomycetes species.
Pleurothecium Höhn., Ber. dt. bot. Ges. 37: 154 (1919).
Notes: Pleurothecium was introduced by Höhnel (1919)
with Pleurothecium recurvatum (Morgan) Höhn as the type
species. There are 11 species listed in Index Fungorum
(June, 2022), and a new species, P. aquisubtropicum, is
described and illustrated here (Fig. 5).
143
Pleurothecium aquisubtropicum J. Ma, Y.Z. Lu & K.D.
Hyde, sp. nov.
Index Fungorum number: IF559508, Facesoffungi number: FoF08709; Fig. 115
Etymology: Referring to the aquatic habitat and collecting
site in subtropical country, China.
Holotype: GZAAS 21–0384
Saprobic on decaying wood in a freshwater stream.
Asexual morph: Colonies on natural substrate superficial,
effuse, brown or dark brown, smooth. Mycelium immersed
or superficial, smooth. Conidiophores 82–177 × 3.5–5.5 μm
( x̄ = 121.3 × 4.3 μm, n = 20), macronematous, mononematous, sucylindrical, straight, unbranched, smooth, septate, brown, paler towards the apex. Conidiogenous cells
25–38 × 2.7–3.8 μm ( x̄ = 29.6 × 3.2 μm, n = 15), holoblastic, polyblastic, integrated, terminal, subhyaline to pale
brown, subcylindrical. Conidia 13.4–15.5 × 3.4–5.5 μm
( x̄ = 14.4 × 4.4 μm, n = 21), acrogenous, solitary, aseptate, pale brown, straight, guttulate, hyaline or pale green,
smooth. Sexual morph: Not observed.
Culture Characters: Colonies growing slowly on PDA,
reaching 37 mm in 35 days at 25 °C, flat, filiform, round,
gray or white, smooth; In reverse, milky at the center, brown
or dark brown at the margin.
Material examined: China, Guizhou Province, Xishui
County, on decaying wood submerged in a freshwater
stream, 13 February 2021, Jian Ma, TL2(GZAAS 21–0384,
holotype); ex–type living culture, GZCC 21–0670.
GenBank numbers: OM339436 (ITS), OM339433 (LSU)
Notes: Our new collection fits well with the generic
concept of Pleurothecium in having macronematous,
mononematous, brown conidiophores, polyblastic, integrated conidiogenous cells and septate, smooth allantoid
or fusiformis conidia. In our phylogenetic analyses, our
new collection of Pleurothecium aquisubtropicum was
placed within Pleurothecium and is basal to other Pleurothecium species (Fig. 116). Pleurothecium aquisubtropicum resembles P. aquaticum in the shape of the conidiophores, and conidia (Luo et al. 2018a, b). However,
Pleurothecium aquisubtropicum differs from P. aquaticum by its darker and longer conidiophores (82–177 μm
vs 53–65 μm) and smaller conidia (13.5–15.5 × 3–5.5 μm
vs 19–21 × 4.5–5.5 μm). Hence, based on both morphology and phylogeny, we introduce our collection as a new
species of Pleurothecium aquisubtropicum.
Subclass Xylariomycetidae O.E. Erikss & Winka.
Amphisphaeriales D Hawksw & OE Erikss.
Amphisphaeriales was introduced by Eriksson and
Hawksworth (1986). However, Amphisphaeriales was synonymized with Xylariales by Eriksson and Hawksworth
(1987). Based on morphological and molecular data,
these two orders were separated and Amphisphaeriales
13
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Fungal Diversity (2022) 117:1–272
Fig. 113 Phaeoisaria goiasensis (UFG 71083, holotype)
a–c Synnemata d Details of a
synnema e–j Conidiophores and
conidia k Conidia l–o Chlamydospores and conidia. Scale
bars: 10 μm
was resurrected (Senanayake et al. 2015). The evidence
for the continuation of Amphisphaeriales and Xylariales
as distinct orders in Xylariomycetidae is also provided
in Maharachchikumbura et al. (2016), Samarakoon et al.
13
(2016a, b), Hongsanan et al. (2017) and Daranagama et al.
(2018). Amphisphaeriales comprises with 17 families viz
Amphisphaeriaceae, Apiosporaceae, Beltraniaceae, Castanediellaceae, Clypeophysalosporaceae, Cylindriaceae,
Fungal Diversity (2022) 117:1–272
Fig. 114 Phylogram generated from Bayesian analysis based on combined ITS and LSU rDNA sequence data of Phaeoisaria conducted
in MrBayes on XSEDE in the CIPRES science gateway. Twenty-nine
strains/isolates are included in the combined analysis, which comprised a total of 1435 characters (ITS = 591 and LSU = 844), including gaps. The substitution model GTR + I + G was used for ITS and
LSU alignments. The tree is rooted with Pleurothecium semifecun-
145
dum CBS 131271 and Pleurothecium recurvatum CBS 138747 and
the scale bar indicates the number of changes. The ex-types (reference strains) are in bold and the new isolates are in blue. Maximum
likelihood bootstrap (ML-BS) support values obtained with RAxML
using 1000 replicates greater than 70% and Bayesian posterior probabilities (BYPP) greater than 0.95 are indicated near nodes
13
146
Fungal Diversity (2022) 117:1–272
Fig. 115 Pleurothecium
aquisubtropicum (GZAAS
21–0384, holotype) a Colony
on decaying wood b–c
Conidiophores with attached
conidia d–g Conidiogenous
cells and conidia h–j Conidia
k Germinating condium l–m
Colony on PDA from above and
below. Scale bars: b, c = 20 μm,
d–k = 10 μm
Hyponectriaceae, Iodosphaeriaceae, Melogrammataceae,
Oxydothidaceae, Phlogicylindriaceae, Pseudomassariaceae, Pseudosporidesmiaceae, Pseudotruncatellaceae,
Sporocadaceae, Vialaeaceae, and Xyladictyochaetaceae
(Hyde et al. 2020a, b, c; Wijayawardena et al. 2020).
Pestalotiopsis, Pseudopestalotiopsis, Pseudosarcostroma,
Robillarda, Sarcostroma, Seimatosporium, Seiridium, Sporocadus, Strickeria, Synnemapestaloides, Truncatella,
Xenoseimatosporium (Hyde et al. 2020a, b, c; Tibproma
et al. 2020; Wijayawardene et al. 2020).
Sporocadaceae Corda, [as 'Sporocadeae'], Icon. fung.
(Prague) 5: 34 (1842).
Sporpcadaceae was first established by Corda (1842)
and typified by Sporodocadus Corda (1839). The members of Sporocadaceae can be found as saprobes, pathogens, and endophytes in a variety of habitats (Nag Raj,
2993, Tanaka et al. 2011, Hyde et al. 2020a, b, c). The
family contains 32 genera viz Allelochaeta, Annellolacinia, Bartalinia, Broomella, Ciliochorella, Diploceras,
Disaeta, Discosia, Distononappendiculata, Diversimediispora, Doliomyces, Heterotruncatella, Hyalotiella,
Hymenopleella, Immersidiscosia, Monochaetia, Morinia,
Neopestalotiopsis, Nonappendiculata, Parabartalinia,
Bartalinia Tassi, Bulletin Labor. Orto Bot. de R. Univ. Siena
3: 4 (1900)
Bartalinia was introduced by Tassi (1900) and belongs
to Sporocadaceae, (Hyde et al. 2020a, b, c; Wijayawardene
et al. 2020). Bartalinia is morphologically similar to Heterotruncatella, Hymenopleella, Morinia, Parabartalinia,
Pestalotiopsis, Pseudosarcostroma, and Truncatella. The
characteristic features of Bartalinia are pycnidial conidiomata and fusiform, 3–4-septate, appendage bearing
conidia with an acute or blunt apex (Senanayake et al.
2015). There are 22 Bartalinia species in the Species Fungorum (2022a, b) and a new species B. kevinhydei was
introduced by Tibpromma et al. (2021). The generic and
species boundaries of Bartalinia-like taxa are complicated
13
Fungal Diversity (2022) 117:1–272
147
Fig. 116 Phylogram generated from maximum likelihood
analysis based on combined
LSU and ITS sequence data.
Twenty–six taxa were included
in the combined analyses, which
comprised 1420 characters
(LSU: 910, ITS: 510) after
alignment. The best scoring
RA × ML tree with a final
likelihood value of is presented.
Bootstrap support values for
ML ≥ 50% and BYPP ≥ 0.95
are given above the nodes. The
tree is rooted with Conioscypha
lignicola CBS 335.93 and C.
japonica CBS 387.84
due to overlapping morphological characteristics. Liu
et al. (2019a, b, c, d) has revised the taxa with appendagebearing conidia in Sporocadaceae based on morphology
and multigene phylogeny. A new genus Parabartalinia was
introduced based on Bartalinia-like taxon.
Bartalinia bidenticola Htet, Mapook & K.D. Hyde, sp.nov.
Index Fungorum number: IF559553; Facesoffungi number: FoF10766; Fig. 117
Etymology: The name refers to the host plant from which
it was collected Bidens pilosa.
Holotype: MFLU 22-0103
Saprobic on dead stems of Bidens pilosa. Sexual
morph: Not observed. Asexual morph: Conidiomata
120–130 × 190–200 μm ( x̄ = 123 × 196 μm, n = 5), uniloculate, solitary, immersed to semi-immersed, globose
to subglobose. Ostiole absent. Peridium 23–30 μm wide,
3–4 layered, comprised of brown cells of textura globulosa. Conidiogenous cells 1–2 μm wide, filiform, hyaline.
Conidia 23–27 × 4–6 µm, ( x̄ ± SD = 24.9 ± 0.8 × 4.9 ± 0.3
µm), fusoid to ellipsoid, straight to slightly curved at the
apex, 4-septate, constricted ate the septa, brown in three
middle cells and hyaline at the basal and apical cells; basal
cells 2–4 µm long, hyaline, obconic to conic with a truncate base, thin-walled; three median cells 14–18 µm long,
( x̄ ± SD = 15.8 ± 0.8 µm), (second cell from the base pale
brown, 6–7 μm long; third cell pale brown, 4–6 μm long;
fourth cell pale brown, 4–7 μm long), doliiform, wall
rugose, versicoloured; apical cells 3–6 µm long, hyaline,
subcylindrical, smooth-walled; with 2–3 tubular appendages
(mostly 2), unbranched, filliform (13–)17–24(–24) µm long,
( x̄ ± SD = 22.7 ± 2.7 µm); basal appendage single, tubular,
unbranched, centric 4–8(–10) µm long.
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Fungal Diversity (2022) 117:1–272
(MFLUCC 12-0384) with 97.32% similarity and the closest
match for ITS sequence was Bartalinia pondoensis (CCTU
459) with 98.85% similarity. Phylogenetic analyses of a
combined ITS and LSU sequence dataset (Fig. 118) show
that Bartalinia bidenticola (MFLUCC 22-0008) is phylogenetically well distinguished and branched off from all other
species in Bartalinia with ML = 100% and BYPP = 1.00 support. We, therefore, identify our isolate as a new species
which was found from Bidens pilosa.
Fig. 117 Bartalinia bidenticola (MFLU 22–0103, holotype). a,b
Appearance of conidiomata on host substrate. c Section through
conidioma. d Peridium. e Conidia on the conidiogenous cells. f–i
Conidia. J–k Germinating conidia. l–m Culture on MEA from surface and reverse. Scale bar a = 300 µm, b = 200 µm, c = 50 µm,
d = 20 µm, e = 10 µm, f,g,h,I,j,k,l,m = 20 µm
Culture characteristics: Spores germinating on MEA
within 24 h, reaching 50 mm after 7 days at room temperature, irregular, undulate, concentric, flat, leathery surface,
grey.
Material examined: Thailand, Chiang Rai Province, Doi
Pui, on the dead stems of Bidens pilosa Linn var radiata
(Asteraceae), 10 July 2020, Zin Hnin Htet SW35 (MFLU
22-0103, holotype), ex-type living culture (MFLUCC
22-0008).
GenBank numbers: ON715467(LSU), ON715520(ITS).
Notes: Bartalinia bidenticola (MFLUCC 22-0008)
resembles Bartalinia in having subcylindrical to fusoid,
pale yellow septate conidia, with an apical cell modified
into branched appendages. According to the BLASTn
results, the closest match for the LSU sequence of Bartalinia
bidenticola (MFLUCC 22-0008) was Bartalinia kevinhydei
13
Bartalinia caryotae Senan., Kular. & K.D. Hyde, sp. nov.
Index Fungorum number: IF558407; Facesoffungi number: FoF10699; Fig. 119
Etymology: Species epithet refers to the host genus.
Holotype: HKAS115853
Associated with leaf-tip spots of Caryota sp. Sexual
morph: Not observed. Asexual morph: Coelomycetous.
Conidiomata 55–75 × 115–140 μm ( x̄ = 62 × 120 μm,
n = 15), pycnidial, superficial or rarely erumpent, solitary,
scattered, dark brown, uniloculate, conical, glabrous, ostiolate, without a papilla. Conidiomata wall 3–8 μm wide
( x̄ = 6 μm, n = 5), comprising several layers of brown,
pseudoparenchymatous cells of textura angularis, paler
towards the inner layers. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3–7 × 1.5–4 μm
( x̄ = 5 × 3 μm, n = 10), enteroblastic, annellidic, integrated,
hyaline to subhyaline, ampulliform to subcylindrical, or
obclavate, aseptate, thin- and smooth-walled. Conidia
17.5–19 × 5–7 μm ( x̄ = 18 × 6 μm, n = 20), subcylindrical,
hyaline when immature, olivaceous when mature, straight
to slightly curved, 4-septate, constricted at septa, smooth,
with longest cell second from base, bearing appendages
at both ends; basal cell 2–4 μm long, conical, hyaline, to
pale brown, paler than middle cells; second cell from base
5–8 μm long, pale brown; third cell 3–5 μm long, pale
brown; fourth cell 3–5 μm long, pale brown; apical cell
2–3 μm long, conical, hyaline, smooth-walled with a tubular, flexuous, divergently 3–4, unbranched, 20–30 μm long,
tubular appendages arising from the tip of apical cell, with
a basal 10–25 μm long, centric to eccentrically located,
unbranched, flexuous, tubular to filiform appendage.
Culture characteristics: Colonies on PDA reaching 2 cm diam. after 5 days at 16 °C, flattened, circular, smooth margin, white with off-white aerial mycelia;
reverse cream.
Material examined: China, Guangdong Province, Shenzhen, Nanshan District, Mountain Yangtai Forest Park, 22°
39′ 21.26″ N 113° 57′ 18.53″ E, living leaves of Caryota
sp (Arecaceae), 4 October 2019, ND Kularathnage, NDK
24, (HKAS115853, holotype), ex-type living culture,
ZHKUCC 21–0093; China, Guangdong Province, Shenzhen, Nanshan District, Mountain Yangtai Forest Park, 22°
39′ 21.26″ N 113° 57′ 18.53″ E, living leaves of Caryota sp
Fungal Diversity (2022) 117:1–272
149
Fig. 118 Phylogram generated from maximum likelihood
analysis based on combined
LSU and ITS sequence data
for Bartalinia. Twenty taxa
were included in the combined
analyses, which comprised
1479 characters (LSU = 897 bp,
ITS = 582 bp) after alignment. Bootstrap support
values for maximum likelihood (ML) ≥ 50% and clade
credibility values ≥ 0.90 from
Bayesian inference analysis are
labelled at each node. Ex-type
strains are in bold and the new
isolate is indicated in blue bold.
Hymenopleella endophytica
(APBSWTPF108, EML AS5-1,
EML AS5-2) were used as the
outgroup taxa
(Arecaceae), 15 September 2018, IC. Senanayake, SI 43,
(ZHKU 21–0005, paratype), ex-paratype living culture,
ZHKUCC 21–0094.
GenBank numbers: ZHKUCC 21–0093
– MZ520792(ITS), MZ520794 (LSU).
ZHKUCC 21–0094 –MZ520793(ITS), MZ520795 (LSU)
Notes: In the combined ITS and LSU gene analysis, Bartalinia caryotae formed a distinct clade with (MP = 100 and
BYPP = 1.00) support, basal to B. kevinhydei (Fig. 118). The
comparison of the DNA sequence of ITS and LSU loci of
Bartalinia caryotae with B. kevinhydei revealed base pair
differences of 0.3% and 0.4% respectively. However, Bartalinia caryotae is morphologically different from Bartalinia
kevinhydei in having small, conical conidiomata with large,
3–4 unbranched apical appendages (Liu et al. 2019a, b, c, d).
Therefore, we introduce Bartalinia caryotae as a novel taxon
from leaves of Caryota species and Bartalinia species are
rarely reported on monocotyledon plants. Our species forms
superficial to erumpent conidiomata on leaves.
Pestalotiopsis Steyaert, Bull. Jard. bot. État Brux. 19: 300
(1949).
Notes: Steyaert (1949), based on the conidial features,
divided Pestalotia into three genera, namely Pestalotia,
Pestalotiopsis and Truncatella. Species with 5-celled conidia
(4-septate) were grouped within Pestalotiopsis. Based on
multilocus phylogenetic and morphological analyses of
Pestalotiopsis-like species, Maharachchikumbura et al.
(2014) divided the complex into three genera: Pestalotiopsis,
Neopestalotiopsis and Pseudopestalotiopsis. Pestalotiopsis
is characterized by median cells concolourous, i.e. three
pale-pigmented median cells. Pestalotiopsis is a complex
genus and has considerable phenotypic diversity (Maharachchikumbura et al. 2014). Thus, the identification is complemented with DNA sequence data and phylogenetic analyses
based on the combination of three gene regions (ITS, tub2
and tef1) (Jeewon et al. 2003; Maharachchikumbura et al.
2011, 2012, 2013; Geng et al. 2013).
Pestalotiopsis piraubensis V.P. Abreu & O.L. Pereira, sp.
nov.
Index Fungorum number: IF556023; Facesoffungi number: FoF04861; Fig. 120
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150
Fungal Diversity (2022) 117:1–272
Fig. 119 Bartalinia caryotae
(HKAS115853, holotype).
a Specimen. b Conidiomata
on substrate. c Cross section
of conidiomata. d-f Conidia
attached to conidiogeneous cells. g-j Conidia. Scale
bars: c = 30 µm, d-f = 20 µm,
g-j = 15 µm
Etymology: Name refers to the city of Piraúba, state of
Minas Gerais, Brazil, where the fungus was collected.
Holotype: VIC 44199
Asexual morph: Culture obtained by direct isolation
from diseased guava fruits (Fig. 120a). On MEA, conidiomata sporodochial, globose, solitary, semi-immersed,
black, exuding globose, dark brown to black conidial masses
(Fig. 120d–e). Conidiophores indistinct, reduced to conidiogenous cells. Conidiogenous cells discrete, cylindrical
or spathulate, hyaline, smooth-walled, 4–10 × 1.5–3 μm.
Conidia fusoid, ellipsoid, straight to slightly curved, 4-septate, 25–33.5 × 5–7.5 μm, basal cell conic, hyaline, smooth
and thin-walled, 4.5–7.5 μm long; three median cells doliiform, 15.5–20.5 μm long, minutely verruculose, concolourous, pale brown, septa darker than the rest of the cell (second
cell from the base 5–7.5 μm long; third cell 4.5–6.5 μm long;
fourth cell 5–7 μm long); apical cell 3.5–6 μm long, hyaline, cylindrical to subcylindrical, thin- and smooth-walled;
with 1–3 tubular apical appendages, arising from the apical
crest, unbranched, filiform, flexuous 12–25.5 μm long; basal
13
appendage single, tubular, unbranched, centric, 2–6.5 μm
long (Fig. 120f–h). Sexual morph: Not observed.
Culture characteristics: Colonies on MEA attaining
43 mm diam after 7 days at 25 °C, with a regular edge, whitish to pale yellow-coloured, with dense aerial mycelium and
dark brown to black conidial masses (Fig. 120b). Colonies
on PDA attaining 68 mm diam after 7 days at 25 °C, with
irregular edge, whitish to pale yellow-coloured, with sparse
aerial mycelium on the central surface and dark brown to
black conidial masses (Fig. 120c).
Pathogenicity test: The inoculation method used consisted
of the wounding of the detached fruits and on these, the
mycelium plug, with the mycelial part facing the surface of
the fruit was added. Causing fruit rot disease, seven days
after the inoculation of healthy ripe guava fruits.
Material examined: Brazil, Minas Gerais, Piraúba, in a
commercial orchard, on fruits of Psidium guajava L. (Myrtaceae), 20 January 2014, V.P. Abreu & O.L. Pereira (VIC
44199, holotype), ex-type living culture COAD 2165.
GenBank numbers: MH627381 (ITS), MH643773 (tub2),
MH643774 (tef1-α).
Fungal Diversity (2022) 117:1–272
151
post-harvest management. The topology of the concatenated
tree was similar to that of individual trees, thus, only the
concatenated tree is presented here (Fig. 121). Molecular
data showed that Pestalotiopsis piraubensis COAD 2165
did not group with any other species reported in the literature. Morphologically, P. piraubensis differs from P. trachicarpicola for presenting larger conidia. Pestalotiopsis
piraubensis has 1–3 apical appendages and the other species have 2–4 (mostly 3). Pestalotiopsis kenyana and P.
biciliata differ from P. piraubensis by having conidiomata
pycnidial in culture on PDA. P. photinicola presents smaller
conidia (18–24 × 4–5 μm) than P. piraubensis. Although
most of the morphological characteristics did not differ so
much, the most striking feature of P. piraubensis was the
size of the conidiogenous cells, which was much smaller
(4–10 × 1.5–3 μm diam), when compared to the other species. Phylogenetic analyses and morphological comparisons
support the introduction of P. piraubensis as a new species
within this genus. Besides that, this study may be helpful for
further studies on the management of guava diseases.
Fig. 120 Pestalotiopsis piraubensis (VIC 44,199, holotype) a disease symptom on the fruit of Psidium guajava in a commercial
orchard in the city of Piraúba, state of Minas Gerais, Brazil. b Colony
on MEA (Malt Extract Agar) after 7 d at 25 ºC with a photoperiod of
12 h in the dark in Petri dishes (90 × 15 mm) (COAD 2165). c Colony
on PDA (Potato Dextrose Agar) after 7 d at 25 ºC with a photoperiod of 12 h in the dark in Petri dishes (90 × 15 mm) (COAD 2165).
d Conidioma sporulating on PDA. e Conidial masses. f Conidia (1–3
apical appendages). Scale bars: e = 50 µm, f–h = 20 µm
Notes: Pestalotiopsis spp. were previously considered
opportunistic pathogens that affect stressed plants (Coyier
and Roane 1987). Pirone (1978) reported different species
of Pestalotiopsis causing leaf spots on a range of ornamentals. However, in recent years, there has been an increase
in reports of these pathogens causing widespread damage
to several economically important crops (Keith et al. 2006;
Ko et al. 2007; Rodrigues et al. 2014; Rosado et al. 2015;
Solarte et al. 2018). Therefore, the increase of guava planting areas has contributed to the emergence of several diseases, and there is no data on the environmental requirements of Pestalotiopsis infection on guavas in Brazil, nor
any studies on field epidemiology for these diseases or
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor &
M.E. Barr, in Hyde, Fröhlich & Taylor, Sydowia 50(1): 23
(1998).
Notes: Apiosporaceae was introduced by Hyde et al.
(1998), with Apiospora as the type genus. Description of
Apiosporaceae has been provided in Hyde et al. (2020a,
b, c), wherein the authors listed five genera in this family,
Appendicospora, Arthrinium, Dictyoarthrinium and Endocalyx, Nigrospora. However, phylogenetic analysis resulted
in the transfer of Dictyoarthrinium and Spegazzinia to
Didymosphaeriaceae (Pleosporales) (Tanaka et al. 2015;
Samarakoon et al. 2020).
Apiospora Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova,
Sér. 4 4: 85 (1875).
Notes: Apiospora is characterized by densely arranged
perithecia arranged in a longitudinal stroma, clavate to
broadly cylindrical asci and apiospores in the sexual morph
and Arthrinium-like asexual morph (Hyde et al. 2020a, b,
c). Species of this genus commonly live as endophytes, epiphytes, saprobes on grass in Poaceae (Samarakoon et al.
2022). Apiospora and Arthrinium has no clear boundary
as to the morphological characteristics. However, Apiospora and Arthrinium sensu stricto were recognized in Apiosporaceae as two independent lineages in this family (Pintos and Alvarado 2021).
Apiospora guiyangensis Samarak., Jian K. Liu & K.D.
Hyde, in Samarakoonet al., Fungal Diversity 112: 19 (2022).
Index Fungorum number: IF558711; Facesoffungi number: FoF10187; Fig. 122
13
152
Fungal Diversity (2022) 117:1–272
Fig. 121 Phylogram generated from Bayesian Inference
analysis based on combined
ITS, tub2 and tef1 sequence
data for several closely related
species in Sporocadaceae.
Sequence data of type cultures,
ex-type or ex-epitype obtained
from Maharachchikumbura
et al. (2014), Chen et al. (2017a,
b), Akinsanmi et al. (2017), Liu
et al. (2017) were included in
this study. The combined genes
sequence analysis included 61
taxa, which comprise total 1994
characters (557 characters for
ITS, 830 characters for tub2,
607 characters for tef1-α), and
outgroup taxon Neopestalotiopsis saprophytica MFLUCC
12–0282. Bayesian posterior
probability are indicated at the
nodes, and values ≥ 0.95 are in
bold. Isolate numbers are indicated after species names. The
ex-type or ex-epitype strains are
in bold and black. The newly
generated sequence is indicated
in bold and blue
Saprobic on a dead twig of Bothriochloa ischaemum. Sexual morph: Stromata visible as fusiform, black, erumpent
pustules with alongitudinal slit on the top. Ascomata
123–165 × 109–185 ( x̄ = 140 × 130, n = 10) μm, subglobose,
unilocular, immersed, papillate, arranged in a linear row
along with the thelongitudinal slit. Peridium 11–15 ( x̄ = 13,
13
n = 10) μm in width, consisting of dark brown cell of textura
angularis, thin at base, becoming thicker near the ostioles,
with a pseudoparenchymatous wall at the most inner layer.
Paraphyses 3.6–7.6 ( x̄ = 5.2, n = 10) μm, hyaline, cylindrical, septate, guttulate. Asci 60–84 × 9.3–13 ( x̄ = 70 × 11,
n = 20) μm, fusiform, unitunicate, 8-spored, apex lacking
Fungal Diversity (2022) 117:1–272
153
Fig. 122 Apiospora guiyangensis (KUN-HKAS 125898,
new host record)a Host. b,
c Appearance of ascostroma
on host. d, e Section through
ascomata. f Peridium. g Paraphysis. h–j Asci (j is stained in
cotton blue). k–m Ascospores.
n Germinal spore. o Upper and
reverse view of cultures on PDA
at 8 days incubation. Scale bars:
d = 300 μm, e = 50 μm, f, h–j,
n = 20 μm, k–m = 10 μm
apical mechanism, with a short basal pedicel. Ascospores
22–28 × 4.6–6.7 ( x̄ = 25 × 5.4, n = 30) μm, hyaline, ellipsoid
to reniform, slightly curve, smooth-walled, aseptate, guttulate. Asexual morph: see description in Samarakoon et al.
(2022).
Culture characteristics: Colonies on PDA rapid growing,
reaching 45 mm diam. after 8 days at 20–25 °C, colonies
medium dense, circular, mycelium superficial in media, cottony, round aspect, white from above and reverse.
Material examined: China, Guizhou Province, Qianxinan
Buyei and Miao Autonomous Prefecture, Ceheng County,
Gaofeng Village, on dead culms of Bothriochloa ischaemum
(Poaceae), 8 August 2018, D. P. Wei, GFC613 (KUN-HKAS
125898), living culture KUNCC22-12539.
Known hosts and distribution: Poaceae (Guizhou, China)
(Samarakoon et al. 2022).
GenBank numbers: OQ029540 (ITS), OQ029613(LSU),
OQ061263(SSU), OQ186444(tef1), OQ186446 (tub2).
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154
Fungal Diversity (2022) 117:1–272
Fig. 123 RAxML tree inferred
from combined ITS, LSU, rpb2
and tub2 sequence. Bootstrap
support for maximum likelihood
analysis ≥ 50% and Bayesian
posterior probabilities ≥ 0.95
are denoted next to the notes in
this order. The new collection is
marked in blue bold font
Notes: Apiospora guiyangensis was introduced by
Samarakoon et al. (2022) from an unidentified Poaceae
species in Guizhou, China. Our species colonizing on dead
column of Bothriochloa ischaemum (Poaceae) was collected
from the same province of type strains of A. guiyangensis.
Its sexual morph fits well to the description of A. guiyangensis in the linear stromata with a slit-like opening, immersed,
subglobose, gregarious, ascomata and ellipsoid to reniform
ascospores. Additionally, the phylogenetic analysis of a combination of ITS-LSU-tef1 -tub2 shows our isolate sisters to
A. guiyangensis with ML = 100% and BYPP = 1.00 support
(Fig. 123). Thus, we determine our isolate as a new collection of A. guiyangensis and this finding indicates that A.
guiyangensis probably specific to Poaceae.
Xylariales Nannf.
Notes: Xylariales belongs to subclass Xylariomycetidae
and the placement is confirmed by different phylogenetic
and evolutionary studies (Maharachchikumbura et al. 2016;
Samarakoon et al. 2016a, b; Hongsanan et al. 2017). The
continuous taxonomic studies expand the number of families
13
and genera in Xylariales. Twenty-one families and 183 genera are listed in this order (Konta et al. 2016; Dayarathne
et al. 2017; Wijayawardene et al. 2022; Hyde et al. 2020a,
b, c).
Diatrypaceae Nitschke, [as 'Diatrypeae'], Verh. naturh. Ver.
preuss. Rheinl. 26: 73 (1869).
Notes: Diatrypaceae is a significant family in Xylariales
introduced by Nitschke (1869) with the generic type Diatrype (Augusto et al. 2016; Hyde et al. 2020a, b, c; Boonmee et al. 2021). Numerous saprobic, endopytic and pathogenic diatrypaceaous taxa are available in both terrestrial
and aquatic habitats worldwide (Dissanayake et al. 2021a,
b; Zhu et al. 2021; Wijayawardene et al. 2020). The family is characterized by erumpent or immersed ascostromata
that contain 8-spored or polysporous, long pedicel asci with
allantoid ascospores in the sexual morph and coelomycetous or hyphomycetous asexual morphs (Konta et al. 2020;
Carpouron et al. 2021; Dissanayake et al. 2021a, b). The
early identification of Diatrypaceae species were based only
on morphology. Currently most studies are performed by
Fungal Diversity (2022) 117:1–272
155
Fig. 124 Diatrypella quercina
(MFLU 18–1865, new host
record). a-c. Appearance
of ascostromata on a twig
of Quercus petraea subsp.
polycarpa host. d. Longitudinal
section of an ascoma. e. Ostiole.
f. Paraphyses. g. Peridium
h–k. Asci l-n. Ascospores.
a = 500 μm b-c, f = 200 μm,
d-e = 100 μm, h–k = 20 μm,
g = 10 μm, l-n = 5 μm
using both morphological observations with ITS and tub2
sequence data analyses (Konta et al. 2020; Dissanayake
et al. 2021a, b; Wijayawardene et al. 2022). However, many
genera of the family are polyphyletic and further taxonomic
studies still need to be performed for resolving diatrypaceous
taxa (Dissanayake et al. 2021a, b). There are 22 genera in
Diatrypaceae with more than 1500 species (Carpouron et al.
2021; Wijayawardene et al. 2020). In this study we discuss
a novel host record of Diatrypella species collected from
Russia.
Diatrypella (Ces. & De Not.) De Not., Sfer. Ital.: 29 (1863)
Diatrypella was established to constitute stromatic Sphaeriales with ovoid and multi spored asci (Croxall 1950,
Carpouron et al. 2021). Diatrypella was introduced by
Cesati and De Notaris (1863) and typified D. verruciformis.
The genus is characterized by a libertella-like coelomycetous asexual morph. In the sexual morph, conical to truncate
and discoid, cushion-like stromata are delimited by black
zones on the substrates, umbilicate or sulcate ostiolar necks,
long-stalked, cylindrical and polysporous asci with allantoid,
hyaline or yellowish ascospores (Kirk et al. 2008; Dissanayake et al. 2020; Hyde et al. 2020a, b, c). Recently several
Diatrypella species were introduced by different authors
from different hosts (Dissanayake et al. 2020; Hyde et al.
2020a, b, c; Zhu et al. 2021). There are 84 records under
Diatrypella in Species Fungorum (2022a, b) while, only 24
taxa have molecular data in GenBank. We follow the latest
13
156
treatment for Diatrypaceae in Boonmee et al. (2021) to
resolve the taxonomic placements of our strain and updated
phylogenetic tree is presented in Fig. 124. In this study,
we discuss a new collection of Diatrypella quercina from
Russia.
Diatrypella quercina (Pers.) Cooke, J. Bot., Lond. 4: 99
(1866).
Index Fungorum number: IF215896; Facesoffungi number: FoF11778; Fig. 124
Saprobic on dead twigs on Quercus robur. Sexual
morph: Stromata 1.0–1.5 mm in diam., well-developed, solitary to gregarious, immersed to semi immersed, erumpent
at the maturity, globose to subglobose, black. Ascomata
610–660 μm high, 550–600 μm diam. ( x̄ = 650 × 580 μm,
n = 10), perithecial, surrounded by white entostroma, 6–8
perithecia arranged in a valsoid configuration, conical,
individual ostiole with a long neck. Neck 450–490 μm long
( x̄ = 480 μm, n = 10), cylindrical, with periphyses. Peridium
20–30 μm wide ( x̄ = 26 μm, n = 10), composed outermost
layers of brown, thick-walled cells in textura angularis,
inner layers hyaline, cells forming of textura prismatica.
Hamathecium comprises 1–2 μm wide ( x̄ = 1.5 μm, n = 20)
paraphyses arising from base of perithecia, hyaline, long,
narrow, unbranched, septate, guttulate, narrowing and tapering towards apex. Asci 70–120 × 10–15 μm ( x̄ = 100 × 14 μm,
n = 30), polysporous, unitunicate, strongly curved, apically
round, with a J-apical ring, long pedicellate (50–70 μm).
Ascospores 6–9 × 1.8–2.3 μm ( x̄ = 7.5 × 2.0 μm, n = 30),
overlapping, hyaline, yellowish in mass, allantoid, aseptate,
guttulate, guttules conspicuous near to apex, smooth-walled.
Asexual morph: see Adamčíková et al. (2013).
Material examined: Russia, Sochi, Khostinsky City District, the territory of Subtropic Scientific Centre of Russia
Academy of on a dead branch of Quercus petraea subsp.
polycarpa (Schur) Soó (syn. Q. colchica Czeczott, Q. iberica Steven ex M. Bieb.) (Fagaceae), Timur S. Bulgakov, 4
August 2018, T-7330 (MFLU 18–1865).
GenBank number: ON705330 (ITS), ON713468 (tub2).
Notes: Ruhland (1900) considered Diatrypella quercina
as a species in Diatrype because of strongly developed
ectostromata. Later, Wehmeyer (1926) discussed the possibility of including this taxon in Diatrype. Croxall (1950)
distinguished D. quercina from other Diatrypella species
because of its strongly curved ascospores. Cryptovalsa and
Diatrypella also have polysporous asci and cannot easily
be distinguished, based on morphological comparisons
(Acero et al. 2004; Vasilyeva and Stephenson 2005; Dissanayake et al. 2021a, b). A number of taxonomic studies have
been done for D. quercina species based on morphological observations from different countries in the world (Farr
and Rossman 2022). Adamčíková et al. (2011) reported the
records of Libertella quercina based on their morphology on
13
Fungal Diversity (2022) 117:1–272
Castanea sativa from Slovakia. Libertella quercina was also
reported on bark of Quercus in England and France (Grove
1937; Adamčíková et al. 2011). Saccardo (1906) described
Cytosporina quercina (basionym Libertella quercina) on
branches of Quercus and Castanea in Italy, France, and
Germany and later, the taxon was identified as the asexual
morph of Diatrypella quercina (Grove 1937). Popov et al.
(2008) reported D. quercina on Quercus robur from Russia
based on morphology while our strain MFLU 18–1865, was
from Q. petraea subsp. polycarpa is also known as Georgian, or Colchician oak—the native oak species for western
Caucasus. However, no genetic studies were reported from
Russia for this taxon. There were only two studies providing molecular data for the species from Spain (on Quercus
faginea) (Acero et al. 2004; Vu et al. 2019 taken from GenBank 2022). We provided a comprehensive taxonomic study
for D. quercina based on their morpho-molecular and phylogenetic analyses. In our phylogenetic analyses our strain
(MFLU 18–1865) grouped with other isolates of D. quercina
(CBS 108.18 and DL30M) in Clade B, with 85% MLBS,
0.95 BYPP support. Also, our strain forms a sister lineage to
CBS 108.18 and grouped with high bootstrap support (95%
ML). In comparison of base pair differences between CBS
108.18 and DL30M (D. quercina), 6 bp differences (1.16%)
have revealed by 513 nucleotides in ITS region while tub2
regions are not available. However, we provided tub2
sequence data for D. quercina in this study. We revealed
1 and 5 pb differences from 515 nucleotides (0.19% and
0.97%) in ITS regions when comparing our strain (MFLU
18–1865) with CBS 108.18 and DL30M respectively. Trimen et al. (1866, https://www.biodiversitylibrary.org/page/
16233050) provided incomplete morphology for the taxa
and these characters are matched with the morphology of
our strain. Based on referred morpho-molecular data we
conclude that our stain should be another collection of D.
quercina from Russia. This is the first genetic study on D.
quercina in Russia with detailed morphology and it is also
the new host record on Q. petraea subsp. polycarpa. However, detailed morphological studies are suggested in future
for this taxon (Figs. 124, 125).
Hypoxylaceae DC., in Lamarck & de Candolle, Fl. franç.,
Edn 3 (Paris) 2: 280 (1805)
Xylariales comprises with 26 families. Xylariaceae and
Hypoxylaceae are two of the most diverse families in this
order. Hypoxylaceae is represented by 16 genera including
which comprises sexual morphs 16 and asexual morphs
11 genera (Wijayawardene et al. 2022). Hypoxylaceae are
distributed in tropical, subtropical regions (Lambert et al.
2019), known for its huge species diversity and abundant bioactive secondary metabolites (Helaly et al. 2018)
and plays an important ecological role in protecting host
plants from pathogens (Song et al. 2022). The family is
Fungal Diversity (2022) 117:1–272
Fig. 125 Phylogram generated from maximum likelihood analysis
based on combined ITS and tub2 sequence data representing Diatrypaceae in Xylariales. Related sequences are taken from Boonmee et al. (2021) and additions according to the BLAST searches in
NCBI. Hundred and thirty-five strains are included in the combined
analyses which comprised 912 characters (517 characters for ITS and
157
395 characters for tub2) after alignment. Kretzschmaria deusta (CBS
826.72) and Xylaria hypoxylon (CBS 122,620) in Xylariaceae (Xylariales) were used as the outgroup taxa. Bootstrap support values for
ML ≥ 75% are given above the nodes (left side). Bayesian posterior
probabilities (BYPP) ≥ 0.95 are given above the nodes (right side).
Ex-type strains are in bold and newly generated sequence is in red
13
158
Fig. 125 (continued)
13
Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
159
Fig. 126 Hypoxylon inaequale.
(HKAS123207, holotype)
a Substrate. b, c Stromata
showing ostioles. d, e Vertical section through stromata.
f Perithecium. g Peridium.
h–k asci. l paraphyses. m–q
Immature to mature ascospores.
r, s Germinating ascospores. t,
u Culture on PDA. Scale bars:
e–f, h = 200 μm, g = 100 μm
i–l = 50 μm, m–s = 5 μm
characterised by carbonised stromatal tissue. Stromata surface is usually blackened when mature, olivaceous, with %10
KOH are purplish or orange. Ostioles that are always higher
than the level of stromatal surface (Cruz et al. 2021).
Hypoxylon Bull., Hist. Champ. Fr. (Paris) 1(1): 168 (1791)
Hypoxylon was introduced by Bulliard, that contains primarily saprotrophs and endophytes of angiospermous plants.
The type genus Hypoxylon is the largest genus in the Hypoxylaceae, with more than 200 species (Pourmoghaddam et al.
2020). Members of the genus have a worldwide distribution,
but they display a higher diversity in the tropics and subtropics (Kuhnert et al. 2014). In the twentieth century, the
generic concept of Hypoxylon was based only on morphological characteristics (Ju and Rogers 1996). Currently, morphological, phylogenetic, and chemotaxonomic evidence,
has also been used to infer species limits in inter- and intragenera in Hypoxylaceae (Sir et al. 2016). Hypoxylon is quite
common in China; however, the occurrence of the species
in China has not been confirmed by molecular phylogenetic
13
160
Fungal Diversity (2022) 117:1–272
Fig. 127 Phylogram generated from maximum likelihood analysis
based on combined ITS, tub2 sequence data representing Hypoxylaceae in Xylariales. Related sequences are taken from (Cedeño–
Sanchez 2020) and additions according to the BLAST searches in
NCBI. Seventy-two strains are included in the combined analyses
which comprised 1654 characters (532 characters for ITS,1122 char-
acters for tub2) after alignment. Biscogniauxia nummularia (MUCL
51,395) and Annulohypoxylon annulatum (CBS 140775) in Xylariales were used as the outgroup taxa. Bootstrap support values for
ML ≥ 70% are given above the nodes (left side). Bayesian posterior
probabilities ≥ 0.95 are given above the nodes (right side). Ex-type
strains are in bold and newly generated species is in red
analyses, and the species diversity and distribution of the
genus in China are unclear (Hyde et al. 2020a, b, c). The
aims of this study were to confirm the taxonomic status of
the new species, explore the species diversity of Hypoxylon,
and infer the evolutionary relationships of Hypoxylon.
n = 15), glomerate, fawn, sessile, gregarious or solitary,
with white ostioles. Perithecia 250–300 × 260–450 μm
(x̄ = 275 × 365 μm, n = 15), spherical to obovoid, completely
immersed in stromata. Ostioles 90–120 μm diam, umbilicate, white. Paraphyses 2.5 × 3.1 μm wide, copious, filiform, aseptate, unbranched. Asci 132 − 153 × 9–11 μm
(x̄ = 142 × 10 μm, n = 10), cylindrical, 8-spored, uniseriate,
stipitate, with apical apparatus colorless in Melzer’s reagent.
stipe 40–112 μm (n = 10). Ascospores 9.8–11.8 × 5.0–5.9 μm
(x̄ = 10.45 × 5.45 μm n = 20), pale dark brown when immature, becoming dark brown with age, ellipsoid-inequilateral
with round ends, unicellular, with two round guttulae,
obliquely arranged. Asexual morph: Not observed.
Hypoxylon inaequale S.C. He & Jayaward., sp. nov.
Index Fungorum number: IF900071; Facesoffungi
number:FoF13393; Fig. 126
Etymology: Based on the inequilateral spore character.
Holotypus: HKAS123207.
Saprobic on dead stem plant from Itoa orientalis. Sexual
morph: Stromata 40–50 × 20–30 mm (M = 45 × 25 mm,
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161
Fig. 128 Astrocystis bambusicola (KUN-HKAS 125897,
new record) a Substrate. b, c.
Ascomata. d Section through
ascoma. e–h Asci. i Paraphysis.
j Asci with J.+ apical ring.
k Ascospore. l, m Upper and
reverse view of cultures on
PDA at 8 days incubation. n
Germinating spore. Scale bars:
d, e = 50 μm, f–j = 30 μm, k, n
= 10 μm. (j stained in Melzer's
reagent)
Culture characteristics: Culture was made from germinating ascospores and was incubated with PDA media at
25 ℃, reaching 2.2–2.4 cm in 24 days. The colony white,
occasionally raised, fluffy, with dense mycelia, umbonate
margin, reverse chrome yellow.
Material Examined: China, Yunnan Province, Kunming
City, Kunming Institute of Botany, Chinese Academy of
Sciences, on Itoa orientalis Hemsl. (Salicaceae), 30 August
2021,Shu-Cheng He, HSC20B (HKAS123207, holotype);
ex-type living culture, KUNCC22-10798.
GenBank numbers: ON329812 (ITS).
Notes: Based on the multi-gene phylogenetic results, our
specimen is closely related to Hypoxylon delonicis (Perera
et al. 2020). Based on morphology, our strain differes from
Hypoxylon delonicis by having wider asci and shorter and
woder ascospores. Based on a megablast search of the
NCBIs nucleotide database using the multi-gene sequence,
the highest similarities ITS (GenBank KU683766; Identities = 480/541(89%), Gaps = 17/541(3%)), tub2 (GenBank
AY951740; Identities = 1126/1265(89%), other species of
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162
Fungal Diversity (2022) 117:1–272
Fig. 129 RAxML tree inferred from combined ITS, LSU, rpb2 and tub2 sequence. Bootstrap support for maximum likelihood analysis ≥ 50%
and Bayesian posterior probabilities ≥ 0.95 are denoted next to the notes in this order. The new collection is marked in blue bold font
Hypoxylon delonicis with strong bootstrap support (82/0.99,
Fig. 127). We identified Hypoxylon inaequale as a new species of Hypoxylon.
Xylariaceae Tul. & C. Tul. [as 'Xylariei'], Select. fung. carpol. (Paris) 2: 3 (1863).
Notes: Xylariaceae is the type and largest family of
Xylariales (Hyde et al. 2020a, b, c). Number of genera
in this family has been subjected to multiple revision in
different articles. Kirk et al. (2001) and Eriksson (2007)
estimated that there are 75 genera and about 800 species
in Xylariaceae. Hyde et al. (2020a, b, c) counted 32 genera in this family and provided notes for each accepted
genus. Wijayawardene et al. (2022) listed 38 genera in
13
Xylariaceae, without giving supported annotation. Xylariaceae can live as saprobes, pathogens, or endophytes,
habiting wood, leaves and fruits (Hyde et al. 2020a, b, c).
Xylariaceae is characterized by perithecia embedded in
erect, applanate or effuse-pulvinate, dark-coloured stromata, cylindrical asci with an amyloid apical ring, brown
to black, 1–2-celled, ellipsoidal, subglobose or reniform,
with with germ slits or pores (Tang et al. 2009; Maharachchikumbura et al. 2016; Hyde et al. 2020). Xylariaceae
was divided into two subfamilies including Xylaroideae
and Hypoxyloideae based on their respective anamorphic
types, their stromatal pigments and secondary metabolites.
Xylaroideae generally is considered to comprise species
with Nodulisporium-type anamorph and KOH+ stromatal
Fungal Diversity (2022) 117:1–272
163
Fig. 130 Xylaria venosula
(Herbarium AMH-10068, new
record) a, d, e Stromata on
decaying host, b Vertical section of stromata c Horizontal
section of stromata f Paraphyses, g-i Asci j Textura intricata
k Peridium, l m Germinating
spore n, o Culture on MEA
plates, p Hypha. Scale bars:
k = 50 μm, f-i = 20 μm. j, m, n,
p = 10 μm
pigments, while it is Geniculosporium-type and KOH− in
Hypoxyloideae. Multigene phylogenetic analysis based on
LSU, SSU and rpb2 sequence well reflect these two subfamilies (Samarakoon et al. 2016a, b).
Astrocystis Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74):
123 (1873) [1875].
Notes: Astrocystis was introduced by Berkeley and
Broome (1850) and typified by Astrocystis mirabilis. Index
fungorum (2022a, b) lists 28 epithets in Astrocystis, while
four of them have been excluded from this genus. Species
of this genus mainly exist in saprobes on monocotyledonous substrates, such as bamboo, palm and Smilax (Laessøe
and Spooner 1993). Astrocystis is characterized by uni- to
multi-peritheciate stromata, carbonaceous peridium, asci
with a relatively short stipe and small, amyloid, stoppershaped ascal ring (Pinnoi et al. 2010). It is widely accepted
that Astrocystis bear similar morphology with Rosellinia,
while the former genus can be distinguished from the latter
by small and cylindrical or funnel-shaped apical ring of asci.
Rosellinia has conspicous, barrel-shaped apical apparatus
(Dulymamode et al. 1998).
Astrocystis bambusicola R.H. Perera & K.D. Hyde, in Hyde
et al., Fungal Diversity 87: 173 (2017).
Index Fungorum: IF553799; Facesoffungi number:
FoF10187; Fig. 128
Saprobic on dead culms of Microstegium sp. appearing
as black raised spots on the host. Sexual morph: Ascomata
up to 173 μm in width and 135 μm in high, subglobose,
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164
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Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
◂ Fig. 131 Maximum parsimony tree generated from by using the ITS
sequences belongs to Xylariaceae species. The tree includes the ML,
MP and BYPP values. In the phylogenetic tree the sequence analysis of Xylaria venosula NFCC-4520 (black bold letters) is shown with
other species of Xylaria and Rosellinia australiensis as out-group
black, perithecial, gregarious, superficial, 1–2-loculate,
with flattened top and projecting papillas. Peridium 43–60
(x̄ = 50, n = 10) μm, composed of membranous inner wall
and black, fragile, carbonaceous outer wall. Paraphyses
3.8–8.3 (x̄ = 6.2, n = 25) μm, septate, cylindrical, hyaline,
unbranched, guttulate. Asci 84–121 × 6.8–11 (x̄ = 99 × 8.8,
n = 30) μm, 8-spored, unitunicate, cylindrical, with a
short pedicel and J+ apical ring. Ascospores12–15 × 5.7–8
(x̄ = 14 × 6.5, n = 50) μm, reniform, dark brown to black, uniseriate, aseptate, guttulate, smooth-walled, with a germ-slit,
without a gelatinous sheath. Asexual morph: Not observed.
Culture characteristics: Culture was made from germinal ascospores that germinated on PDA within 24 h,
at 23–25 °C. Colonies rapidly growing on PDA, reaching
30 mm at 8 days, white from above and reverse, cottony,
circular, umbonate, edge irregular.
Material examined: China, Guizhou Province, Qianxinan Buyei and Miao Autonomous Prefecture, Ceheng
County, Gaofeng Village, on dead culms of Microstegium
sp. (Poaceae), 8 August 2018, D. P. Wei, GFC612 (KUNHKAS 125898), living culture KUNCC 22-12539.
Known hosts and distribution: Bamboo (Yunnan, China;
Thailand) (Hyde et al. 2017, 2020a, b, c).
Genbank numbers: OQ029540 (ITS), OQ029613 (LSU),
OQ061263 (SSU), OQ186444 (tef1), OQ186446 (tub2).
Notes: Astrocystis bambusicola has been reported on
bamboo column from China and Thailand (Hyde et al.
2017, 2020a, b, c). Our isolate phylogenentically groups
with Astrocystis bambusicola with great support (100%
ML/1.00 BYPP, Fig. 129). Morphologically our isolate bears
resemblance with Astrocystis bambusicola in the subglobose, black, superficial ascomata, carbonaceous peridium,
cylindrical asci with J+ apical ring and reniform, dark brown
to black ascospores with a germ-slit. We introduce our isolate as a new host record species of Astrocystis bambusicola
from Microstegium sp. in Guizhou Province, China.
Xylaria Hill ex Schrank, Baier. Fl. (München) 1: 200
(1786).
Notes: Xylaria is one of the largest genera in Xylariaceae
and it includes more than 600 species (Hyde et al. 2020a, b, c;
Boonmee et al. 2021). Its occurrence in diverse environments
shows its unique role as saprobes, endophytes and as plant
pathogens (Edwards et al. 2003; Ju et al. 2018; Hyde et al.
2020a, b, c). Xylaria species mostly have long stalked threadlike macro structures. A few species of this genus coexist with
plants as endophytes (Chen et al.2013), which may later turn
into saprobes (Promputtha et al. 2007) when plants die. They
165
also extend their habits as coprophilous and endolichenic
(Piasai and Manoch 2009; Cañón et al. 2019). Most Xylaria
species also serve as economically important compound producers (Ratnaweera et al. 2014; Adeleke and Babalola 2021;
Wangsawat et al. 2021; Becker and Stadler 2021), which act as
antibacterial, antifungal and/or biocontrol agents. Therefore,
there is a need for the discovery of novel species and new
geographical records of this genus. Annually ten or more, new
species are introduced to this genus. For instance, in 2020, ten
new species were discovered and in the next year, 15 species
were discovered (Index Fungorum 2022a, b) by morphology
and molecular data.
Xylaria venosula Speg. Boletín de la Academia Nacional de
Ciencias en Córdoba 11 (4): 511 (1889).
Index Fungorum number: IF 247711; Facesoffungi number:
FoF09866; Fig. 130
Saprobic on decaying twig. Sexual morph: Ascostromata 1 cm long, superficial, aggregated in clusters, surface undulated, rarely with parallel cracks, with acute apices. Ascomata 340–385 × 370–450 μm ( x̄ = 362 × 406 μm,
n = 5), globose, erumpent, pulvinate, with central periphysate necks 90–120 × 60–100 μm ( x̄ = 101 × 75 μm,
n = 5). Peridium 32 μm wide, with brown to hyaline,
textura porrecta cell layers. Hamathecium: paraphyses
septate, branched, 2.8 μm wide, longer than asci, sparsely
present. Asci 110 –132 × 5.6–8 μm ( x̄ = 117.5 × 6.6 μm,
n = 25), unitunicate, 8–spored, cylindrical, apically
rounded with J + apical rings, rings 2.6–3.6 × 1.7–2.3 μm
( x̄ = 3.2 × 2 μm, n = 25), long-pedicellate, persistent.
Ascospores 12.5 –15.5 × 4.7–7 μm ( x̄ = 13.6 × 5.7 μm, n = 25),
overlapping uniseriate, hyaline to brown at maturity, oblong
to navicular, with straight germ slits, uni-guttulate, obtuse
ends, smooth-walled. Asexual morph: Not observed.
Distribution: Brazil, China, Ecuador, India and USA.
GenBank numbers: MZ292933 (ITS).
Material examined: India, Andaman and Nicobar
Islands, South Andaman, Mount Harriet, (11° 71′ 09.8″ N
92° 73′ 30.6″ E), recorded on an unidentified decaying log,
7 December, 2017, M. Niranjan and V. V. Sarma (PUFNI
1763). Herbarium submitted in Ajrekar Mycological Herbarium-AMH (AMH-10068) and Living culture (NFCC-4520)
deposited in National Fungal Culture Collection of India
(NFCCI), Pune.
Notes: The references for descriptions of X. venosula
could be found in Index Fungorum (https:// www. biodi
versi tylib rary. org/ page/ 29371 43# page/ 534/ mode/ 1up)
and the global fungal red list (http:// iucn. ekoo. se/ iucn/
speci es_ view/ 247711). The present taxon has morphological characteristics that are similar to the type and
other with slight differences. The present collection consists of paraphyses, smaller asci (109 –132 × 5.6–8 vs.
90–400 × 7–9) and oblong to navicular, smaller ascospores
13
166
Fig. 132 Chlorophyllum squamulosum (SDBR-CMUNK0585, new
record). a. Basidiomata. b. Basidiospores. c. Basidia. d. Cheilocystidia. Scale bars: a = 50 mm; b = 5 μm; c, d = 10 μm
(12.5 –15.5 × 4.7–7 vs. 14 –18 × 6–7 μm). Geographical
distribution of X. venosula in five countries mentioned in
distribution (https://www.gbif.org/occur rence/search?q=
Xylaria%20venosula&taxon_key=5487903) and this is the
first report of X. venosula from the Andaman Islands, India
and as such the present collection extends the geographical
distribution and range of this taxon globally (Fig. 131).
Basidiomycota R.T. Moore.
We follow the latest treatments of Basidiomycota in
Zhao et al. (2017), He et al. (2019) and Wijayawardene
et al. (2022).
Agaricomycotina Doweld.
Agaricomycetes Doweld.
Agaricales Underw.
Agaricaceae Chevall.
Notes: Agaricaceae was erected by Chevallier (1826)
based on the type genus Agaricus L. Previously, this family
contained only gilled fungi. However, research-based on
molecular data transferred many non-gilled fungal families
such as Lycoperdaceae, Nidulariaceae and Tulostomataceae to Agaricaceae. Now, it is represented by more than
1300 species belonging to 85 genera (Kirk et al. 2008; He
et al. 2019; Wijayawardene et al. 2022). Some gasteroid
pufball genera included in the Agaricaceae are Arachnion
Schwein., Bovista Pers., Calvatia Fr. and Lycoperdon Pers.
Chlorophyllum Massee, Bull. Misc. Inf., Kew (no. 138):
135 (1898).
Chlorophyllum (Agaricaceae, Agaricales) was introduced by Massee (1898) with C. molybdites (G. Mey.)
Massee as the type species. Chlorophyllum species are
widely distributed in tropical, subtropical, and temperate
13
Fungal Diversity (2022) 117:1–272
areas throughout the world as saprobes (Ge and Yang
2006; Kirk et al. 2008; Crous et al. 2015a, b, c; Ge et al.
2018; Dutta et al. 2020). This genus is characterized by
agaricoid, secotioid or sequestrate habits, a hymenidermal
pileus covering and smooth stipe, white, green or brown
basidiospores either lacking a germ pore or with a germ
pore that is caused by a depression in the episporium (Ge
and Yang 2006; Vellinga 2002, 2003a, 2004; Crous et al.
2015a, b, c; Loizides et al. 2020). Most Chlorophyllum
species are known to be poisonous (Vellinga and de Kok
2002; Leudang et al. 2017). There are 28 accepted species of Chlorophyllum in Index Fungorum (2022a, b).
Chlorophyllum is divided into six infrageneric sections:
Chlorophyllum Massee, Ellipsoidospororum Z.W. Ge,
Endoptychorum (Czern.) Z.W. Ge, Parvispororum Z.W.
Ge, Rhacodium Z.W. Ge and Sphaerospororum Z.W. Ge
based on morphological and phylogenetic analyses (Ge
et al. 2018). Only four Chlorophyllum species, C. globosum (Mossebo) Vellinga, C. hortense (Murrill) Vellinga,
C. molybdites (G. Mey.) Massee and C. rhacodes (Vittad.)
Vellinga, have been reported from Thailand (Chandrasrikul et al. 2011; Leudang et al. 2017; Ge et al. 2018;
Sysouphanthong et al. 2021; Suwannarach et al. 2022).
Chlorophyllum squamulosum A.K. Dutta, Soumili Bera &
K. Acharya, Phytotaxa 451: 121 (2020).
Index Fungorum number: IF835117; Facesoffungi number: FoF10684. Fig. 132
Basidiomata agaricoid, medium to large. Pileus
50–75 mm in diam., convex to broadly convex, often with
a shallow central depression, sometimes with an upturned
margin with age; surface white to cream, covered with
squamules, entire at the disc, elsewhere disrupting in some
specimens, mostly small plate-like, arranged in a concentric manner from center towards the margin, flat or curved
upwards, greyish brown (8E3) to reddish-brown (8E4) or
dark brown (8F5) at the center, elsewhere greyish brown
(8D3) to brownish grey (7C2) or dull red (9C3). Lamellae
4–6 mm broad, adnexed, crowded with two series of lamellulae, white to cream, concolorous; edge even to slightly
wavy, yellowish with KOH. Stipe 60–90 × 9–12 mm, central, cylindrical, gradually broader towards the base, at base
15–27 mm wide and bulbous to subbulbous, hollow, white
(6A1), turning brown (6D6–6E7) on bruising or with KOH.
Annulus double; upper portion concolorous with the stipe
surface; lower portion white to cream, sometimes with
greyish brown (8D3) to reddish-brown (8E4) at the margin,
rarely with a brownish border; edge sometimes floccose.
Basidiospores 7.5–12 × 5–7.5 μm (n = 50), Q = 1.25–1.85,
Qm = 1.5 ± 0.13, ellipsoid, smooth, hyaline, dextrinoid,
with a prominent wide germ-pore, truncated, thick-walled;
apiculus short, 0.5–1 μm long. Basidia 25–50 × 8–10 μm,
cylindrical to clavate or subclavate, thin-walled, 4-spored,
Fungal Diversity (2022) 117:1–272
167
Fig. 133 Phylogenetic tree derived from maximum likelihood analysis of a combined ITS and LSU genes of 38 sequences and the
aligned dataset was comprised of 1613 characters including gap. The
average standard deviation of the split frequencies of the BI analysis
was 0.00481. Agaricus megacystidiatus (MFLU 12–0137) and Xanthagaricus flavosquamosus (GDGM50918) were used as outgroup
taxa. The numbers above branches are the bootstrap statistics percentages (left) and Bayesian posterior probabilities (right). Branches with
bootstrap values ≥ 70% are shown at each branch and the bar represents 0.1 substitutions per nucleotide position. Hyphen (-) represents
support values ≤ 70%/0.90. Ex-type strains are in black bold. The
newly generated sequences are indicated in blue
sterigmata upto 10 μm long, cylindrical. Pleurocystidia
absent. Cheilocystidia 17–25 × 7.5–12 μm, clavate or
spheropedunculate, hyaline, thin-walled. Annulus hyphae
3–7 μm broad, tightly arranged, hyaline, often branched,
hyphal end obtuse to clavate, thin-walled. Pileipellis (pileal
squamules) a tightly packed hymeniderm, with cylindrical
and flexuous, or narrowly clavate terminal elements, measuring 5–12.5 μm broad, with pale brown intracellular pigments, often incrusted, sometimes branched, thin-walled.
Pileus trama hyphae 7–14 μm broad, incrusted, thin-walled.
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Fungal Diversity (2022) 117:1–272
Fig. 134 Lepiota metulispora
in habitat. a-c HNL503136 d-e
HNL503151
Fig. 135 Lepiota metulispora (HNL503151, new record). a Basidiospores b Cheilocystidia c Pileus covering
13
Stipitipellis hyphae 5–10 μm broad, parallel to subparallel, light yellowish with KOH, sometimes branched, nonincrusted, thin-walled. Stipe trama hyphae 5–12 μm broad,
incrustations present, parallel to subparallel, thin-walled.
Caulocystidia absent. Clamp connections absent in all the
tissues.
Material examined: Thailand, Chiang Mai Province, Muang District, Chiang Mai University, 18°48′2″N
98°57′18″E, elevation 335 m, solitary on soil in grassland,
3 August 2019, J. Kumla, SDBR-CMUNK0585, 18°48′14″N
98°57′15″E, elevation 333 m, solitary on sandy humus
mixed soil, 26 July 2020, J. Kumla, SDBR-CMUNK0731.
Habitat: Solitary, on sandy humus mixed soil in the dry
deciduous forests and grassland.
Distribution: Known from India and Thailand (Dutta
et al. 2020; this study).
GenBank numbers: SDBR-CMUNK0585- MZ4502085
(ITS), MZ452086 (LSU)
Fungal Diversity (2022) 117:1–272
SDBR-CMUNK0731- MZ4502084 (ITS), MZ452070
(LSU)
Notes: Chlorophyllum squamulosum belongs to the
Chlorophyllum section Rhacodium based on a combination
of morphological and molecular data (Fig. 133). Morphologically, C. squamulosum is similar to C. nothorhacodes,
C. rhacodes, C. olivieri and C. brunneum by its brownish
to reddish colouration of the stipe upon bruising, truncated basidiospores and clavate cheilocystidia. The phylogenetic tree indicated that C. squamulosum formed a sister
taxon to C. nothorhacodes. However, C. nothorhacodes
differs from C. squamulosum by its much larger basidiocarp (pileus of up to 280 mm in diam. and stipe up to
250 mm × 25–60 mm) (Vellinga 2003a). Furthermore,
C. rhacodes differs from C. squamulosum by its broader
basidiospores (9.8–11.1 × 6.3–7.7 μm), comparatively
larger cheilocystidia (16–43 × 8.5–25 μm) and the presence of clamp connections in the basidia, cystidia and
tramal hyphae (Vellinga 2001, 2003b). Chlorophyllum
olivieri has clamped basidia and broader cheilocystidia
(35–45 μm) when compared with C. squamulosum (Vellinga 2001). Chlorophyllum brunneum differs from C.
squamulosum by the presence of clamp connections at the
base of the basidia and cystidia (Bougher and Syme 1998;
Vellinga 2002, 2003a).
Lepiota (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 601
(1821).
Lepiota belongs to Agaricaceae, and is consisted of 450
species (He et al. 2019). Vellinga (2001) accepted Lepiota
for six sections based on the morphology, which are Sect.
Echinatae Fay., Sect. Fuscovinaceae Bon & Candusso,
Sect. Lepiota (Pers.) Gray, Sect. Lilaceae M. Bon, Sect.
Ovisporae (J.E. Lange) Kühner and Section Stenosporae
(J.E. Lange) Kühner. However, the genus is not monophyletic according to many molecular studies (Vellinga 2003a,
b; Liang et al. 2011; Hou and Ge 2020). In this study, three
species of Lepiota are recorded for the first time in Laos, and
a new species is described from Thailand.
Lepiota metulispora (Berk. & Broome) Sacc., Syll. fung.
(Abellini) 5: 38 (1887).
Index Fungorum number: IF461315; Facesoffungi number: FoF09887; Figs. 134, 135
Pileus 30–65 mm, sub umbonate to umbonate, expanding to plano-concave, with inflexed margin; greyish orange
to brownish orange (6B5-8, 6C7-8) glabrous at umbo, with
concolorous squamules toward margin on white fibrillose background; margin sulcate, with partial veil remnants, with concolorous squamules on surface. Lamellae
free, broadly ventricose, 3–5 mm wide, white, moderately
crowded, with 3 length lamellulae, with eroded edge. Stipe
60–80 × 5–4 mm, cylindrical or slightly tapering to apex,
169
completely fibrillose or cortinate at annular zone, white,
with white to greyish orange to brownish orange (6B5-8,
6C7-8) squamules under annular zone downward base, on
white to orange-white (5A2) background. Annulus an annular zone, cortinate with white fibrils. Context in pileus white,
up to 4 mm wide; in stipe hollow, concolorous with surface.
Smell and taste unknown. Spore print white.
B a s i d i o s p o re s [ 5 0 , 2 , 2 ] 1 3 – 1 6 . 5 × 4 – 5 µ m ,
avl × avw = 17.7 × 4.7 µm, Q = 2.91–3.30, Qav = 3.20, in
side-view cylindrical, with attenuate or rounded apex, with
straight abaxial side, with an inflexed hilar appendage, with
suprahilar depression, fusiform to cylindrical in frontal
view, slightly thick-walled, hyaline, dextrinoid, congophilous. Basidia 22–33 × 7–9 µm, clavate, slightly thick-walled,
hyaline, 4-spored. Cheilocystidia 15–35 × 5–20 µm, clavate
to broadly clavate, sometimes utriform, branched or with
septate under element cell, thick-walled, hyaline. Pileus
covering a trichoderm made up of two layers of element;
upper layers made up of cylindrical elements with rounded
or attenuate apex, 70–190 × 5–13 µm, hyaline to pale brown,
slightly thick-walled, smooth-walled, with parietal pigment;
under layers made up of shortly clavate to clavate elements,
35–50 × 8–14 µm, smooth or rough-walled, with hyaline to
parietal pale brown pigment. Stipe covering of squamules
similar to pileus covering. Clamp connections present.
Material examined: Laos, Oudomxay Province, Xay District, Houay Houm Village, N 20° 32′ 00.67″, E 101° 53′ 48.
17.16″. 917 m., 15 June 2014, P. Sysouphanthong, PS20141465 (HNL503136); ibidem, 20 July 2014, P. Sysouphanthong, PS2014-1480 (HNL503151).
Habitat and distribution: solitary or grow in a small cluster with few basidiomes, on dead leaves and humus soil,
saprotrophic. The species was only reported from tropical
regions viz. Sri Lanka (Pegler 1972), India (Kumar and
Manimohan 2009), China and Hong Kong (Liang et al.
2011), Tanzania (Pegler 1977), Thailand (Sysouphanthong
et al. 2012). This is the first report of L. metulispora in
northern Laos.
GenBank numbers: HNL503136–MT436261 (ITS).
HNL503151–MT436262 (ITS).
Notes: Lepiota metulispora has a trichodermal pileus covering and penguin-shaped basidiospores, and it is placed in
the section Lepiota (Vellinga 2001). The species is widespread in tropical countries. Lao specimens were found in
the mature stage, colour of squamules on pileus is paler than
Thai specimens, but the morphology is identical (Sysouphanthong et al. 2012). The type specimen of the species
from Sri Lanka is closer to Lao and Thai specimens in morphology, but basidiospores are slightly larger and cheilocystidia are undetermined (Pegler 1972). Liang et al. (2011)
studied the type material from Sri Lanka and compared it
with the Chinese specimen, and the type specimen has a
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170
13
Fungal Diversity (2022) 117:1–272
Fungal Diversity (2022) 117:1–272
171
◂ Fig. 136 Maximum likelihood phylogenetic tree based on nrITS
sequences of Lepiota species. Bootstrap support ≥ 70% is indicated at
the nodes. New sequences from this study are in blue. The GenBank
accession numbers are indicated after the species name. Abbreviation
L = Lepiota, M = Macrolepiota. The tree is rooted in Macrolepiota
orientiexcoriata
smaller basidiospore size. However, it seems that the size of
basidiospores is not much different in all specimens found in
China, Laos, Sri Lanka and Thailand; and the morphology
and size can be minorly different in different specimens.
According to the analysis of nrITS sequence data (Fig. 136),
Lao specimens are identical to specimens from China with
high (100%) bootstrap support.
Lepiota thrombophora from Thailand is most similar to
L. metulispora in morphology, but differ in smaller basidiospores size (10–14 × 3–5 μm) (Hyde et al. 2021); and the
type specimen of L. thrombophora from Sri Lanka is different in dark brown squamules on pileus and shorter elements of pileus covering (25–100 × 5–15 µm). Liang et al.
(2011) described L. thrombophora from China, based on its
longer elements (up to 330 µm long). The analysis of nrITS
sequence data showed that Chinese and Thai specimens of
L. thrombophora are identical, and related to L. metulispora (Fig. 136). The type specimen of L. attenuata from
China is closer to L. metulispora in morphology but differs
in much longer elements on pileus covering (231 µm long),
and basidiospores are more attenuated at the apex (Liang
et al. 2011). Thai specimens of L. attenuata from Thailand
have longer and typically attenuate elements (300 µm long)
(Hyde et al. 2021). Other similar species to L. metulispora
were discussed in Sysouphanthong et al. (2012).
Lepiota pongduadensis Sysou., Hyde & Vellinga in Sysouphanthong et al., Cryptog. Mycol. 33(1): 37 (2012).
Index Fungorum number: IF519961; Facesoffungi number: FoF09886 Figs. 137, 138
Fig. 138 Lepiota pongduadensis (HNL503131, new record). a
Basidiospores b Cheilocystidia c Pileus covering
Pileus 30–45 mm, campanulate, expanding to convex
or umbonate with small umbo, applanate with low umbo,
straight margin; glabrous to rough at umbo, brown to dark
brown (6E5-8, 7F7-8), with light brown to brown (7D68, 7E7-8) glabrous around umbo, later surface broken in
radial streaks from around umbo towards the margin, with
concolorous tomentose to crowded squamules towards the
margin, on white to yellowish-white (4A2) black ground;
marginal zone broken, split, fringed, cortinate with write
bibrils and light brown to brown (7D6-8, 7E7-8) partial veil
remnants. Lamellae free, slightly crowded, ventricose to
broadly, 3–5 mm wide, with 2 length lamellulae, white to
yellowish-white (4A2), with concolorous eroded edge. Stipe
35–45 × 4–6 mm, cylindrical, fibrillose or cortinate at middle
zone, then with light brown to brown (7D6-8, 7E7-8) squamules downwards base, with white to yellowish-white (4A2)
Fig. 137 Lepiota pongduadensis in habitat. a-b HNL503131 c
HNL503150
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172
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Fig. 139 Lepiota subthailandica in habitat. a-b MFLU
09–0166 c-d MFLU 10–0616
(holotype)
Fig. 140 Lepiota subthailandica (MFLU 10-0616, holotype). a
Basidiospores b Basidia c Cheilocystidia d Pileus covering
black ground from middle to apex, with reddish-white (7A2)
below middle zone downwards base. Annulus an annular
zone, cortinate with white fibrils. Context white and up to
3 mm wide in pileus; hollow and concolorous with surface.
Smell and taste unknown. Spore print white.
Basidiospores [50,2,2] 11.5–16.5 × 4–5.5 µm,
avl × avw = 13.4 × 4.6 µm, Q = 2.8–3, Qav = 2.91, in sideview cylindrical amygdaliform, with attenuate apex, with
straight abaxial side, with hilar appendage, with superhilar
depression, in frontal view fusiform, hyaline, slightly thickwalled, dextrinoid, congophilous. Basidia 16–26 × 7–9 µm,
13
clavate, hyaline, thick-walled, 4-spored. Cheilocystidia
abundant, 27–35 × 6.5–15 µm, mostly fusiform or clavate,
sometimes utriform, thick-walled, hyaline. Pileus covering
a trichoderm made up of narrowly cylindrical elements, normally wider at middle and narrow to apex, with attenuate
apex 70–400 × 6–13.5 µm, thick-walled, with brown parietal
and intracellular pigment, smooth, sometimes incrusted at
base of element and hyphae. Stipe covering of squamules a
trichoderm similar to pileus covering. Clamp-connections
present.
Material examined: Laos, Oudomxay Province, Xay
District, Houay Houm Village, N 20° 32′ 00.67″, E 101°
53′ 48. 17.16″. 917 m., 10 July 2014, P. Sysouphanthong,
PS2014-1460 (HNL503131); ibidem, 12 August 2014, P.
Sysouphanthong, PS2014-1479 (HNL503150).
Habitat and distribution: Growing solitary to a small
group; saprotrophic and terrestrial on humus soil; originally
described from northern Thailand (Sysouphanthong et al.
2012). This is the first report of L. pongduadensis in Laos.
GenBank numbers: HNL503131–MT436259 (ITS).
HNL503150–MT436260 (ITS).
Notes: Lepiota pongduadensis is a new record from Laos;
two Lao specimens were collected from Oudomxay Province
of northern Laos, and they show similar morphology and
nrITS sequences to the type specimen (Fig. 136). Lepiota
pongduadensis was originally described from Chiang Mai
and Chiang Rai Provinces of northern Thailand. The species
is placed in the section Lepiota by Sysouphanthong et al.
(2012) and is distinguished from other species in the section. Only a few species are similar to L. pongduadensis
(Sysouphanthong et al. 2012). Lepiota attenuata, the type
specimens from China, is similar to L. pongduadensis in
morphology, but different in lacking a dark brown surface
on umbo, lighter colour of squamules on pileus, and stipe
covering (brownish-yellow to yellowish-brown), larger
Fungal Diversity (2022) 117:1–272
basidiospores (14.5–19 × 4–5.5 mm), shorter elements of
pileus covering (80–231 × 3.8–13 mm) (Liang et al. 2011).
Lepiota subthailandica Sysouph., K.D. Hyde & Thongkl.,
sp. nov.
MycoBank no: MB839987, Facesoffungi number:
FoF09889; Figs. 139, 140
Etymology: the morphological characteristics of this species are similar to L. thailandica.
Holotype: MFLU 10–0616
Diagnosis: similar to L. thailandica in morphology, but
the difference in larger basidiomata, lacking of utriform or
fusiform cheilocystidia and nrITS sequences.
Pileus 10–16 mm diam., convex to umbonate, expanding to plano-concave, with straight margin; rough or with
crowded squamules at center, light brown to brown (7D6-8,
7E7-8) at center, with concolorous squamules around umbo
towards margin, slightly distant at marginal zone on white
to orange-white (5A2) background; margin broken, sulcate
or striate, appendicular, with concolorous squamules on the
surface and white fibrillose remnants. Lamellae free, broadly
173
ventricose, 3–4 mm wide, white, moderately crowded, with 1
length lamellulae, with eroded edge. Stipe 25–30 × 3–4 mm,
cylindrical, slightly wider at base; completely fibrillose,
crowded at annular zone down toward base, white, with light
brown to brown (7D6-8, 7E7-8) squamules at base zone, on
white to orange-white (5A2) background. Annulus with an
annular zone or cortinate with white fibrils. Context white in
pileus, up to 1 mm wide; hollow in stipe, concolorous with
surface. Taste and smell unknown. Spore print white.
Basidiospores [50,2,2] 5–6 × 3.2–4 µm,
avl × avw = 5.54 × 3.64 µm, Q = 1.45–1.57, Qav = 1.52,
in side-view ellipsoid ovoid, in frontal view ovoid to,
slightly thick-walled, dextrinoid, congophilous, cyanophilous, not metachromatic. Basidia 15–18 × 5–7 µm, clavate,
slightly thick-walled, hyaline, 4-spored. Cheilocystidia
18.5–29 × 5–10 µm, narrowly clavate to clavate, rarely
broadly clavate, slightly thick-walled, hyaline. Pleurocystidia absent. Pileus covering a trichoderm made up of two
layer of elements; upper layer made up of cylindrical to
narrowly cylindrical elements with rounded apex, sometimes swollen at middle and tapering to base and apex,
Fig. 141 Lepiota subvenenata
in habitat. a-e HNL503121
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174
Fig. 142 Lepiota subvenenata (HNL503121, new record). a Basidiospores b Basidia c Cheilocystidia d Pileus covering
60–140 × 9–17.5 µm, with pale brown parietal and intracellular pigment; underpayer made up of shortly clavate elements, 30–45.0 × 10.0–20 µm, with pale brown parietal and
intracellular pigment. Stipe covering of squamules at base
zone similar to pileus covering. Clamp connections present
in all tissues.
Material examined: Thailand, Chiang Mai Province:
Mae Taeng district, Pha Deng village, N 19° 07.13′, E 98°
43.52′, 905 m, 04 July 2010, P. Sysouphanthong P98 (MFLU
09–0616, holotype); ibidem, 25 July 2008, P. Sysouphanthong PS093 (MFLU 09–0166, paratype).
Habitat and distribution: solitary, saprotrophic, on
decayed humus soil; found in high elevation deciduous forests of northern Thailand.
GenBank numbers: MFLU 09–0616–MT436254 (ITS).
MFLU 09–0166–MT436253 (ITS).
Note: Lepiota subthailandica has a tiny basidioma, a
trichodermal structure of pileus covering and ellipsoid ovoid
basidiospores; and the species is located in Lepiota sect.
Ovidsporae (J.E. Lange) Kühner (Vellingar 2001). In the
same section, Lepiota subthailandica is very similar to L.
thailandica Sysouph., K.D. Hyde, J.C. Xu & P.E. in morphology; and they are widespread in the same location of
Chiang Mai, northern Thailand. However, L. thailandica
has smaller basidiomata (3–4 mm diam. in pileus), utriform
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Fungal Diversity (2022) 117:1–272
and fusiform cheilocystidia, and shorter elements of pileus
covering (55–95 × 5.5–22 µm) (Sysouphanthong et al. 2016).
The second species, L. microcarpa Sysouph., K.D. Hyde &
Vellinga, is similar in micromorphology with L. subthailandica. However, L. microcarpa has penguin-shaped basidiospores, and belongs to sect. Lepiota (Sysouphanthong et al.
2012). Based on the nrITS sequences analysis, two samples of L. subthailandica are not related to L. thailandica,
and are separated from other species in the sect. Ovisporae
(Fig. 136).
Lepiota subvenenata Hai J. Li, Y.Z. Zhang & C.Y. Sun.
Index Fungorum number: not found; Facesoffungi number: FoF09888; Figs. 141, 142
Pileus 30–50 mm diam., first subglobose, expanding
to parabolic to convex or umbonate, plano-convex when
mature, with straight margin; when young glabrous to
granulose, completely light brown to brown (7D7-8), when
mature surface breaking and leaving concolorous glabrous
or granulose umbo, with concolorous squamules around
umbo towards margin, on white to orange-white (5A2) background; margin with concolorous squamules and white partial veil remnants. Lamellae free, broadly ventricose, with
3 length lamellulae, up to 4 mm wide, white, crowded, with
white eroded edge. Stipe 40–70 × 5–7 mm, cylindrical, with
white fibrillose and light brown to brown (7D7-8) squamules
at annular zone, with concolorous squamules under annular
zone downwards base, on white to orange-white (5A2) background. Annulus an annular zone, with white fibrillose and
concolorous squamules. Context in pileus white, 3–5 mm
wide; in stipe hollow, concolorous with surface. Taste and
smell unknown. Spore print white.
Basidiospores [50,1,1] 4.5–6.3 × 2.5–3.5 µm,
avl × avw = 5.02 × 3 µm, Q = 1.7–1.8, Qav = 1.75, in
side-view oblong ovoid, in frontal view oblong, slightly
thick-walled, hyaline, dextrinoid, congophilous. Basidia
18–23 × 7–10 µm, clavate, slightly thick-walled, hyaline,
4-spored, rarely 2-spored. Cheilocystidia 13–25 × 6–10 µm,
cylindrical to irregular cylindrical, narrowly clavate to
clavate, sometimes fusiform or utriform, slightly thickwalled, hyaline. Pleurocystidia absent. Pileus covering a
trichoderm made up of cylindrical elements with rounded or
attenuate apex, 60–170 × 7–13 µm, thick-walled, with parietal pale brown and intracellular pigment. Stipe covering of
squamules similar to pileus covering. Clamp connections
present.
Material examined: Laos, Oudomxay Province, Xay District, Houay Houm Village, N 20° 32′ 00.67″, E 101° 53′ 48.
17.16″. 917 m., 25 July 2014, P. Sysouphanthong, PS20141450 (HNL503121).
Habitat and distribution: Growing in a small group, on
humus soil mixed with dead leaves; known from Yunnan
province of Southwest China (Zhang et al. 2019). We report
the first record of L. subvenenata from Laos in this study.
Fungal Diversity (2022) 117:1–272
175
Fig. 143 a-c Field symptoms of
sclerotial wilt under field conditions. d-f Collar region showing
fungal pathogen colonizing the
stem soil interface. g–i Sclerotia
infecting leaves
GenBank numbers: HNL503121–MT436258 (ITS).
Notes: Lepiota subvenenata was described from Yunnan
province of Southwest China. It is closely related to L. venenata Z. H. Chen & Zhu L. Yang in morphology. Lao specimens clustered with the type specimens in the nrITS phylogenetic analyses and share similar morphology (Fig. 136).
Atheliales Jülich.
Atheliales is an order mostly composed of corticioid
fungi (Sulistyo et al. 2021). Based on phylogenetic analyses
Sulistyo et al. (2021) accepted five families: Atheliaceae,
Byssocorticiaceae, Lobuliciaceae, Pilodermataceae, and
Tylosporaceae in this order. However, Wijayawardene et al.
(2022) accepted only Atheliaceae and Lobuliciaceae in this
order.
Atheliaceae Jülich, Biblthca Mycol. 85: 355 (1982) [1981].
This is the type family of Atheliales. Jülich (1982)
included the genera Athelopsis, Caerulicium, Confertobasidium, Leptosporomyces, and Luellia in Atheliaceae.
Wijayawardene et al. (2022) accepted 20 genera in this family. Atheliaceae mainly consists with saprotrophic taxa, with
one lichenicolous species (Athelia arachnoidea) (Sulistyo
et al. 2021).
Athelia Pers., Traité champ. Comest. (Paris): 57 (1818).
Athelia is a genus of corticioid fungi. Some species are
facultative parasites of plants and of lichens (Sulistyo et al.
2021). A species of Athelia also engaged in a symbiotic relationship with termites (Reticulitermes), in which the fungus
forms sclerotia that mimic termite eggs and worker termites
13
176
Fig. 144 a Stem part showing the mycelial cords colonized on the
surface. b Sclerotia observed under stereo zoom microscope. c-d Pure
cultures of Athelia rolfsii on PDA medium. e–f Pathogenicity test performed under greenhouse showing the color rot symptoms
handling the sclerotia as if they were eggs (Matsuura et al.
2005). Wijayawardene et al. (2022) accepted 32 species in
this genus.
Athelia rolfsii (Curzi) C.C. Tu & Kimbr., Botanical
Gazette Crawfordsville 139: 460 (1978).
Index Fungorum number: IF309351; Faceoffungi number
FoF13394; Figs. 143, 144
Pathogenic on roots and stem of Lablab purpureus. Sexual morph: not observed. Asexual morph: numerous globoid sclerotial bodies were developed on the collar region of
infected Lablab purpureus plants at the stem soil interface.
Sclerotia measured 1–3 mm in diam. (n = 50), initially whitish and later turned to brownish upon maturity.
Cultural characteristics: On PDA, reaching 90 mm at
28 °C after 7 d in 12/12 h light/dark, dense white fluffy
colonies developed rapidly over the culture plates and produced characteristic sclerotia near the edges of plates and
on the upper lid surface. After, 12–15 days of incubation
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Fungal Diversity (2022) 117:1–272
sclerotia were recorded per plate ranging to 180 – 490 (mean
358 ± 24, n = 20). Colonies on PDA reaching 90 mm at 28 °C
after 14 d in 12/12 dark, colonies appeared white to pink
with abundant aerial mycelium.
Material examined: India, Karnataka, Mysore, Doddamaragowdanahally, on infected plants of Lablab purpureus (L.) Sweet (Fabaceae), 17 April 2013, S. Mahadevakumar (UOM-Sr-13–4), living culture Sr-LP3.
Habitat: Wide range of hosts (Farr and Rossman 2022),
Lablab purpureus (This study).
Distribution: Brazil, China, Cuba, Fiji, New Zealand,
Panama, Papua New Guinea, Phillippines, South Africa,
Spain, Sri Lanka, West Indies and USA (Farr and Rossman
2022), India (This study).
GenBank number: KJ002765 (ITS).
Notes: The characteristic foot-rot and leaf blight disease of Lablab purpureus are reported in the present study
and the diagnostic features are presented. The disease was
observed at all stages of plant growth. In the early infection stage, the collar rot affected seedlings collapsed. Initial symptoms appeared as tan water-soaked lesions usually
near the stem-soil interface, with lesions that enlarged and
expanded towards the shoot apex, causing rotting. Similarly,
lesions expanded towards the roots causing root decay and
death of the host plant. The pathogen produced numerous
globoid sclerotial bodies over the surface of the host plants.
The cabbage heads' rotting was recorded with sclerotia
development over the head surface. The disease was most
prevalent during the rainy season. In dry weather, infected
tissues showed the presence of mycelial threads with very
few hard, melanized sclerotial bodies. In USDA host fungal
database, the record 51387 specifies a geographical record
for India, which shows the sequences submitted by our group
to GenBank in 2013 and referred by Paul et al. (2017), a new
geographical record for Athelia rolfsii on Ipomoea batatas
from Korea. Based on the morphological, and cultural characteristics and ITS-rDNA analysis, the fungal pathogen was
identified as Athelia rolfsii (Fig. 145). This is the first report
on the association of A. rolfsii causing root rot and leaf
blight disease in L. purpureus from India and worldwide.
Hymenochaetales Oberw.
The concept of hymenochaetaceous fungi was initiated
by Patouillard (1900) in Série des Igniaires, which includes
xanthochoroid polypores with simple, clampless hyphae
and setal elements. Later, Oberwinkler (1977) raised the
order Hymenochaetales, based on the characteristics of the
Hymenochaetoidae, Hymenochaetaceae (proposed earlier by
Donk (1948)). Hymenochaetales comprises with homobasidiomycetes with diverse basidiomatal characters, annual to
perennial, resupinate to pileate, stipitate to spathulate with
smooth, poroid or hydnoid hymenophore. Microscopically,
simple, clampless, mono to di or trimitic hyphal system with
Fungal Diversity (2022) 117:1–272
177
Fig. 145 Phylogenetic tree for species of Athelia generated from
maximum likelihood (RAxML) based on ITS gene regions. The Maximum likelihood bootstrap value ≥ 50% are given at the nodes. The
newly generated sequences are in blue. The tree is rooted to Rhizoctonia solani (FX1 AG4.HG.1)
smooth to ornamented, thin to thick-walled, hyaline to coloured, globose to cylindrical basidiospores are characteristic
features. The presence or absence of sterile elements such as
cystidia, cystidioles or seate also plays a vital role in identification (Fiasson and Niemela 1984).
MycoBank recorded 19 associated families (http://www.
mycobank.org) whereas 1530 taxa were submitted in GenBank (https:// www. ncbi. nlm. nih. gov) (as of 17 January
2022). Wijayawardene et al. (2022) accepted six families
in this order.
surface, simple mono to dimitic hyphal system, presence or
absence of setae, thin to thick-walled, hyaline or coloured
basidiospores (Ryvarden 1991; Dai 2010). Sharma (1995)
reported 16 genera and 91 species of Hymenochaetaceae
under Indian Aphyllophorales. Hymenochaetaceae is one
of the largest families in Basidiomycota (Kirk et al. 2008).
Dai (2010) mentioned Phellinus sensu lato, Inonotus sensu
lato and Hymenochaete as the three largest genera of Hymenochaetaceae. There are 71 associated genera documented
in MycoBank (http://www.mycobank.org) and 1,103 taxa
submitted in GenBank (https://www.ncbi.nlm.nih.gov) (as
of 1 January 2022). Wijayawardene et al. (2022) accepted
42 genera in this family.
Coltricia Gray, Nat. Arr. Brit. Pl. (London) 1: 644 (1821).
The genus is typified by C. perennis (L.) Murrill. Species of this genus are cosmopolitan and have few species that have been found to be associated with plant
roots (Tedersoo et al. 2007). Coltricia is characterized
by poroid and stipitate basidiocarps, a monomitic hyphal
system lacking clamp connections and coloured, ellipsoid
Hymenochaetaceae Donk, Bull. Bot. Gdns Buitenz. 17(4):
474 (1948).
Donk (1948) classified the xanthochroid Aphyllophorales
under Hymenochaetaceae which comprises wood-decaying
white-rot fungi and medicinal fungi with styrylpyrone pigments (which are responsible for a positive xanthochoric
reaction) (Dai et al. 2007, 2009). Hymenochaetaceae are
described by yellow to deep brown, resupinate to pileate
or stipitate basidiomata with smooth to hydnoid hymenial
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178
Fungal Diversity (2022) 117:1–272
Fig. 146 Coltricia insularis. a-c
Basidiocarps. d Basidiospores.
e, f Pileipellis. g Hymenial
trama, longitudinal section.
h Stipitipellis. All from LIP
0,401,817 (holotype, pictures
by D. Borgarino (b) and P.-A.
Moreau) except a (Hal-BP-72,
sequence MT594499, picture by
A. Rinaldi) and c (Hal-BP-19,
sequence MT594498, picture by
A. Rinaldi). Scale bars = 5 mm
(a-c), 10 µm (d-g)
to subglobose, smooth basidiospores (Dai 2010). Wijayawardene et al. (2022) accepted 40 species in this genus.
Coltricia insularis P.-A. Moreau, Bellanger, Loizides & A.
Rinaldi, sp. nov.
Index Fungorum number IF900072; Faceoffungi number FoF13395; Fig. 146
Etymology. Insularis = of islands, a Latin adjective
referring to Corsica, Cyprus and Sardinia, three Mediterranean islands in which the species was discovered.
Holotype: LIP 0401817
Pileus 15–60 mm, irregularly infundibuliform, rough,
sometimes weakly lobed, subtomentose or more distinctly
tomentose to subhirsute towards the centre, radially corrugated or furrowed and usually with well-defined concentric
13
chromatic zones; colours ranging from brown, red-brown,
grey-brown, purple, umber or ochraceous; margin often paler
and somewhat undulating. Hymenial surface poroid, comprised of adnexed to irregularly decurrent, shallow, angular,
or somewhat elongated pores, mostly 2–3 per mm, becoming
poorly defined towards the margin and sometimes fusing to
form sterile tomentose patches or a sterile marginal zone;
warm grey to buff or ochraceous. Stipe 20–40 × 3–8 mm,
usually irregularly furrowed, densely tomentose, buff or pale
brown at the apex but quickly staining ochraceous or brown,
darker chestnut to red-brown or orange-brown lower, deeply
submerged into the substrate and often with sand and litter
tightly adhered to it. Context corky, thick and indistinctly
zonate in the pileus, darker in the stipe, chestnut to redbrown or umber.
Fungal Diversity (2022) 117:1–272
179
Fig. 147 Phylogenetic analyses were conducted online at www.phylo ▸
geny.fr (Dereeper et al. 2008). Multiple sequence alignments were
performed with MUSCLE v. 3.7 (Edgar 2004). Maximum likelihood
(ML) phylogenetic analysis was achieved with PhyML v. 3.0 (Guindon et al. 2010), using the GTR + I + Γ model of evolution and the
Shimodaira Hasegawa version of the approximate likelihood-ratio test
(SH-aLRT) of branch support (Anisimova et al. 2011). Phylogram
was built using TreeDyn 198.3 (Chevenet et al. 2006) and edited with
Inkscape 0.91 (https://inkscape.org/fr). Newly generated sequences
for this study are in bold. The tree is rooted by mid-point rooting
method under PhyML (Guindon et al. 2010)
Spores (6.8) 7.5–9 (10.2) × 3.2–3.9 (4.1) µm, ellipsoid to
broadly ellipsoid or ovoid when immature, cylindrical to
fusiform when projected, with a light supra-apicular depression, pale yellow in KOH and in Melzer’s, wall < 0.3 µm
thick, smooth. Basidia 12–19 × 7–7.5 µm, 4-spored (partly
2-spored on immature specimens) with spindle-shaped
straight sterigmata, shortly cylindrical, hyaline. Hymenophoral setae absent. Edges sterile, made of bunches of slender
thin-walled hairs 2.3–3 µm wide, encrusted with yellowish granular deposits. Pileipellis 60–80 µm-thick, convoluted trichocutis, made of generative hyphae 3.5–6(7) µm
wide, terminal elements 45–80 µm long, branching often
with right-angled furcations, rounded to mucronate at the
apex. Stipitipellis a trichocutis made of flexuose, mostly
unbranched skeletoid hairs, 70–180 × 3.5–5.5 µm, with occasional secondary septa; wall smooth or with few hyaline
mucoid deposits towards the apex, 0.8–1.2 µm thick, yellow
in 5% KOH; apex rounded. Clamps absent from all observed
septa. Smell weak, faintly acidic.
Habitat and Distribution: Associated with Cistaceae
(Cistus spp., Halimium halimifolium) shrubs in dry, sandy
places. So far collected from Cyprus, France (Corsica), Italy
(Sardinia) and Spain (Andalucia), but probably widespread
in xerothermic localities throughout the Mediterranean
basin.
Material examined: France: Corsica (Haute-Corse),
Monaccia d’Aullène, Réserve naturelle du Mucchiu Biancu,
25 Nov. 2006, D. Borgarino, L. Hugot, C. Lavoise, P.-A.
Moreau & F. Richard, PAM06112616 (LIP 0401817,
holotype).
Other material examined: Cyprus: Trimiklini, M.
Loizides, 28 Feb. 2019, ML91292CO (LIP 0401819). Italy:
Sardinia, Gonnesa, A. Rinaldi, 5 Dec. 2019, Hal-BP-135,
LIP 0401740. Spain: Andalucia, Huelva, Almonaster-laReal, under Pinus halepensis and Cistus monspeliensis in a
semi-open heathland, A. Gasch Illescas & P.-A. Moreau, 28
Dec. 2019, PAM1912814 (LIP 0401740).
Notes: Coltricia insularis is a xerophilic Mediterranean species characteristic of Cistaceae shrublands,
found especially among sands, where it may grow in dense
clusters adhered to the bases of rockroses. Also known
from the thermo-mediterranean zone in Cyprus and the
13
180
supramediterranean zone in Andalucia, where a slenderer
form of this species was collected on sandy ground, at the
edge of a pine forest. In Sardinia, it occurs in pure Halimium stands in coastal areas. However, in places where more
potential hosts are present, it might be part of the extensive ectomycorrhizal networks that are common in several
Mediterranean ecological settings (Taudiere et al. 2015;
Leonardi et al. 2020). From what is known about Coltricia mycorrhizal biology, members of the genus appear to
establish ectomycorrhizal associations with a range of hosts
(Rinaldi et al. 2008). Coltricia insularis phylogenetically
belongs to the group of Coltricia perennis, which has not
Fig. 148 Fulvifomes jawadhuvensis (MUBL4011, holotype).
a-b Basidiocarp. c Stratified
tube layer. d Tramal hyphae. e
Context hyphae. f Basidioles.
g Basidia. h Basidiospore i
Basidiospore in H2O. j Basidiospore in KOH. k Basidiospore
in cotton blue. Scale: a = 3 cm.
c = 1 cm. d–k = 10 µm
13
Fungal Diversity (2022) 117:1–272
been the object of monographic revision since the advent of
DNA studies. The phylogenetic analysis (Fig. 147) shows
the unexpected diversity unravelled in this lineage, in which
at least three main subclades can be identified among the
available sequences in GenBank and UNITE. Most of these
are devoid of Linnaean names and the need of typification of
C. perennis itself (type of the genus) is obvious, considering
this name has been arbitrarily applied to nearly all sequences
available in the’Perennis-clade’. Coltricia montagnei Fr. (in
Montagne 1836, p. 341), originally described from Northern
France (Ardennes, near Sedan), has been variously interpreted and was only recently described in detail (Rivoire
Fungal Diversity (2022) 117:1–272
2020). Coltricia confluens Keizer (1997, p. 389), from which
the isotype collection Keizer 93060 was kindly provided by
the author, but unfortunately, could not be sequenced. Coltricia insularis differs from all currently recognized European
species by spore dimensions and shape: mature specimens
display spores with an average Q of around 2.3 and a typical fusiform profile. Like C. perennis and C. confluens (but
not C. montagnei), the hyphae of pileipellis are frequently
T-branched, but the taxonomic importance of this feature
requires further observations. Coltricia’confluens’ and a
sister Mexican species (Fig. 147), represent an independent
well-supported clade. Amongst the subclade 3 (Fig. 147),
C. insularis has an American sister species (sequences ITS
MH211968 and MK966429 from pine forests in Oregon);
both sequences form a well-supported subclade, distinct
from the two other subclades of Clade 3 representing apparently circumboreal North European, North American and
Asian species. None of the collections available in these
clades suggests a thermophilic or xerophilic Mediterranean origin. A detailed revision of collections representing
each clade is required before a thorough comparison of this
cluster of species with C. insularis and within the whole
“Perennis-clade” can be made.
Fulvifomes Murrill, Northern Polypores (5): 49 (1914).
Fulvifomes (typified as Fulvifomes robiniae Murrill)
was described by Murrill (1914) to include perennial,
sessile, ungulate, woody xanthochoroid fungus with dimitic hyphal system, colored basidiospores and lacks setae.
Earlier Fulvifomes was treated as a synonym of Phellinus Quél. (Ryvarden and Johansen 1980; Dai 1999) later,
many validated the generic rank of Fulvifomes, by studying
the macro-microscopical description, nuclear behaviour
and molecular data (Fiasson and Niemelä 1984; Wagner
and Fischer 2002). Fulvifomes was revaluated and extensive ranges of characteristics were added to the classical
description which includes annual to perennial, resupinate to pileate, applanate to ungulate, smooth to rimose
basidiomata, homogenous to duplex context and poroid
hymenial layer with mono-dimitic to dimitic hyphal system, presence or absence of setae, cystidioles, smooth,
thick-walled, colored, globose to ellipsoid, cyanophilic to
acyanophilic and inamyloid basidiospores (Murrill 1914;
Wagner and Fischer 2002; Dai 2010; Salvador-Montoya
et al. 2018; Tchoumi et al. 2020). Salvador-Montoya et al.
(2018) and Zheng et al. (2021) proposed keys to American
and Chinese Fulvifomes spp., respectively, since few samplings were done in other parts of the world.
Approximately 331 sequences belonging to 50 taxa were
submitted in NCBI (https://www.ncbi.nlm.nih.gov) and 65
associated taxa are available in MycoBank (http:// www.
mycobank.org) (as of 1 January 2022). Wijayawardene et al.
(2022) accepted 33 species in this genus.
181
Fulvifomes jawadhuvensis Kezo, K., Gunaseelan, S., &
Kaliyaperumal, M., sp. nov.
Index Fungorum: IF558179; Facesoffungi Number:
FoF10745; Fig. 148
Etymology: The species epithet jawadhuvensis refers to
the type locality of basidiomata collection.
Holotype: MUBL4011
Basidiocarps perennial, solitary, pileate, sessile, light in
weight, hard when dry. Pileus dimidiate, convex to meagrely
ungulate, with no distinct crust, projecting up to 5.7 cm,
9.5 cm wide and 3.4 cm thick near the base. Pilear surface
velvety, light brown (6D6) to rust-brown (6E8) and meagerly warted when young, on maturity pilei becoming rough,
weakly rimose, concentrically, narrowly sulcate, weakly
zonate. Margin entire, round to obtuse, brown (6E4) to dark
brown (6F5), velutinate when young, developing into brownish grey (6F2) to greyish brown (6E3), glabrous on maturity.
Pore surface raw umber brown (5F8) to brown (6E7). Pores
round to angular, regular, 4–8 per mm. Dissepiments entire,
thick. Context up to 2.7 cm, homogeneous, yellowish-brown
(5E8) to dark brown (6F7). Tubes yellowish-brown (5D8)
to light brown (6D6), up to 0.7 cm thick, tube layers stratified, each stratum 0.2 to 0.4 cm with a thin layer of context
in between.
Hyphal system strictly dimitic, Generative hyphae
dominant; both skeletal and generative hyphae acyanophilous; tissue darkening with KOH without swelling.
Context Generative hyphae, thin to thick-walled, hyaline
to yellow, simple septate, branched, 2–6.5 μm diam.; skeletal hyphae, thick-walled with narrow lumen, unbranched,
yellowish-brown, aseptate, 2–5.7 μm diam. Trama Generative hyphae, thin to thick-walled, yellow to brown,
septate, rarely branched, 2–5.2 μm dia.; skeletal hyphae,
thick-walled with narrow to wide lumen, yellowish-brown,
aseptate, unbranched, 2–5.2 μm dia. Setae, cystidioles
absent. Basidioles dominant, clavate, 7.7–14 × 3.8–7.2 μm.
Basidia clavate to broadly clavate, with four sterigmata,
8.5–15 × 5–8.5 μm. Basidiospores broadly ellipsoid to subglobose, thick-walled, smooth, yellow in water, turning
rust-brown in KOH, (4.8–) 5.1–6.4 (–6.9) × (4.1–) 4.4–4.9
(–5.2) μm (n = 50/2), Q = 1.1–1.3, CB ̄, IKI ̄. Chlamydospores globose to subglobose, thick-walled rust brown to
reddish-brown, 5.2 – 8.5 × 4.1–7 μm, CB ̄, IKI ̄.
Specimen examined: India, Tamil Nadu, Thiruvannamalai district, Jawadhu hills, Jamunamarathur, 12.64° 54′
19.1″ N 79° 18′ 33″ E, on living angiosperm tree (Albizia amara (Roxb.) Boiv., Fabaceae), 09 February 2018,
Kezhocuyi Kezo (MUBL4011, holotype).
GenBank numbers: MW040079 (ITS), MW048886
(LSU), MW690924 (tef1).
Notes: Fulvifomes jawadhuvensis shares similarities with F. grenadensis by having dimidiate to ungulate
pileus, round to obtuse margin, absences of cystidioles, but
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Fungal Diversity (2022) 117:1–272
Fig. 149 Fulvifomes malaiyanurensis (MLCASB020,
holotype). a–b Basidiocarp. c
Stratified tube layer. d Tramal
hyphae. e Context hyphae. f
Basidioles. g Basidia. h Basidiospore. i Basidiospore in H2O. j
Basidiospore in KOH. k Basidiospore in cotton blue. Scale:
a = 3 cm, c = 1 cm, d–k = 10 µm
the former lacks distinct crust after velvety pileus wears
off and no. of pores per mm. Larger broadly ellipsoid to
subglobose spores in F. jawadhuvensis differ from smaller
spores in F. grenadensis (4–6 × 3–4 µm) (Ryvarden 2004).
The new Indian species share other common characteristics with F. elaeodendri, F. hainanensis, F. thailandicus,
in having dimidiate to ungulate basidiomata, zonate pileus
with rimose patten (except F. hainanensis), pores per mm,
distinctly thick-walled, colored basidiospores, absence
of setae, but significantly varies basidiospores size and
absences of cystidioles (Zhou 2014; Tchoumi et al. 2020).
Fulvifomes jawadhuvensis and F. centroamericanus share
only dimidiate pilei and absences of cystidioles and differ in other features; the former varies entirely from F.
13
krugiodendrii (Ji et al. 2017). Fulvifomes jawadhuvensis
shows variations with F. nonggangensis and F. tubogeneratus in basidiomata characters and microscopic illustrations (Zheng et al. 2021).
Fulvifomes malaiyanurensis Gunaseelan, S., Kezo, K. &
Kaliyaperumal, M., sp. nov.
MycoBank: MB821517; Facesof fungi number:
FoF10744; Fig. 149
Etymology: The species epithet ″malaiyanurensis″ refers
to the type locality of the collection site.
Holotype: CAL 1618
Basidiocarps perennial, solitary, sessile, broadly attached
to substrate, hard, light when dry. Pilei dimidiate, ungulate
Fungal Diversity (2022) 117:1–272
183
Fig. 150 Fulvifomes thiruvannamalaiensis (MUBL4013,
holotype). a Habitat. b Pilear
surface. c Pore surface. d Stratified layer (yellow arrows and
lines indicates each stratum). e
Contextual hyphae. f Tramal hyphae. g Cystidioles. h
Basidioles. i Basidia. j. Basidiospores. Scale bars: a–d = 1 cm,
e–j = 10 µm
to triquetrous, projecting up to 5 cm, 12 cm wide and 4 cm
thickness near the attachment. Pilear surface velutinate
towards the margin, partially, concentrically zonate, brown
(6E6) to greyish brown (6F3), cracked during collection.
Margin obtuse often round, velvety shining, brown (5E8) to
yellowish brown (5E4). Pore surface umber-brown (5E8).
Pores regular, circular to angular, 5–7 per mm. Dissepiments
thick, entire. Context yellowish-brown (5C7) to dark golden
brown (5C7), woody corky, up to 2 cm thick. Tube layer
brown (6D7), up to 2 cm long in distinctly stratified, individual layer up to 0.5 cm in length.
Hyphal system strictly dimitic, skeletal and generative
hyphae acyanophilous, tissue darkening with KOH without swelling. Context: Hyphal system subparallel, generative hyphae dominant hyaline to yellow, simple septate,
thin to thick-walled, 2.2–6.4 μm in diam, rarely septate
and branched, skeletal hyphae rare, yellow to brown, thickwalled, with a narrow to wide lumen, infrequently septate,
unbranched, 1.8–5.2 μm in dia. Trama Hyphae interwoven,
color not distinct from context; generative hyphae thin to
thick-walled, frequently septate 2–5.2 μm wide, skeletal
hyphae thick-walled, unbranched, aseptate with narrow to
wide lumen 2.7–5.2 μm. Hymenial setae, cystidia and cystidioles absent. Basidioles clavate, 5–16 × 3–7 μm. Basidia
broadly clavate, with four sterigmata, 7–16 × 5–8 μm. Basidispores broadly ellipsoid to subglobose, smooth, yellow
to golden yellow in water, turning golden brown to rustbrown in KOH, thick-walled, (4.2–) 4.6–5.4 (–5.7) × (3.9–)
4.2 – 4.9 (–5.2) μm, Q = 1.09, (n = 50/2), Q = 1.05–1.25, CB
̄, IKI .̄ Chlamydospores globose to subglobose, thick-walled,
rust-brown to reddish-brown, 5.7– 12.8 × 5.2–11.6 μm, CB
̄, IKI ̄.
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184
Material examined: India, Tamil Nadu, Vizhupuram district, Malaiyanur, 37° 25′ 19.1″ N 23° 36′ 33″ W, on living
angiosperm tree (Tamarindus indica L., Fabaceae), 25 October 2016, Sugantha Gunaseelan, MLCASB020, holotype.
Additional specimen examined: India, Tamil Nadu,
Vizhupuram district, Malaiyanur, 37° 25′19.1″ N 23°
36′33″ W, on living angiosperm tree (Tamarindus indica,
Fabaceae), 25 October 2016, Sugantha Gunaseelan,
(MLCASB021, Isotype).
GenBank numbers: MLCASB020-MF155651 (ITS),
MF155652 (LSU).
MLCASB021- MW048883 (LSU), MW690925 (tef1).
Notes: Fulvifomes malaiyanurensis is similar to F. thailandicus (Zhou 2015) in sharing yellowish-brown, broadly
attached basidiomata, pore, dimitic hyphal and absence of
cystidioles. However, F. malaiyanurensis is distinct by having ungulate to triquetrous, yellowish-brown, velutinate
basidiocarp, acyanophilic subglobose basidiospores, lacking
cystidioles. Fulvifomes malaiyanurensis differs from F. grenadensis (Ryvarden 2004), F. hainanensis (Zhou 2014) and
F. imbricatus (Zhou 2015) in pileus character, pore per mm,
shape and size of basidiospore. F. malaiyanurensis share a
similar pileial character with F. robinae (Salvador-Montoya
et al. 2018) but differs in hyphal system and basidiospore
but shares pores/mm. Fulvifomes malaiyanurensis shares
similarities with F. elaeodendri and F. yoroui in triquetrous
up to ungulate basidiomata and pores per mm (Tchoumi
et al. 2020; Olou et al. 2019) and differs in other characters.
Macroscopically, F. malaiyanurensis may resemble African
(Tchoumi et al. 2020), Asian (Zhou 2014, 2015; Liu et al.
2020; Du et al. 2021) and American (Ji et al. 2017) known
species but shows variation in other taxonomic characters.
Fulvifomes thiruvannamalaiensis Gunaseelan, S., Kezo, K.
and Kaliyaperumal, M., sp. nov.
Index Fungorum number: IF558486; Facesoffungi number: FoF10726; Fig. 150
Etymology: The species epithet ″thiruvannamalaiensis″
refers to the type locality of collection site.
Holotype: MUBL4013
Basidiocarp perennial, solitary, sessile, applanate,
broadly attached to the substrate by the narrow side, hard,
woody when dry. Pileus dimidiate, convex to meagrely ungulate, lacks crust, projecting up to 4.8 cm in length, 10.4 cm in
width and 3.4 cm thick near the attachment. Pilear surface
partly covered with microalgae, glabrous, light brown (6D8,
6E7), rust-brown (6E8) to dark brown (6F7), finely cracked
with small brownish grey scales (6E3, 6F2) concentric and
radially sulcate, but coarse and deep sulci, scrupose zones
near the attachment/older region, meagrely wavy near the
margin. Margin entire, round to obtuse, velutinate to smooth,
dark brown (6F7), brownish grey (6F2), often wavy. Pore
surface yellowish raw umber brown (5F8), yellowish-brown
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Fungal Diversity (2022) 117:1–272
(5E8) to brown (6E7, 6F8). Pores round, regular, 4–7 per
mm. Dissepiments entire, thick. Context up to 1.6 cm, homogeneous, fibrous to corky, brown (6E7, 6F8) to dark brown
(6F7). Tubes yellowish-brown (5E8), light brown (6D6) to
brown (6E7, 6F8), up to 1.3 cm thick, tube layers stratified
with thin-walled bright yellowish generative hyphae usually
running between the old tubes, each stratum up to 0.4 cm.
Hyphal system strictly dimitic, skeletal and generative
hyphae acyanophilous; tissue darkening with KOH without swelling. Context Generative hyphae dominant, thin to
thick-walled, simple septate, branched, hyaline to brown,
1.2–4.8 μm diam.; skeletal hyphae, thick-walled with
narrow to wide lumen, unbranched, aseptate, yellow to
brown, 2.4–4.3 μm diam. Trama Generative hyphae, thin
to thick-walled, septate, rarely branched, hyaline to brown,
1.2–4.8 μm diam.; skeletal hyphae, thick-walled with narrow
to wide lumen, aseptate, unbranched, brown, 2.4–4.8 μm
diam. Setae absent. Cystidioles thin-walled, hyaline, varies
in shape, fusoid to ventricose with elongated apical portion,
12.9–27 × 2.8–7.2 μm. Basidioles clavate to broadly clavate,
8.5–15.5 × 3.5–6.5 μm. Basidia clavate to broadly clavate,
with four sterigmata, 9.4–15.7 × 5.2–8 μm. Basidiospores
broadly ellipsoid to subglobose, thick-walled, smooth,
yellow in water, turning brown in KOH, (5.3–) 5.5–6.7
(–6.9) × (4.6–) 4.8–5.1(–5.5) μm (n = 30/2), Q = 1.05–1.3,
CB ̄, IKI ̄.
Material examined: India, Tamil Nadu, Thiruvannamalai district, Jawadhu hills, Jamunamarathur, 12.64° 54′
19.1″ N 79° 18′33″ E, on living angiosperm tree (Albizia
amara, Fabaceae), 09 February 2018, Sugantha Gunaseelan,
MUBL4013, holotype.
GenBank numbers: MZ221598 (ITS), MZ221600 (LSU).
Notes: Fulvifomes thiruvannamalaiensis, characterized by
perennial, solitary, dimidiate, applanate to ungulate basidiomes, significantly cracked pilear surface with brownishgrey scales, stratified tube layer with dimitc hyphal system
and thick-walled yellow to brown, acyanophilic subglobose
to ellipsoid basidiospores.
African species, F. yoroui (Olou et al. 2019) shows close
resemblance with F. thiruvannamalaiensis, in sharing perennial, pileate, ungulate basidiomata, dimitic hyphal system
and presence of fusoid cystidioles, but the size and shape
of the basidiospores (subglobose to globose basidiospores
5.5–6.5 × 4.7–5.6 μm), differ from F. thirvannamalaiensis.
Fulvifomes krugiodendri (Ji et al. 2017), F. rimosus (Hattori et al. 2014) and F. thailandicus (Zhou 2015) are closely
similar to F. thirvannamalaiensis morphologically by sharing dimidiate, concentrically sulcate, cracked basidiocarp,
dimitic hyphal system and presence of fusoid cystidioles, yet
differs in the number of pores and the size of basidiospores.
Despite few morphological resemblances between F.
elaeodendri (Tchoumi et al. 2020) such as sulcate, glabrous
pileus becoming cracked with age, stratified tube layer the
Fungal Diversity (2022) 117:1–272
185
Fig. 151 Phylogram of Fulvifomes species, obtained from
maximum likelihood (RAxML)
of combined ITS, LSU, and tef1
datasets. Bootstrap values (BS)
from maximum likelihood (ML,
left) and Maximum parsimony
(MP, middle) greater than
70% and Bayesian posterior
probabilities (PP), greater than
0.95, are indicated above the
nodes as ML/MP/BYPP. The
tree is rooted with Phylloporia
nodostipitata FLOR 51153.
New species and new records
are indicated in black bold
13
186
Fig. 152 Psilocybe keralensis (SDBR-CMUNK0448, new record).
a. Basidiomata. b. Basidiospores. c. Basidia. d. Cheilocystidia. Scale
bars: a = 10 mm; b–d = 5 μm
distinct microscopical traits viz., absence of cystidioles and
the size of the basidiospores are distinguishing F. thiruvannamalaiensis from F. elaeodendri. Fulvifomes centroamericanus (Ji et al. 2017), F. hainanensis (Zhou 2014) and F.
imbricatus (Zhou 2015) besides having dimitic hyphal systems are significantly distinct from F. thiruvannamalaiensis in morphology and microscopic features, by having
uncracked pilear surface, shape and size of basidiospores.
Fulvifomes thiruvannamalaiensis shares a few similar
morphological traits with the American species, F. cedrelae,
F. robinae and F. squamosus (Salvador-Montoya et al. 2018)
by having perennial, applanate, ungulate, sessile basidiomata with sulcate, cracked pilei, homogenous context and
stratified tubes, however, the former differs from the later in
hyphal system, number of pores and size of the basidiospore.
The Chinese species F. submerrillii (Liu et al. 2020), shows
high variations both morphologically and microscopically
from the newly described Indian F. thiruvannamalaiensis
(Fig. 151).
Hymenogastraceae Vittad, [as 'Hymenogastereae'], Monogr. Tuberac. (Milano): 11 (1831)
Note: Hymenogastraceae was erected by Vittadini (1893)
with contained gilled and false-truffle fungi. According to
the recent study by He et al. (2019), Hymenogastraceae is
one of the larger families in the Agaricales and comprises
10 genera, namely Anamika K.A. Thomas, Peintner, M.M.
Moser & Manim., Flammula (Fr.) P. Kumm., Galerina
Earle, Gymnopilus P. Karst., Hebeloma (Fr.) P. Kumm.,
Hymenogaster Vittad., Naucoria (Fr.) P. Kumm., Phaeocollybia R. Heim, Psathyloma Soop, J.A. Cooper & Dima and
Psilocybe (Fr.) P. Kumm. Now, it is represented by more
than 1500 species in Hymenogastraceae (Matheny et al.
2006; Redhead et al. 2007; Kirk et al. 2008; He et al. 2019).
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Fungal Diversity (2022) 117:1–272
Psilocybe (Fr.) P. Kumm., Führ. Pilzk. (Zerbst): 21 (1871)
Psilocybe was first introduced by Kummer (1871) with
P. semilanceata (Fr.) P. Kumm. as the type species. This
genus is saprotrophic and widely distributed in both tropical and temperate areas (Singer and Smith 1958; Guzmán
1978, 1983; Redhead et al. 2007; Kirk et al. 2008), while
Psilocybe sensu lato is known to include Deconica. Both
Psilocybe and Deconica have been characterized by typically hygrophanous basidiomata, brown to yellow–brown
pileus, lilac-brown to dark brown to dark purple-brown spore
prints, ellipsoid to rhomboid to subhexagonal basidiospores
with a distinct apical germ pore (Singer and Smith 1958;
Guzmán 1978, 1983; Redhead et al. 2007). Most Psilocybe
contains psychedelic compounds, e.g. baeocystin, psilocin
and psilocybin (Stamets 1996), whereas Deconica possesses
none of these compounds (Marcano et al. 1994). Traditionally, Psilocybe and Deconica belong to Strophariaceae,
order Agaricales (Guzmán 1978, 1983). However, multiphylogenetic analyses have revealed that Psilocybe formed
a monophyletic genus in Hymenogastraceae, order Agaricales, which clearly separates it from Deconica (RamírezCruz et al. 2013). There are more than 300 accepted species
of Psilocybe in the Index Fungorum (2022a, b), however,
only eight Psilocybe species have been reported in Thailand (P. cubensis (Earle) Singer, P. magnispora E. Horak,
Guzmán & Desjardin, P. samuiensis Guzmán, Bandala &
J.W. Allen, P. subaeruginascens Höhn, P. thaiaerugineomaculans Guzmán, Karun. & Ram.-Guill., P thaicordispora
Guzmán, Ram.-Guill. & Karun., P. thaiduplicatocystidiata
Guzmán, Karun. & Ram.-Guill., and P. thailandensis E.
Horak, Guzmán & Desjardin and P. thaizapoteca Guzmán,
Karun. & Ram.-Guill.) (Guzmán et al. 1993, 2012; Horak
et al. 2009; Chandrasrikul et al. 2011).
Psilocybe keralensis K.A. Thomas, Manim. & Guzmán,
Mycotaxon 83: 196, 2002.
Index Fungorum number: IF380972; Facesoffungi number: FoF10681;Fig. 152
Pileus 13–25 mm diameter, hemisphearic, subconic or
campanulate, hygrophanous, brownish-orange (6C4) to greyish orange (5B6), fading to light orange (5A4) to orangewhite (5A2), surface glabrous, lucidus when dry and often
somewhat bluish when touched or on drying, especially at
the edge, not viscid and the margin finely translucent striate
when moist; context pale yellow (2A3) or yellowish-white
(2A2), bruising ink blue. Lamellae adnate to slightly sinuate, close, light orange (5A4) to orange (5A6) to brownish grey (9C2) or reddish-brown (9E5), often with ink
blue tinge, edges serrulate and remaining whitish. Stipe
45–80 × 1.5–3 mm, equal, sometimes flattened and becoming tapered toward the base, nearly concolorous with the
pileus, darker below, often with ink blue to blackish tinge
when touched or when dry, and shiny when dry; surface
Fungal Diversity (2022) 117:1–272
187
Fig. 153 Phylogenetic tree
derived from maximum likelihood analysis of a combined
ITS and LSU genes of 25
sequences and the aligned
dataset was comprised of 1640
characters including gap. The
average standard deviation of
the split frequencies of the BI
analysis was 0.00962. Panaeolus acuminatus (CBS 270.47)
and Pa. subbalteatus (CBS
326.34) were used as outgroup
taxa. The numbers above
branches are the bootstrap
statistics percentages (left) and
Bayesian posterior probabilities
(right). Branches with bootstrap
values ≥ 70% are shown at each
branch and the bar represents
0.1 substitutions per nucleotide
position. Hyphen (-) represents
support values ≤ 70/0.90. Extype strains are in black bold.
The newly generated sequences
are indicated in blue
longitudinally striate and covered with appressed whitish
fibrils or flocculose, sometimes uneven and with scrobicula
and grooves; base with white mycelium and often radicating;
annulus absent; context fragile, yellowish and yellowishbrown towards the surface and base, staining somewhat ink
blue when bruised. Spore print dark brown.
Basidiospores 6–10 × 5–7 × 4–6 μm, often subrhomboid
or ovoid, sometimes ellipsoid, occasionally inconspicuous
subhexagonal in face view, Q = 1.1–1.7, Qm = 1.35 ± 0.13,
ellipsoid or subovoid, sometimes nearly oblong inside
view, Q = 1.2–1.8, Qm = 1.55 ± 0.15; yellowish-brown with
a purple tinge in water, dark yellow to yellowish-brown in
KOH, dark purplish-brown in deposit; wall smooth, slightly
thick (0.5–1 μm), complex, with 0.8–1.5 μm wide apical
germ pore. Basidia 18–28 × 5–8 μm, hyaline, long subcylindric to clavate, often constricted in middle and narrowed
in the lower half, 4-spored and 2-spored, rarely 1-spored;
sterigmata up to 6 μm. Pleurocystidia are relatively rare
and scattered, 13–22 × 4–6.5 μm, hyaline, ventricose to
sublageniform, sometime fusoid or clavate, occasionally nearly cylindric-clavate or obclavate, often with a
1–7.5 × 1–2 μm neck or rostrum, not branched, the top or
apex often seems wall thickened or contain some matter.
Cheilocystidia abundant, 14–35.5 × 4–8 μm, hyaline, similar to pleurocystidia, mostly with a 1.5–13 × 1–3 μm rostrum or neck, sometimes with an acuminate apex, the top
or apex often seems wall thickened or contain some matter.
Pileipellis an ixocutis, 15–80 μm thick, made up of creeping,
interwoven, 2–6 μm wide filiform to slender tubular hyphae,
hyaline and colourless, wall-smoothed and thin; subpileipellis more pigmented to dark yellow in KOH. Subhymenium
subcellular, hyaline, composed of irregular vesiculose to
polygonal or subglobose cells. Caulocystidia abundant,
14–49 × 3.5–12 μm, scattered, gregarious to clustered at
the upper part of the stipe, hyaline, similar to cheilocystidia, hyaline, thick-walled. Clamp connections present in
all tissues.
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Fungal Diversity (2022) 117:1–272
Fig. 154 Marasmius pallidoaurantiacus BKF10248
(holotype). a Basidiomata, b
Basidiospores, c Basidium,
d Checlocystidia, e Pileipellis cell. Scale bars: a = 2 mm,
b–e = 10 μm. Photographs and
drawing by N. Wannathes
Material examined: Thailand, Ubon Ratchathani Province, Phibun Mangsahan, District, 15° 3′ 25″ N 105° 26′ 50″
E, elevation 164 m, on soil, 4 May 2019, N. Suwannarach &
J. Kumla, SDBR-CMUNK0448.
Habitat: Growing solitary to scattered on dung or soil of
meadows, or grassland in an open area.
Distribution: India (Thomas et al. 2002), Southwestern
China (Ma et al. 2016) and Thailand (This study).
GenBank numbers: MZ452082 (ITS), MZ452083 (LSU).
Notes: Morphologically, P. keralensis is similar to P.
columbiana Guzmán; however, P. columbiana differs significantly in the absence of pleurocystidia. It has been known
to only be from Colombia (Guzmán 1978, 1983; Guzmán
et al. 2005). Phylogenetically, P. keralensis formed a sister taxon to P. thaizapoteca (Fig. 153). However, the large
pileus (20–50 mm in diameter) and small basidiospores
(6–7 × 3–4.5 × 3–3.5 μm) of P. thaizapoteca clearly differ
from P. keralensis (Guzmán et al. 2012).
Marasmiaceae Roze ex Kühner.
Note: Marasmiaceae was erected by Kühner (1980) with
a combination of Marasmieae, Collybieae and Myceneae
of Tricholomataceae R. Heim in Singer (1986) classification. Molecular phylogeny has revealed that Marasmiaceae
is monophyletic, containing ten genera (Amyloflagellula
Singer, Brunneocorticium Sheng H. Wu, Campanella Henn.,
Chaetocalathus Singer, Crinipellis Pat., Hymenogloea Pat.,
Marasmius Fr., Moniliophthora H.C. Evans, Stalpers,
Samson & Benny, Neocampanella Nakasone, Hibbett &
Goranova and Tetrapyrgos E. Horak) with more than 1590
species, Marasmius being the generic type (Matheny et al.
2006; Kirk et al. 2008; He et al. 2019) of this family.
Marasmius Fr., Fl. Scan.: 339 (1836).
Marasmius is a large genus of plant debris decomposers
(Singer 1976a, b, 1986; Antonín and Noordeloos 2010) that
is represented by approximately 600 species worldwide (He
et al. 2019). The genus was first introduced by Fries (1835)
13
and Marasmius rotula (Scop.). Fr. was later designated as
the type species by Singer and Smith (1946). Marasmius is
characterized by small or possibly large to robust basidiomata, convex or campanulate, striate to sulcate, white to
strongly pigmented pilei, well-developed lamellar, thin and
typically tough, cylindrical, often brown or darkly fuscous
stipe, white spore print, hyaline, smooth and inamyloid
spores and hymeniform pileipellis of smooth or diverticulate cells (Singer 1976a, b, 1986; Antonín 2007). Generic
circumscription of Marasmius sensu stricto was restricted
(Wilson and Desjardin 2005) and the infrageneric classification has been revised based on the results of a phylogenetic
analysis of combined sequences of LSU and ITS incorporated with other notable morphological features (Oliveira
et al. 2020b).
Marasmius pallidoaurantiacus Wannathes, N. Suwannarach, J. Kumla & S. Lumyong, sp. nov.
MycoBank number: MB 840,379; Facesoffungi number:
FoF 10682;Fig. 154
Etymology: ‘pallio’ = pale, ‘aurantiacus’ = orange colour, refers to the pileus colour
Holotype: BKF10248
Pileus 1–2 mm diam., hemispherical, umbilicate, with a
tiny dark brown papilla at center; dull, dry, glabrous, slightly
striate; orange white (5A2) to pale orange (5A3) with darker
disc. Context thin, white. Lamellae adnate to a small collarium, distant (10–11), broad (1 mm), orange white (5A2),
non-marginate. Stipe 8–13 × 0.2 mm, cylindrical, central;
glabrous, dull, dry; insititious; dark brown overall, sometime
stipe arising directly from dark brown rhizomorph.
Basidiospores 7–10(–11) × 4–5(–6) µm
[x = 8.20 ± 1.35 × 4.18 ± 0.58, Q = 1.75–2.57, q = 1.97 ± 0.30,
n = 25] ellipsoid, fusoid, smooth, hyaline, inamyloid, thinwalled. Basidia 14–16 × 4–6 µm, clavate, with 4 sterigmata,
thin-walled, inamyloid. Cheilocystidia abundant, of Siccus- type broom cells; main body 5–7 × 4–6 µm, cylindrical
to clavate, hyaline, inamyloid, thin-walled; apical setulae
Fungal Diversity (2022) 117:1–272
189
Fig. 155 Phylogenetic tree
derived from maximum likelihood analysis of a combined
ITS gene of 65 sequences
and the aligned dataset was
comprised of 1116 characters
including gap. The average
standard deviation of the split
frequencies of the BI analysis
was 0.00825. Marasmius siccus
KG 039 and KG 119 were used
as outgroup. Numbers above
branches are the bootstrap
statistics percentages (left) and
Bayesian posterior probabilities
(right). Branches with bootstrap
values ≥ 70% are shown at each
branch and the bar represents
0.1 substitutions per nucleotide
position. Hyphen (-) represents
support values ≤ 70%/0.95. Extype strains are in black bold.
The newly generated sequences
are indicated in blue and bold
type species
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190
Fungal Diversity (2022) 117:1–272
Fig. 156 Marasmius tangerinus BKF10249 (holotype). a
Basidiomata, b Basidiospores,
c Basidium, d Checlocystidia, e Pileipellis cell. Scale
bars: a = 5 mm, b–e = 10 μm.
Photographs and drawing by N.
Wannathes
1–4 × 1 µm, cylindrical to conical, obtuse, hyaline to pale
yellow, thin- to thick-walled. Pleurocystidia absent. Pileipellis hymeniform, mottled, composed of Siccus-type broom
cells; main body 10–14 × 4–6 µm, clavate to broadly clavate,
flexuose, pale yellow, inamyloid, thin- to thick-walled; apical setulae (1–) 2–6 × 1 µm, cylindrical, regular in outline,
obtuse, brown, thick-walled. Pileus trama interwoven, inamyloid. Lamellar trama interwoven, hyphae 2–5 µm diam.,
cylindrical, inflated, smooth, hyaline, inamyloid, thinwalled, non-gelatinous. Stipitipellis subparallel, hyphae
2–4 µm diam., cylindrical, dark brown, smooth, inamyloid,
thick-walled (up to 1 µm), non-gelatinous. Stipe trama subparallel, hyphae 2–5 µm diam., cylindrical, hyaline to pale
yellow, smooth, dextrinoid, thin-walled, non-gelatinous.
Caulocystidia absent. Clamp connections present in almost
all tissues.
Material examined: Thailand, Nakhon Ratchasima
Provinve, Khao Yai National Park, 33 km marker on Hyw
2090, 22 Sep 2018, collectors N Wannathes, N Suwannarach
J Kumla, S Lumyong, BKF10248 (holotype), NW1108
(isotype).
Habit, habitat and known distribution: Gregarious on
dicotyledonous leave, known only from Thailand.
13
GenBank numbers: MZ452673 (ITS).
Notes: Marasmius pallidoaurantiacus is characterized
by a tiny, hemispherical with dark brown conical papilla
at the center, slightly striate, plae orange pileus, collariate, distant lamellae, thin stipe, sometimes arising directly
from the rhizomorph, ellipsoid to fusoid basidiospores
with mean 8.2 × 4.2 µm, and the presence of Siccus-type
broom cell cheilocytidia. This new species is morphologically similar to Marasmius pallenticeps Singer, which was
originally described from Argentina. The latter differs by
forming a pure white pileus, stipe usually arising directly
from rhizomorphs and the presence of nodes (Desjardin and
Ovrebo 2006). Marasmius pallidoaurantiacus pilei appear
pigmented like those of M. longibasidiatus J.S. Oliveira,
a species recently described from Brazil. This species differs by forming bigger (1.3–4 mm diam.), sulcate pileus,
longer basidia (33–42 × 7.8–10 µm) and pileipellis composed of mixed Siccus type broom cell and non-setulost cells
(Oliveira 2020). Phylogenetic analyses inferred from ITS
sequence data (Fig. 155) confirmed that M. pallidoaurantiacus is a distinct species from other related morphological
species and other taxa within this genus.
Fungal Diversity (2022) 117:1–272
191
Fig. 157 Basidiomata of Rhizomarasmius cunninghamietorum a. HGASMF01-10,709 (holotype), b. HGASMF01-10,708 (paratype). Scale
bars: = 1 cm
Marasmius tangerinus Wannathes, N. Suwannarach, J.
Kumla & Lumyong, sp. nov.
MycoBank number: MB840380; Facesoffungi number:
FoF10683; Fig. 156
Etymology: ‘tangerinus’ = tangerine colour, refers to the
pileus colour
Holotype: BKF10249
Pileus 2–3 mm diam., hemispherical to convex, umbilicate, with a tiny dark brown papilla at center; dull, dry, glabrous, striate; orange red (8A8) overall when young, orange
red (8A8) at disc, orange (6B7) to tangerine (6B8) at margin in age. Context thin, white. Lamellae adnate to a small
collarium, distant (10–11), narrow (0.5 mm), pale orange
(5A3), marginate with orange. Stipe 13–20 × 0.3 mm, cylindrical, central, glabrous, dull, dry, insititious, golden brown
(5D7) overall, sometimes stipe arising directly from golden
brown rhizomorph.
Basidiospores
8–10 × (4–)4.5–5
µm
[x = 8.56 ± 0.71 × 4.92 ± 0.28, Q = 1.6–2.0, q = 1.74 ± 0.16,
n = 25] ellipsoid, fusoid, smooth, hyaline, inamyloid, thinwalled. Basidia 16–18 × 6–8 µm, clavate, with 4 sterigmata,
thin-walled, inamyloid. Cheilocystidia abundant, of Siccus- type broom cells; main body 10–14 × 5–7 µm, cylindrical to clavate, sometime branched, hyaline, inamyloid,
thin-walled; apical setulae 1–4 × 1(–2) µm, cylindrical to
conical, obtuse, pale yellow, thin- to thick-walled. Pleurocystidia absent. Pileipellis hymeniform, not mottled,
composed of 2 type of cells: (a) Siccus-type broom cells;
main body 18–38 × 10–16 µm, clavate to broadly clavate or
turbinate, irregular in outline, hyaline to pale yellow, inamyloid, thin- to thick-walled; apical setulae 2–10 × 1–4 µm,
cylindrical to conical, regular in outline, obtuse, yellow to
pale brown, inamyloid, thick-walled; (b) non-setulose cell,
8–32 × 10–15 µm, clavate to broadly clavate, sometime
branched, yellow to pale brown, inamyloid, thick-walled.
Pileus trama interwoven, inamyloid. Lamellar trama interwoven, hyphae 4–8 µm diam., cylindrical, inflated, smooth,
hyaline, inamyloid, thin-walled, non-gelatinous. Stipitipellis subparallel, hyphae 4–6 µm diam., cylindrical, pale
brown, smooth, dextrinoid, thick-walled (up to 1 µm), nongelatinous. Stipe trama subparallel, hyphae 2–4 µm diam.,
cylindrical, hyaline, smooth, dextrinoid, thin-walled, nongelatinous. Caulocystidia absent. Clamp connections present
in almost all tissues.
Material examined: Thailand, Nakhon Ratchasima
Provinve, Khao Yai National Park, 33 km marker on Hyw
2090, 22 Sep 2018, collectors N Wannathes, N Suwannarach
J Kumla, S Lumyong, BKF10249 (holotype), NW1224
(isotype).
Habit, habitat and known distribution: Gregarious on
dicotyledonous wood, known only from Thailand.
GenBank numbers: MZ452087 (ITS).
Notes: Marasmius tangerinus is characterized by a
tiny, hemispherical, bright orange pileus, with dark brown
papilla at the center, collariate, distant lamellae, thin stipe,
sometimes arising directly from the rhizomorph, ellipsoid
to fusoid basidiospores with mean 8.6 × 4.9 µm, pileipellis
composed of mixed Siccus type broom cell and non-setulost cells, with the presence of cheilocytidia, and ligicolous
habit. Marasmius tangerinus is morphologically similar to
M. trichorhizus Speg. which has been characterized from a
neotropic realm. The latter species differ by forming more
lamellae (12–15), longer stipe (4–71 mm long), and smaller
basidiospores with mean 6.9 × 3.4 µm (Oliveira 2020). The
phylogenetic analyses of the ITS sequence data (Fig. 155)
indicate that M. tangerinus is a distinct species from other
related morphological species and other taxa within this
genus.
Physalacriaceae Corner, Beih. Nova Hedwigia 33: 10
(1970).
Notes: The family was typified by Physalacria Peck
(Peck 1883), which contained a number of marasmoid
fungi, including the important tree pathogen Armillaria (Fr.)
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192
Fungal Diversity (2022) 117:1–272
Fig. 158 Microscopic structures of Rhizomarasmius cunninghamietorum (holotype). a.
Basidiospores, b. Basidia and
Basidioles, c. Cheilocystidia, d.
Caulocystidia, e. Pileiocystidia,
f. Pleurocystidia, g. Terminal cells of pileipellis, Scale
bars = 10 μm
Staude and edible fungi Flammulina P. Karst., this group
usually have pilocystdia and non-dextrinoid tissue (Petersen
and Hughes 2010). More than 28 genera included in this
family by Index Fungorum (2022a, b) and are accepted by
Wijayawardene et al. (2022).
R. setosus (Sowerby) Antonín & A. Urb. To date, there are
five species described within this genus. Petersen (2000),
Petersen and Hughes (2010) and Ronikier and Ronikier
(2011), showed Rhizomarasmius to form a well-supported
clade in phylogenetic analyses.
Rhizomarasmius R.H. Petersen, Mycotaxon 75: 333 (2000).
Notes: Petersen (2000) accommodated two Marasmius
species, M. pyrrhocephalus Berk. and M. undatus (Berk.)
Fr. to form a new genus, Rhizomarasmius R.H. Petersen.
This genus is characterized by convex to hemispherical
pileus, distant, thick, ascending, white lamellae and central
stipe apically pale and darkening downward to brown-black,
smooth basidiospores, pileipellis forming a hymeniderm
layer of clavate, globose to sphaeropedunculate, smooth
elements and scattered, elongate pileocystidia, as well as
the presence of cheilo-, pleuro- and caulocystidia (Petersen
2000; Antonín and Noordeloos 2010). Most members of
marasmoid are saprotrophic or parasitic fungi (Pacioni
and Lalli 1989; Filippi 1991; Ronikier and Ronikier 2011;
Moreau et al. 2015). Rhizomarasmius epidryas (Kühner
ex A. Ronikier) A. Ronikier and M. Ronikier is a common
arctic-alpine saprotrophic fungus (Ronikier 2009, 2011), and
this species was originally described by Kühner as Marasmius epidryas Kühner in 1935 (Kühner 1936). Wilson
and Desjardin (2005) and Noordeloos and Antonín (2008)
transferred this species into Mycetinis Earle. However, later
established that the species belongs to the new genus Rhizomarasmius by Anna & Macheł Ronikier and changed the
current name accordingly (Ronikier 2009, 2011). Recently
Moreau et al. (2015) added two members, Rhizomarasmius
oreinus (Pacioni & Lalli) Vizzini, Antonín & A. Urb. and
Rhizomarasmius cunninghamietorum Chun Y. Deng, J.P.
Li & Gafforov, sp. nov.
Index Fungorum number: IF558751; Facesoffungi number: FoF10408; Figs. 157, 158
Etymology: the species epithet refers to the host genus
Cunninghamia.
Holotype: HGASMF01-10709
Basidiomata gregarious, marasmioid, marasmielloid or
collybioid. Pileus 9–36 mm in diameter, convex to planoconvex when young, expanding to applanate with age,
obtuse or slightly depressed at the centre, often wrinkled,
slightly sticky, sometimes hygrophanous, translucent when
mosit, radially striate or sulcate from the margin up to 3/4
of the radius nearly to the centre, with transversely sulcate
between each radial striate when old, involute then inflexed
to reflexed at margin, with marginal zone often undulating
with age, almost white overall when young, with somewhat
milk white (1A2) pigment, then change to generally orange
white (6A2), pale orange (6A3) or cinnamon (6D6) overall,
paler towards margin, henna (7E8) to dark brown (8E8) at
disc when old, darker in the groove, paler at margin. Lamellae adnate, sometimes with slightly decurrently tooth, furcate, intervenose, or anastomosing, often split, linear to
ventricose, subdistant, L = 10–14, l = 1–2, white overall,
often with light brown (6D4) to reddish brown (8E8) tint,
up to 3 mm broad. Stipe 6–23 × 1–3 mm, central, insititious,
13
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193
Fig. 159 Phylogenetic tree derived from maximum likelihood analysis of a combined ITS and LSU sequence data. The tree is rooted with
Laccariopsis mediterranea and Hydropus mediterraneus. Branches
with bootstrap values ≥ 70% are shown at each branch and the bar
represents 0.1 substitutions per nucleotide position. Ex-type strains
are in black bold. The newly generated sequences are indicated in
blue and bold type species
hollow, pruinose, more or less cylindrical above, sometimes
broadened in somewhere, often tapering towards base, pallid
at the apex, orange white (6A2) or pale orange (6A3) to dark
brown (8E8) below, black at base. Basidiospores (3.5)5–6.5
(7) × 3–4 μm (average = 5.77 × 3.42 μm, E = 1.43–2(–2.16),
Q = 1.69), ellipsoid, amygdaliform, thin-walled, nondextrinoid, hyaline, smooth. Basidia 21–35 × 4.5–6 μm,
4-spored, clavate, hyaline. Basidioles 21–35.5 × 4–6 μm,
clavate, cylindrical, hyaline. Cheilocystidia sparse to abundant, 11–50 × 7–17 μm, ellipsoid, cylindrical, irregular
clavate, narrowly utriform to narrowly fusiform with capitate apex, thin-walled. Pleurocystidia scattered, sparse,
narrowly utriform to narrowly fusiform with capitate apex,
39–64.5 × 7–12.5 μm. Pileipellis a hymeniform layer of
clavate to sphaeropedunculate smooth cells measuring
12.5–29.5 × 9–15.5 μm, possibly gelatinized. Pileiocystidia 35.5–51 × 8–16.5 μm, fusiform, narrowly lageniform
to lageniform with capitate apex, thin-walled. Stipitipellis a cutis, of cylindrical, parallel, slightly thick-walled,
non-dextrinoid, up to 19.5 μm wide hyphae. Caulocystidia
35–59 × 6–12 μm, cylindrical to broadly cylindrical with or
without capitate apex, sometimes lobed, thick-walled. Clamp
connections present in all tissue.
Material examined: China, Guizhou Province, Qiandongnan Miao Autonomous Prefecture, Liping County, Dongfeng
tree farm, on the dead trunk of Cunninghamia lanceolata
(Lamb.) Hook., (Cupressaceae), 17 October 2020, Ji-Peng Li
(LJP563), HGASMF01-10,709 (holotype), ibid., 17 October
2020, Ji-Peng Li (LJP564), HGASMF01-10,708 (paratype).
GenBank numbers: HGASMF01-10709- MW326779
(ITS), MW332104 (LSU).
HGASMF01-10708- MW326780 (ITS), MW332105
(LSU)
Notes: Rhizomarasmius cunninghamietorum has an insititious stipe, a pileipellis as a hymeniform layer of clavate
to sphaeropedunculate smooth cells, with narrowly lageniform to lageniform with capitate apex pileocystidia. These
morphological characteristics of R. cunninghamietorum
are consistent with the circumscription of Rhizomarasmius
(Moreau et al. 2015). The phylogenetic evidence showed
that R. cunninghamietorum formed a single branch and was
clearly separate from the known species of Rhizomarasmius and closer to its related genera Cibaomyces and Gloiocephala (Fig. 159). However, Cibaomyces was accepted
as a monotypic genus based on the subglobose to broadly
ellipsoid basidiospores with conspicuous spines and Gloiocephala sensu Singer with smooth spores, subcapitate to
capitate cystidia, a gelatinized trama (Singer 1976a, b, 1986)
is probably polyphyletic (Moncalvo et al. 2002; Binder et al.
2006; Vizzini et al. 2012; Hao et al. 2014). Thus, Rhizomarasmius is probably polyphyletic based on the phylogenetic analysis generated here. Morphologically, among the
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Fig. 160 Grammothele taiwanensis (GC 1704–17, holotype).
a, b Basidiomata. c Generative
hyphae from trama. d Skeletal
hyphae from trama. e Generative hyphae from context. f
Skeletal hyphae from context. g
Dendrohyphidia. h Hyphidia. i
Cystidioles. j Basidia. k Basidiospores. Scale bars: a–b = 1 cm,
b–j = 10 μm, k = 5 μm
known species of Rhizomarasmius, R. epidryas has entirely
brown-black and distinctly velutinose stipe, bigger basidiospores (average 8.85 × 5.7 μm) and narrowly clavate or
cylindrical pleurocystidia (Ronikier 2009); R. setosus with
different habitat on leaf petioles and its microscopic features without clamps (Antonín and Noordeloos 2010); Rhizomarasmius oreinus has smooth red-brown pileus, entirely
tomentose pubescent stipe and bigger basidiospores (average 12.5 × 8.8 μm, Moreau et al. 2015); R. pyrrhocephalus
13
has pubescent to tomentose stipe, and bigger basidiospores
(average 8.2 × 3.7 μm, Desjardin 1989) and R. undatus has
longer (5.0–6.3 cm) and strigose stipe (Smith 1836).
Polyporales Gäum.
Notes: Polyporales is a large group of Agaricomycetes,
accommodating about 2500 species (He et al. 2019). The
latest treatments and updated accounts of Polyporales are
followed in Justo et al. (2017), He et al. (2019), and Wijayawardene et al. (2022).
Fungal Diversity (2022) 117:1–272
Polyporaceae Fr. ex Corda [as 'Polyporei'], Icon. fung.
(Prague) 3: 49 (1839).
Notes: Polyporaceae, typified by Polyporus P. Micheli
ex Adans, currently includes 90 genera, most of which are
polypores, rarely corticioid species (Justo et al. 2017; He
et al. 2019; Wijayawardene et al. 2022). Microscopically,
the hyphal system is usually dimitic or trimitic, rarely monomitic; the generative hyphae are usually with clamp-connections; cystidia are mostly absent; basidiospores are thin- to
thick-walled, smooth to ornamented and colorless to brown.
All species produce a white rot (Justo et al. 2017).
Grammothele Berk. & M.A. Curtis, Journal of the Linnean
Society. Botany 10: 327 (1869).
Notes: Grammothele, typified by G. lineata Berk. &
M.A. Curtis, accommodates 24 species (Zhou and Dai
2012; Ryvarden 2015; Yuan 2015; Wu et al. 2016; Hyde
et al. 2019). The genus is characterized by having resupinate basidiomata with shallow poroid hymenophore consisting of angular, partly sinuous, irregular or incomplete
pores, dimitic or trimitic hyphal system with clamped
generative hyphae and dextrinoid skeletal hyphae, usually
presence of dendrohyphidia, and ellipsoid to cylindrical
basidiospores that are thin-walled, smooth, colorless and
not reacting both in Melzer’s reagent and Cotton Blue.
Species of Grammothele commonly occur on hardwoods
and monocotyledons in tropical to subtropical regions,
causing a white rot (Ryvarden 1979, 2015). Previous phylogenetic studies have shown that Grammothele is polyphyletic and closely related to some other genera in the
Polyporaceae, such as Porogramme (Pat.) Pat., Theleporus Fr. and Tinctoporellus Ryvarden (Zhou and Dai 2012;
Yuan 2015; Wu et al. 2016; Hyde et al. 2019).
Grammothele taiwanensis C.C. Chen, sp. nov.
Index Fungorum number: IF900073; Facesoffungi number: FoF13396; Fig. 160
Etymology: Referring to the type locality, Taiwan.
Holotype: GC 1704-17
Basidiomata annual, resupinate, effused, adnate, corky
when dry, up to 15 cm long, 6 cm wide, and 1.3 mm thick
in section, sparsely cracked, growing beneath the bark;
margin sterile, cottony or slightly fimbriate, sometimes
indistinct. Pore surface cream, buff-yellow or buff upon
drying, not changing in KOH; pores angular, 3–5 per
mm; dissepiments thin, entire, sometimes lacerate. Tubes
concolorous with pore surface, corky, up to 1 mm deep.
Context cream, corky, up to 0.3 mm thick, or sometimes
invisible. Hyphal system dimitic in both context and trama;
generative hyphae nodose-septate; tissues not changing in
KOH. Context generative hyphae colorless, unbranched,
2.5–3.5 µm diam, thin-walled; skeletal hyphae, colorless,
slightly flexuous, occasionally branched, 2.5–3.5 µm diam,
195
thick-walled with a wide lumen or almost solid, weekly
dextrinoid. Tramal generative hyphae colorless, flexuous,
frequently branched, 2–3.5 µm diam, thin-walled; skeletal hyphae colorless, slightly flexuous, rarely branched,
3–4.5 µm diam, thick-walled with a wide lumen or subsolid, often agglutinated in bundles, weekly dextrinoid.
Cystidia absent. Cystidioles fusoid, colorless, thin-walled,
14–20 × 4.5–5 µm. Dendrohyphidia colorless, thin-walled,
9–23 × 1.5–2.5 µm. Hyphidia colorless, thin-walled,
19–26 × 2–3 µm. Basidia clavate or suburniform, with
four sterigmata and a basal clamp connection, colorless,
thin-walled, 17–25 × 6–7 µm. Basidiospores cylindrical,
colorless, thin-walled, smooth, inamyloid, nondextrinoid,
acyanophilous, mostly 7.9–10.5 × 3.3–4.4 µm. (7.8–)8
.4–10.5(–12.3) × (3.5–)3.7–4.4(–5.1) µm, L = 9.4 µm,
W = 4.1 µm, Q = 2.23–2.43 (n = 30) (holotype). (7–)7.9–9
.4(–10.6) × (3–)3.3–3.9(–4.3) µm, L = 8.6 µm, W = 3.6 µm,
Q = 2.3–2.45 (n = 30) (GC 1704–16).
Material examined: China: Taiwan, Nantou County,
Yuchih Township, Shuishetashan Trail, 23° 51′ N, 120° 56′
E, 980 m, on angiosperm branch, 8 April 2017, GC 1704–17
(TNM F31469, holotype).
GenBank number: MW440487 (ITS).
Additional specimen examined: China: Taiwan, Nantou
County, Yuchih Township, Shuishetashan Trail, 23° 51′ N,
120° 56′ E, 980 m, on angiosperm branch, 8 April 2017, GC
1704–16 (TNM F31468).
GenBank number: MW440486 (ITS).
Notes: Grammothele taiwanensis is characterized
by having resupinate, cream to buff basidiomata with
larger pores (3–5 per mm), dimitic hyphal system with
clamped generative hyphae and weekly dextrinoid skeletal hyphae, presence of fusoid cystidioles, dendrohyphidia and hyphidia, and cylindrical basidiospores
(7.9–10.5 × 3.3–4.4 µm). Grammothele taiwanensis resembles G. pulchella (Bres.) Ryvarden in having similar pore
and basidiospore sizes [4–5 per mm, 7–10 × 3–4 µm in
Hjortstam and Ryvarden (1982), as G. ochracea Ryvarden]
and presence of dendrohyphidia; however, G. pulchella
differs from this species in having cork-colored to woodcolored pore surface and pale brown context, and absence
of cystidioles and hyphidia (Hjortstam and Ryvarden
1982). Grammothele taiwanensis is also similar to G. hainanensis F. Wu & L.W. Zhou and G. hondurensis (Murrill)
Ryvarden, but the latter two species have hyphal pegs and
smaller basidiospores [G. hainanensis: 7–8.1 × 2.3–2.9 µm
in Wu et al. (2016); G. hondurensis: 5–8 × 3–3.5 µm in
Ryvarden (1985)]. Phylogenetically, two sequences of G.
taiwanensis formed an isolated lineage with high support
(Fig. 161).
Incrustoporiaceae Jülich, Biblthca Mycol. 85: 373 (1982)
[1981].
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196
Fig. 161 Phylogram generated from maximum likelihood analysis
based on ITS sequence data of Grammothele taiwanensis (in bold)
and related species. The selection of strains and species consulted
Zhou and Dai (2012), Yuan (2015), Wu et al. (2016) and Hyde et al.
(2019). Funalia trogii is used as the outgroup taxa. Forty-six strains
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Fungal Diversity (2022) 117:1–272
are included in the sequence analyses, which comprise 665 characters with gaps. The tree topology of the ML analysis was similar to
the BI. The best scoring RAxML tree with a final likelihood value of
is presented. Bootstrap support values for ML ≥ 70%, BYPP ≥ 0.9 are
given above the nodes
Fungal Diversity (2022) 117:1–272
197
Notes. Incrustoporiaceae was established by Jülich (1981)
and typified by Incrustoporia Domański. This family was
treated as a synonymy of Polyporaceae according to the 10th
edition of Dictionary of the Fungi (Kirk et al. 2008). Justo
et al. (2017) provided a revised family-level classification
of the Polyporales based on phylogenetic analyses inferred
from ITS, nLSU and rpb2 genes, and four genera were
accepted in Incrustoporiaceae, viz., Incrustoporia, Piloporia Niemelä, Skeletocutis Kotl. & Pouzar and Tyromyces P.
Karst.
Skeletocutis Kotlába & Pouzar, Ceská Mykologie 12 (2):
103 (1958).
Notes: Skeletocutis Kotl. & Pouzar was established in
1958 and its type species is S. amorpha (Fr.) Kotl. & Pouzar (Kotlába and Pouzar 1958). The genus has resupinate to
pileate basidiocarps, encrusted generative hyphae covered
by fine crystals and tiny basidiospores, these are the most
important characteristics of the genus. In addition, species
in Skeletocutis cause white rot disease (Niemelä 1998). Species in this genus have wide distribution in the world, but the
majority of the known species so far are found in the Northern Hemisphere (Gilbertson and Ryvarden 1986; Núñez and
Ryvarden 2001; Ryvarden and Melo 2014). There are more
than 60 taxa recorded in Skeletocutis and 25 of them occur
in China (Cui and Dai 2008; Dai 2012; Zhou and Qin 2012;
Bian et al. 2016; Fan et al. 2017; Du and Dai 2020).
Recently, taxonomic and phylogenetic studies of polypore
fungi from China have been extensively carried out, and
many new genera or new species have been found (Zhao
et al. 2015; Han et al. 2016; Chen and Cui 2017; Chen et al.
2017a, b; Song and Cui 2017; Zhou and Cui 2017; Xing
et al. 2018; Cui et al. 2019; Shen et al. 2019; Wu et al. 2019;
Zhu et al. 2019; Sun et al. 2020a, b; Liu et al. 2021a, b),
however, only very few studies have been focused on Skeletocutis of the Incrustoporiaceae. In this study, two new
species of Skeletocutis are described from China based on
morphological characteristics and phylogenetic analyses
inferred from ITS + nLSU sequences.
Skeletocutis cangshanensis B.K. Cui & Shun Liu, sp. nov.
Index Fungorum number: IF559465; Facesoffungi number: FoF10674; Figs. 162a, 163
Differs from other Skeletocutis species by its white pore
surface when fresh, white to buff-yellow upon drying, small
and circular to angular pores (7–10 per mm), and cylindrical basidiospores (2.7–3.5 × 0.8–1.5 μm).
Etymology. Cangshanensis (Lat.): refers to the type locality (Cangshan Park) of the type specimen.
Holotype: Cui 17978.
Fruiting body. Basidiocarps annual, resupinate, not easily separated from substrate, soft leathery, without odour or
taste when fresh, becoming corky upon drying, up to 8.5 cm
Fig. 162 Basidiocarps of Skeletocutis species. a. S. cangshanensis;
b. S. subchrysella (scale bars: a, b = 2 cm)
long, 3.5 cm wide, and 1 mm thick at center. Pore surface
white when fresh, becoming white to buff-yellow upon drying; pores circular to angular, 7–10 per mm; dissepiments
thick, entire. Subiculum white, corky, up to 0.2 mm thick.
Tubes darker than poroid surface, corky, up to 0.5 mm long.
Hyphal Structure. Hyphal system dimitic; generative
hyphae with clamp connections, hyaline, thin- to slightly
thick-walled, dominant at dissepiment edge; skeletal hyphae
thick-walled with a wide to narrow lumen; IKI–, CB–,
unchanged in KOH.
Subiculum. Generative hyphae frequent, hyaline, thinto slightly thick-walled, rarely branched and bearing fine
crystals, 1.5–2.8 μm in diameter; skeletal hyphae dominant,
thick-walled with a narrow lumen, flexuous, unbranched,
interwoven, 2–3.5 μm in diameter.
Tubes. Generative hyphae frequent, thin-walled, frequently branched, usually covered by fine crystals,
sharply pointed encrustations, especially at dissepiment
edge, 1.5–2.5 μm in diameter; skeletal hyphae dominant,
thick-walled with a wide to narrow lumen, occasionally
branched, subparallel along the tubes, not agglutinated,
2–3 μm in diameter. Dissepiment edge dimitic with
smooth skeletal hyphae, and dominant winding, encrusted
generative hyphae. Cystidia absent, cystidioles abundant,
bottle-shaped, with a conical apex, 7.5–17 × 3.2–4.7 μm.
Basidia clavate, with a basal clamp connection and four
sterigmata, 9.6–13.5 × 3.2–4.7 μm; basidioles in shape
similar to basidia, but slightly smaller.
Spores. Basidiospores cylindrical, hyaline, thinwalled, smooth, IKI–, CB–, (2.6–)2.7–3.5 × 0.8–1.5 μm,
L = 3.02 μm, W = 1.02 μm, Q = 2.76–3.23(n = 90/3).
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Fungal Diversity (2022) 117:1–272
Fig. 163 Microscopic structures
of Skeletocutis cangshanensis
(drawn from the holotype). a.
Basidiospores; b. Basidia and
basidioles; c. Cystidioles; d.
Section of dissepiment edge; e.
Hyphae from trama; f. Hyphae
from subiculum. Scale Bars:
a = 5 µm; b–f = 10 µm
Material examined: China, Yunnan Province, Dali,
Cangshan Park, on angiosperm stump, 4 November 2019,
Cui 17978 (holotype, BJFC).
13
Additional specimens examined. China, Yunnan Province, Dali, Cangshan Park, on fallen angiosperm branch, 4
November 2019, Cui 17990, 17994 (Paratypes, BJFC).
GenBank numbers: Cui 17978—MZ327279 (ITS),
MZ348535 (LSU),
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Cui 17990—MZ327280 (ITS), MZ348536 (LSU).
Cui 17994—MZ327281 (ITS), MZ348537(LSU).
Notes: In the phylogenetic tree, the three specimens
of Skeletocutis cangshanensis formed a highly supported lineage (Fig. 164), and grouped together with S.
bambusicola L.W. Zhou & W.M. Qin. Morphologically,
both S. cangshanensis and S. bambusicola have similar
pores, but S. bambusicola differs by having wider basidiospores (2.5–3 × 1.5–2 μm) and growth on Bambusa
(Zhou and Qin 2012). Skeletocutis lepida A. Korhonen
& Miettinen, S. mopanshanensis C.L. Zhao and S. yunnanensis L.S. Bian, C.L. Zhao & F. Wu were also discovered from Yunnan Province. Skeletocutis lepida differs
from the new species by having narrower basidiospores
(2.9–3 × 0.5–0.6 µm; Korhonen et al. 2018); S. mopanshanensis differs by having larger pores (4–5 per mm) and
basidiospores (4.7–6.6 × 3.2–4.5 µm; Wu et al. 2017); S.
yunnanensis differs by having larger pores (5–6 per mm)
and basidiospores (3.4–4.5 × 1–1.2 µm; Bian et al. 2016).
Skeletocutis subchrysella B.K. Cui & Shun Liu, sp. nov.
Index Fungorum number: IF559466; Facesoffungi number: FoF10675; Figs. 162b, 165
Diagnosis: Differs from other Skeletocutis species
by its white to cream pore surface when fresh, cream to
cinnamon-buff upon drying, and allantoid basidiospores
(2.7–3.2 × 0.7–1 μm).
Etymology. Subchrysella (Lat.): refers to the new
species resembling Skeletocutis chrysella Niemelä in
morphology.
Holotype: Cui 17748.
Fruiting body. Basidiocarps annual, resupinate, not easily separated from substrate, soft leathery, without odour or
taste when fresh, becoming corky upon drying, up to 6 cm
long, 1.5 cm wide, and 3 mm thick at center. Pore surface
white to cream when fresh, becoming cream to cinnamonbuff upon drying; pores angular, 6–8 per mm; dissepiments
slightly thick, entire to lacerate. Subiculum cream, corky, up
to 0.5 mm thick. Tubes darker than poroid surface, corky,
up to 2 mm long.
Hyphal Structure. Hyphal system dimitic; generative
hyphae with clamp connections, hyaline, thin-walled, dominant at dissepiment edge; skeletal hyphae with a wide to
narrow lumen; IKI–, CB–, unchanged in KOH.
Subiculum. Generative hyphae frequent, hyaline, thinwalled, unbranched and bearing fine crystals, 1.5–2.8 μm
in diameter; skeletal hyphae dominant, thick-walled with a
narrow lumen, flexuous, unbranched, interwoven, 2–4 μm
in diameter.
Tubes. Generative hyphae frequent, thin-walled,
unbranched, usually covered by fine crystals, sharply pointed
encrustations, especially at dissepiment edge, 1.5–2.5 μm
in diameter; skeletal hyphae dominant, thick-walled with a
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wide to narrow lumen, unbranched, subparallel along the
tubes, not agglutinated, 2–3.7 μm in diameter. Dissepiment
edge dimitic with smooth skeletal hyphae, and dominant
winding, encrusted generative hyphae. Cystidia absent,
cystidioles abundant, bottle-shaped, with a conical apex,
8–13.8 × 2–3.6 μm. Basidia clavate, with a basal clamp
connection and four sterigmata, 10.3–15.6 × 3.2–4.5 μm;
basidioles in shape similar to basidia, but slightly smaller.
Spores. Basidiospores allantoid, hyaline, thin-walled,
smooth, IKI–, CB–, (2.6–)2.7–3.2 × (0.5–)0.7–1 μm,
L = 2.93 μm, W = 0.8 μm, Q = 3.33–4.5(n = 60/2).
Material examined: China, Sichuan Province, Shimian
County, Liziping National Nature Reserve, on fallen angiosperm trunk, 14 September 2019, Cui 17748 (holotype,
BJFC).
Additional specimen examined: China, Yunnan Province,
Baoshan, Gaoligongshan Nature Reserve, on fallen angiosperm trunk, 8 November 2019, Cui 18141 (paratype,
BJFC).
GenBank numbers: Cui 17748- MZ327278 (ITS),
MZ348534 (LSU).
Cui 18141- MZ327278 (ITS), MZ348538(LSU).
Notes: In the phylogenetic tree (Fig. 164), Skeletocutis subchrysella grouped with S. chrysella, S. delicata
Niemelä & Miettinen, S. exilis Miettinen & Niemelä and
S. kuehneri A. David. Morphologically, they all have
allantoid basidiospores; but S. chrysella differs from S.
subchrysella in its trimitic hyphal system and longer basidiospores (2.8–4.5 × 0.7–1 μm; Niemelä 1998); S. delicata
and S. exilis differ by having larger pores (3–6 per mm
in S. delicata, 3–5 per mm in S. exilis) and basidiopores
(3.2–4.2 × 1.1–1.4 µm in S. delicata, 3.2–3.9 × 0.9–1.1 µm
in S. exilis; Miettinen and Niemelä 2018); S. kuehneri differs by having thin and brittle basidiocarps and growth
on dead wood of Picea and Pinus (David 1982). Skeletocutis bambusicola, S. lepida, S. mopanshanensis and S.
yunnanensis were also discovered from Yunnan Province.
Skeletocutis bambusicola differs from the new species by
having smaller pores (8–11 per mm) and ellipsoid basidiospores (2.7–3.1 × 1.5–1.9 µm; Zhou and Qin 2012); S.
lepida differs by having half-resupinate basidiocarps
with ochraceous upper surface when dry (Korhonen et al.
2018); S. mopanshanensis differs by having larger pores
(4–5 per mm) and basidiospores (4.7–6.6 × 3.2–4.5 µm;
Wu et al. 2017); S. yunnanensis differs by having larger
basidiospores (3.4–4.5 × 1–1.2 µm; Bian et al. 2016).
Psathyrellaceae Vilgalys, Moncalvo & Redhead, in Redhead et al., Taxon 50(1): 226 (2001).
Wächter and Melzer (2020) based on phylogenetic and
morphological characteristics, introduced six new monophyletic genera to this family. Wijayawardene et al. (2022)
accepted 21 genera in Psathyrellaceae.
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◂ Fig. 164 Maximum likelihood tree illustrating the phylogeny of Skel-
etocutis based on the combined sequences dataset of ITS + nLSU.
Branches are labeled with maximum likelihood bootstrap higher
than 50%, parsimony bootstrap proportions higher than 50% and
Bayesian posterior probabilities more than 0.90 respectively. Bold
names = New species
Fig. 165 Microscopic structures of Skeletocutis subchrysella (drawn
from the holotype). a. Basidiospores; b. Basidia and basidioles; c.
Cystidioles; d. Section of dissepiment edge; e. Hyphae from trama; f.
Hyphae from subiculum. Scale bars: a = 5 µm; b–f = 10 µm
Coprinopsis cinerea (Schaeff.) Redhead, Vilgalys & Moncalvo, in Redhead et al., Taxon 50 (1): 227 (2001).
Index Fungorum number: IF474379; Faceoffungi number: FoF11681; Figs. 166, 167, 168
Saprobic on roots of Vigna unguiculata. Sexual morph:
Basidiomycetous. Basidiocarp 0.3–4 cm long, formed on
the roots of cowpea, mycelial strands encircled the host
tissues on the root region, and basidiocarp emerged upon
incubation in moist chamber. Pileus strongly convex, and/
or parasol-shaped to flat or depressed, hymenium borne
201
on gills, becoming deliquescent and inky, lamellae thin,
basidia unmodified, basidiospores 7.5–12.2 × 5 − 10 μm
diam., ballistosporic, blue-black, smooth with a distinct
germ pore. Asexual morph: not obesrved.
Cultural characteristics: On PDA medium, colony
were white and free from fruiting bodies till 5 days but
later, development of basidiocarp was noticed which later
enlarged and produced typical fruiting body. Upon examination of the basidiocarp from the PDA plates, microscopic examination of basidiospores revealed that they
were same in morphological features.
Material examined: India, Karnataka, Mysore, Doddamaragowdanahally, on infected leaves of Vigna
unguiculata (L.) Walp. (Fabaceae) placed on PDA as
secondary saprophytes, 18 May 2017 Mahadevakumar
(UOM-IOE-18/24), living culture (MD8).
Habitat: commonly found on dung or wood chips (Kamada 2002), roots and leaves of Vigna unguiculate (This
study).
Distribution: worldwide.
GenBank number: OM812073(ITS).
Notes: Coprinopsis species are known to grow on various
substrates as secondary saprophytes and are also known to
occur as primary components of soil and leaf litter. However,
an association of Coprinopsis species has not been reported
on any crop plants. Cowpea plants affected with a characteristic white cottony mycelium profusely grown on roots were
observed which were eventually wilted and dried off which
caused the death of the host plants. Infected materials were
collected and subjected to pathogen isolation. Few infected
parts were incubated at room temperature and observed
for mycelial development. To our surprise, a characteristic
mushroom fruiting bodies were developed. Therefore, the
spores were photographed and subjected for identification.
Here, the mushrooms are developed in three parts. Infected
roots collected from the field, root samples incubated at
room temperature and leaves incubated in a moist chamber
(in a Petri plate) showed the development of small basidiocarp. Upon microscopic examination, it was identified as
Coprinopsis sp. based on fruiting bodies, basidia, basidiospores, sterigmata and other associated structures. This
is the first time that C. cinerea is reported from Fabaceae,
Vigna unguiculata representing a new host record (Fig. 169).
Thelephorales Corner ex Oberw.
Notes: The order was established by Oberwinkler (1976)
based on the type family Thelephoraceae Chevall. Thelephorales is reported to incorporate two families: Thelephoraceae
and Bankeraceae Donk, and ten major clades: Amaurodon,
Boletopsis, Hydnellum/Sarcodon, Lenzitopsis, Odontia,
Phellodon/Bankera, Pseudotomentella/Polyozellus, Sarcodon, Thelephora/Tomentella and Tomentella (Stalpers 1993;
Vizzini et al. 2016). Twelve genera and about 800 described
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Fig. 166 Coprinopsis cinerea
associated with root rot
of cowpea: a Cowpea root
showing fungal mycelium. b
Close view of root showing
white mycelium. c Root sample
incubated at room temperature
showing the formation of young
basidiocarp. d Root samples
showing emergence of basidiocarps on incubation at room
temperature. e–f Close view of
basidiocarp showing characteristic feature of Coprinopsis cinerea. g Basidium of
Coprinopsis cinerea recorded.
Scale bar: g = 20 µm
species are accommodated within the order according to
Index Fungorum 2022a, b (http://www.indexfungorum.org).
Wijayawardene et al. (2022) accepted 14 genera in this order.
Thelephoraceae Chevall. [as’Thelephoreae’], Fl. Gén. Env.
Paris (Paris) 1: 84 (1826).
Notes: Thelephoraceae was proposed by Chevall (1826),
and Thelephora Ehrh. ex Willdwas regarded as its type
genus. Thelephoraceae, as a relatively large one, comprises
13
nine genera within the Thelephorales (Wijayawardene et al.
2022). Many species in the family, as ectomycorrhiza formers, are believed to be of great ecological importance in
maintaining the balance of terrestrial ecosystems (Haug
et al. 2005; Jakucs and Erős-Honti 2008; Kuhar et al. 2016).
Tomentella Pers. ex Pat., Hyménomyc. Eur. (Paris): 154
(1887).
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203
Fig. 167 Coprinopsis cinerea
developed on cowpea leaves: a
– e Cowpea leaves showing the
mycelial cords and development
of basidiomata upon incubation
at room temperature. f–h Mycelium colonizing the young root
system under in-vitro condition
Notes: Tomentella was validated by Patouillard (1887).
Species of the genus usually form cottony or spider web–likereproductive structures and grow on fallen wood, leaf litter, soil and other substrates (Larsen 1974; Tedersoo et al.
2003). Around 400 names have been recorded and about
200 species were described. The species of Tomentella are
reported to be widely distributed throughout the world and
have been immensely taxonomically studied from Eurasia, North America, South America, Australia, Asia and
WestAfrica (Thind and Rattan 1971; Larsen 1998; Agerer
et al. 2001; Yorou and Agerer 2008; Alvarez-Manjarrez
et al. 2015; Kuhar et al. 2016). During the investigation of
resupinate-thelephoroid fungi from the subtropical forests
in China, several Tomentella specimens were collected, and
two undescribed species have been identified using morphological characteristics and molecular phylogenetic analyses
(Fig. 173).
Tomentella exiguelata Y.H. Mu & H.S. Yuan, sp. nov.
Index Fungorum number: IF900080; Facesoffungi number: FoF13398; Figs. 170, 171, 172
Etymology: Refers to the presence of slightly thick-walled
basidia.
Holotype: IFP 019495
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204
Fig. 168 Micro-morphology of basidiospores of Coprinopsis
cinerea associated with root rot of cowpea: a-c Pure cultures of
Coprinopsis cinerea established on PDA medium. d-f Microscopic
view of basidiospores of Coprinopsis cinerea
Basidiocarps annual, resupinate, adherent to the substrate,
mucedinoid, without odour or taste when fresh, 0.5–1 mm
thick, continuous. Hymenophoral surface granulose, grayish brown to light brown (6D3–6D4) and concolorous
withsubiculum when dry. Sterilemargin often determinate,
farinaceous, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
clamped and rarely simple septate, thin- to slightly thickwalled, moderately branched, 3–5μmdiam, without encrustation, yellow in KOH, cyanophilous in sightly thick-walled
hyphae, inamyloid. Subhymenial hyphae clamped and rarely
simple septate, thin-to slightly thick-walled, frequently
branched, 3–5 μm diam; hyphal cells more or less uniform, yellow in KOH, cyanophilousin sightly thick-walled
hyphae, inamyloid. Cystidia absent. Basidia10–47 μm long
and 4–8 μm diam at apex, 3–5μmat base, with a clamp connection or simple septate at base, utriform, thin-walled and
rarely slightly thick-walled, not stalked, sinuous, yellow in
KOH, yellow in distilled water, 4-sterigmate; sterigmata
2.5–4.5μmlong and 0.5–1 μm diam at base. Basidiospores
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thick-walled, (6.2–)7.1–8.1(–8.3) × (5.4–)5.8–6.9(–7.1) μm,
L = 7.33 μm, W = 6.21 μm, Q = 1.18–1.21 (n = 60/2), subglobose, triangular or lobed in frontal view and subglobose to
ellipsoid in lateral view, echinulate to aculeolate, yellow in
KOH and in distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up
to 2.5 μm long.
Material examined: China, Zhejiang Province, Kaihua
County, Gutianshan National Nature Reserve, on fallen
angiosperm debris, 24 July 2018, Yuan 12805 (IFP 019495,
holotype); on fallen angiosperm branch, 25 July2018, Yuan
12900 (IFP 019496, paratype).
GenBank numbers: IFP 019495- MZ329771(ITS),
MZ329775 (LSU).
IFP 019496- MZ329772 (ITS), MZ329776 (LSU)
Notes: Tomentella exiguelata forms a close phylogenetic
relationship with T. galzinii and T. subtestacea (Fig. 173).
Tomentella galzinii and T. exiguelata share the following similar morphological and anatomical characteristics:
mucedinoid basidiocarps adherent to the substrate, granulose
hymenophoral surface, farinaceous sterile margin, similarwide subhymenial hyphae, and not stalked and sinuous
basidia. However, T. galzinii has discontinuous and dull
green to olive-brown basidiocarps, indeterminate sterile
margin, and the presence of cystidia (Bourdot and Galzin
1924). Tomentella subtestacea resembles T. exiguelata in
mucedinoid basidiocarps adherent to the substrate, clamped
and rarely simple septate subicular hyphae, thin-to slightly
thick-walled subhymenial hyphae, utriform and not stalked
basidia and echinulate basidiospores of similar length. However, T. subtestacea can be distinguished by reddish-brown
to grayish buff hymenophoral surface, arachnoid sterile margin, and the presence of cystidia (Svrček 1958).
Tomentella fuscoaraneosa Y.H. Mu &H.S. Yuan, sp. nov.
Index Fungorum number: IF900079; Facesoffungi number: FoF13399; Figs. 174, 175, 176
Etymology: Refers to the brown and arachnoid
basidiocarps.
Holotype: IFP019493
Basidiocarps annual, resupinate, separable to the substrate, arachnoid, without odour or taste when fresh,
0.8–1.5 mm thick, continuous. Hymenophoral surface
smooth, light brown to brown (7D8–7E8) and concolorous
with subiculum when dry. Sterile margin often determinate,
byssoid, concolorous with hymenophore. Rhizomorphs present in subiculum and margin, 10–35 μm diam; rhizomorphic surface more or less smooth; hyphae in rhizomorphmonomitic, undifferentiated, of type B, compactly arranged
and of uniform; single hyphae with both clamps and simple
septa, thick-walled, unbranched 2–3 μm diam, yellow in
KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphaeclamped and rarely simple septate,
Fungal Diversity (2022) 117:1–272
205
Fig. 169 Phylogenetic tree for species of Coprinopsis generated from
maximum likelihood (RAxML) based on ITS region. The Maximum
likelihood bootstrap value ≥ 50% are given at the nodes. The newly
generated sequences are in blue. The tree is rooted to Parasola auricoma (SZMC.NL:0087) and Parasola conopilea (LO186.02)
thin- to slightly thick-walled, frequently branched, 2–3 μm
diam, withoutencrustation, yellow in KOH, cyanophilous in
sightly thick-walled hyphae, inamyloid. Subhymenial hyphae
clamped and rarely simple septate, thin-walled, occasionally
branched, 2.5–4 μm diam, without encrustation; hyphal cells
short and not inflated, yellow in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia10–65 μm long and 4–7 μm
diam at apex, 2–4 μm at the base, with a clamp connection
or simple septate at the base, utriform, thin-walled, stalked,
sinuous, occasionally with transverse septa, yellow in KOH,
yellow in distilled water, 4-sterigmate; sterigmata 2–5 μm
long and 1–2 μm diam at the base. Basidiospores thickwalled, (5–)5.5–7.5(–8) × (4–)4.5–6(–7) μm, L = 6.29 μm,
W = 5.32 μm, Q = 1.12–1.18 (n = 60/2), subglobose to ellipsoid in frontal view and subglobose to ellipsoid in lateral
view, echinulateto aculeolate, yellow in KOH, yellow in
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 1.5 μm long.
Material examined: China, Zhejiang Province, KaihuaCounty, Gutianshan National Nature Reserve, on fallen
angiosperm branch, 25 July 2018, Yuan 12875(IFP 019493,
holotype); on fallen angiosperm branch, 25 July2018, Yuan
12910 (IFP 019494, paratype).
GenBank numbers: IFP 019493-MZ329769 (ITS),
MZ329773 (LSU).
IFP 019493-MZ329770 (ITS), MZ329774 (LSU)
Notes: Tomentella fuscoaraneosa is closely related to T.
aureomarginata in the phylogeny (Fig. 173). In morphology,
T. fuscoaraneosa is similar to T. aureomarginata in having a
determinate and byssoid sterile margin, the presence of type
B rhizomorphs, monomitic generative hyphae with clamps
and simple septa, and the absence of cystidia. However, T.
aureomarginata differs from T. fuscoaraneosa by having
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Fig. 170 A basidiocarp of Tomentella exiguelata (IFP 019495, holotype). Scale bars = 0.25 cm
pelliculose basidiocarps adherent to the substrate, golden
brown to yellowish-brown hymenophoral surface turning
darker when dry, not stalked basidia and wider basidiospores
(6–6.5 μm vs. 4.5–6 μm in T. fuscoaraneosa) with shorter
echinuli (up to 1 μm vs. up to 1.5 μm in T. fuscoaraneosa)
(Yuan et al. 2020). Tomentella fuscoaraneosa shares common features with T. brunneoflava in arachnoid and continuous basidiocarps, the presenceof rhizomorphs and clamped
and rarely simple septate subicular hyphae and short and
not inflated subhymenial hyphae. However, T. brunneoflava
differs from T. fuscoaraneosa by having brownish-yellow
basidiocarps adherent to the substrate and clavate and not
stalked basidia (Yuan et al. 2020).
Agaricales genera incertae sedis
Gerronema Singer, Mycologia 43(5): 599 (1951).
Gerronema is a minor genus of lignicolous agaric and
is distributed worldwide (Singer 1986). There are 66 epithets (excluding synonyms) listed in the Index Fungorum
(2021). Gerronema. melanomphax Singer is the specific
type species. Historically, different circumscriptions of
Fig. 171 SEM of basidiospores
of Tomentella exiguelata (IFP
019495, holotype)
13
Fig. 172 Microscopic structures of Tomentella exiguelata (IFP
019495, holotype). a Section through a basidiocarp. b Basidiospores
in frontal view. c Basidiospores in lateral view
Gerronema based on pigmentations established by Singer
(1964) and Bigelow (1970) have been problematic. Subsequently, Redhead (1986) and Norvell et al. (1994)
restricted the delimitation of Gerromena and characterized
its descriptive features as elastic to fleshy, omphalinoid to
clitocyboid basidiomata, white spore print, smooth, thinwalled, inamyloid basidiospores, cutis pileipellis often
with intracellular pigment, sarcodimitics trama tissue and
lignicolous habit. This circumscription correlates with the
molecular phylogenetic analysis of the combined nrITS
Fungal Diversity (2022) 117:1–272
207
Fig. 173 Phylogram generated from maximum likelihood analysisbased on combined ITS and LSU sequence data. Related sequences
were obtained from GenBank and Unite. One hundred and ten strains
are included in the combined sequence analyses, which comprise
1451 characters withgaps. Single gene analyses were also performed
and topology andclade stability compared from combinbed gene analyses. Odontia ferruginea (TU110988) is used as the outgroup taxon.
Bootstrap support values for ML ≥ 50% are given. The newly generated sequences are in bold
and nrLSU dataset and revealed that Gerronema is monophyletic and forms a clade with Megacollybia and Trogia
in Hydropoid calde (Antonín et al. 2019).
Etymology: ‘atro’ = dark; ‘virens’ = green, refers to the
dark green colour of basidiomata.
Holotype: BKF10264
Pileus 21–51 mm diam., convex with depress center
when young and deeply infundibuliform in age, elastic,
glabrous, translucent-striate to striate at margin, dull green
(27E3) at center, dark green (27F3) at margin, grey (5E1) to
brownish grey (5E2) in old specimen; Lamellar decurrent,
Gerronema atrovirens Wannathes, N. Suwannarach, J.
Kumla, Phonrob & S. Lumyong, sp.nov.
MycoBank number: MB840183; Facesoffungi number:
FoF10685, Figs. 177a, 178
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208
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Fig. 174 A basidiocarp of Tomentellafuscoaraneosa (IFP 019493,
holotype). Scale bars = 0.25 cm
subdistant to close (24–28) with 1–2 series of lamellulae,
narrow (up to 1.5 mm), withe (27A1), non-marginate; Stipe
14–36 × 2.5–4.0 mm, cylindrical, slightly broadened at base,
usually flatten, flexuose, hollow, elastic, central, pubescent,
greenish grey (27F2) at apex fading paler to brownish grey
(5C2) at base, basal mycelium. Context thin, elastic. Odor
and taste not distinctive.
Basidiospore 7–8 × 4–5 µm [x = 7.56 ± 0.51 × 4.36 ± 0.49,
Q = 1.4–2.0, q = 1.75 ± 0.21, n = 25, s = 3] ellipsoid, smooth,
hyaline, inamyloid, thin-walled. Basidia 21–27 × 6–7 μm,
clavate, with 4 sterigmata, sometime with 2 sterigmata, thin-walled, inamyloid. Cheilocystidia abundant,
21–51 × 5–11 µm, cylindrical with 1–2 slight constrictions,
flexuose, irregular in shape, sometimes 2-celled, hyaline,
inamyloid, thin-walled. Pleurocystidia absent. Lamellar
trama subregular to interwoven, arranged in two directions,
hyphae 4–8 µm diam., cylindrical, smooth, hyaline, inamyloid, thin-walled, gelatinous. Pileipellis composed with cutis
of repent hyphae, radially arrangement, 3–10 µm, cylindrical, incrusted, greyish green in KOH, inamyloid, thinwalled, true pileocystidia absent. Pileus trama sarcodimitic,
Fig. 175 SEM of basidiospores
of Tomentella fuscoaraneosa
(IFP 019493, holotype)
13
Fig. 176 Microscopic structures of Tomentella fuscoaraneosa (IFP
019493, holotype). a Hyphae from a rhizomorph. b Section through
a basidiocarp. c Basidiospores in frontal view. d Basidiospores in lateral view
subregular, composed of 2 type of hyphae: a) sarco-hyphae,
elongate fusoid cell 162–195 × 5–15 µm, hyaline, smooth,
inamyloid, thick-walled (up to 1 µm); b) generative hyphae
2–6 µm wide, cylindrical, branched, hyaline, smooth, inamyloid, thin-walled. Stipitipellis cutis, hyphae 3–8 µm diam.,
parallel, cylindrical, greenish brown in KOH, smooth, inamyloid, thin-walled. Stipe trama sarcodimitic, subparallel,
Fungal Diversity (2022) 117:1–272
Fig. 177 Basidiomata and habit. a Gerronema atrovirens (holotype: BKF10264), b G. kuruvense (BKF10266), c G. flavum (holotype: BKF10253), Scale bars = 10 mm. Photographs by N. Wannathes
composed 2 type of hyphae: a) sarco-hyphae, elongate fusoid
cell 125–200 × 3–25 µm, hyaline, smooth, inamyloid, thickwalled (up to 1 µm); b) generative hyphae 2–13 µm wide,
cylindrical, branched, hyaline, smooth, inamyloid, thinwalled. Caulocystidia abundant, 19–34 × 5–9 µm, cylindrical
with 1–2 constrictions, flexuose, irregular in shape, hyaline
to pale green in KOH, inamyloid, thin-walled. Clamp connections present in all tissues.
Habit, habitat and known distribution: Gregarious on
decayed bamboo wood, known only from Thailand.
Material examined: Thailand, Sukhothai Provinve, Si
Satchanalai National Park, Natural trail, 22 Aug 2020,
collector N Wannathes N Suwannarach and J Kumla,
BKF10264 (holotype).
Additional material examined: Thailand, Sukhothai Province, Si Satchanalai National Park, Natural trail, 23 Aug
2020, collector N Wannathes N Suwannarach and J Kumla,
BKF10265, NW1372 (isotype).
GenBank numbers: BKF10264- MZ452088(ITS),
MZ452671(LSU).
BKF10265- MZ452668(ITS), MZ452672 (LSU)
Notes: Gerronema atrovirens is characterized by a
medium size of omphaliod, elastic basidiomata, convex
with depress center to deeply infundibuliform, translucent
striate at margin, dull green pileus, decurrent, subdistant
lamellae, cylindrical, central, greenish grey stipe, ellipsoid
basidiospores with mean 7.6 × 4.4 µm, pileipellis composed
of incrusted hyphae with greyish green colour, present of
209
Fig. 178 Gerronema atrovirens (BKF10264, holotype). a Basidiomata, b Basidiospores, c basidium, d Cheilocystidia, e Caulocystidia f Pileipellis cell. Scale bars: a = 20 mm, b–e = 10 μm, f = 20 μm.
Drawing by W. Phonrob
irregular cylindrical cheilocystidia and cualocystidia, sarcodimitic trama tissue, and grow on decayed bamboo wood.
A new species is morphologically similar to G. cyathiforme
(Berk. & M.A. Curtis) Singer, species originally described
from a Neotropic, differs in forming a distinct radial stripes
pileus, absent of cystidia, and mahogany red (in KOH)
lamellar trama (Singer 1970). Gerronema atrovirens is
also closely related G. indigoticum T. Bau & L.N. Liu, a
green–blue species from subtropical China. The latter species differs in forming smaller pilei (9–16 mm wide) with
green–blue, shorter (20–27 × 8–12 µm), simple clavate
cheilocystidia, and simple clavate cualocystidia (Lui et al.
2019), and the phylogenetic analyses inferred from combined sequences (Fig. 179) confirmed that G. atrovirens is
closely allied with G. indigoticum and it distinct species
from related morphological species and other taxa in this
genus.
Gerronema flavum Wannathes, N. Suwannarach, J. Kumla,
Phonrob & S. Lumyong, sp.nov.
MycoBank number: MB 840184; Facesoffungi number:
FoF10686; Figs. 177c, 180
Etymology: ‘flavum’ = yellow, refers to the colour of
basidiomata.
Holotype: BKF10253
Pileus 4–11 mm diam., convex, umbonate when young
and plano-convex with depress in center to infundibuliform
in age, glabrous, radially fibrillose when young, translucent
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Fig. 180 Gerronema flavum (BKF10253, holotype). a Basidiomata,
b Basidiospores, c Basidium, d Pleurocystidia, e Cheilocystidia, f
Caulocystidia, g Pileipellis cell. Scale bars: a = 20 mm, b–f = 10 μm,
g = 20 μm. Drawing by W. Phonrob
Fig. 179 Phylogenetic tree derived from maximum likelihood analysis of a combined ITS and LSU genes of 21 sequences and the
aligned dataset was comprised of 1700 characters including gap. The
average standard deviation of the split frequencies of the BI analysis was 0.00612. Togia infundibuliformis KUN HKAS56709 and
T. venenata KUN HKAS56679 were used as outgroup. Numbers
above branches are the bootstrap statistics percentages (left) and
Bayesian posterior probabilities (right). Branches with support values ≥ 70%/0.90 are shown at each branch and the bar represents 0.1
substitutions per nucleotide position. Hyphen (-) represents support
values ≤ 70%/0.95. Ex-type strains are in black bold. The newly generated sequences are indicated in blue and bold type species
striate to striate at marging in age, dull, light yellow (4A5)
overall when young, sunflower yellow (4A7) overall in
age, hygrophanous, become yellowish white. Lamellar
decurrent, subdistant (14–16) with 2–3 series of lamellulae, narrow, yellowish withe (4A2), non-marginate; Stipe
4–19 × 1–2 mm, tapering upward when young, cylindrical
in age, elastic, hollow, central, flexuose, pubescent, yellowish withe (4A2) overall. Context thin, Odor and taste not
distinctive.
Basidiospore
7–8(–9) × 4–5(–6)
μm
[x = 7.68 ± 0.63 × 4.68 ± 0.56, Q = 1.3–2.25, q = 1.66 ± 0.23,
n = 25, s = 1] broadly ellipsoid, smooth, hyaline, inamyloid,
thin-walled. Basidia 25–32 × 6–9 μm, clavate, with 4 sterigmata, thin-walled, inamyloid. Cheilocystidia abundant,
25–40 × 5–8 µm, knobby cylindrical to clavate, irregular
in shape, hyaline, inamyloid, thin-walled. Pleurocystidia
abundant, 24–33 × 9–11 µm, clavate to cylindrical, sometime
13
knobbed, hyaline, inamyloid, thin-walled. Lamellar trama
interwoven, hyphae 3–6 µm diam., cylindrical, smooth, hyaline, inamyloid, thin-walled. Pileipellis composed with cutis
of repent hyphae, radially arrangement, 4–15 µm, cylindrical, non-incrustation, hyaline in KOH, inamyloid, thinwalled, true pileocystidia absent. Pileus trama sarcodimitic,
interwoven, composed of 2 type of hyphae: a) sarco-hyphae,
elongate fusoid cell 50–60 × 10–14 µm, hyaline, smooth, inamyloid, thin-walled; b) generative hyphae 3–11 µm wide,
cylindrical, branched, hyaline, smooth, inamyloid, thinwalled. Stipitipellis cutis, hyphae 3–10 µm diam., parallel,
cylindrical, hyaline to pale yellow in KOH, smooth, inamyloid, thin-walled. Stipe trama sarcodimitic, parallel, composed 2 type of hyphae: a) sarco-hyphae, elongate fusoid cell
87–165 × 4–15 µm, hyaline, smooth, inamyloid, thick-walled
(1–4 µm); b) generative hyphae 3–12 µm wide, cylindrical,
branched, hyaline, smooth, inamyloid, thin-walled. Caulocystidia abundant, 28–47 × 7–9 µm, cylindrical to clavate,
irregular in shape, flexuose, hyaline KOH, inamyloid, thinwalled. Clamp connections present in all tissues.
Habit, habitat and known distribution: Gregarious on
decayed wood, known only from Thailand.
Material examined: Thailand, Nakhon Ratchasima
Provinve, Khao Yai National Park, trial to Pha Kluai Mai
waterfall, 21 Sep 2018, collector N Wannathes, N Suwannarach J Kumla, S Lumyong, BKF10253 (holotype).
GenBank numbers: BKF10253- MZ1452142 (ITS),
MZ452170 (LSU).
Fungal Diversity (2022) 117:1–272
Fig. 181 Gerronema keralense (BKF10263) a Basidiomata, b Basidiospores, c Basidium, d Cheilocystidia, e Caulocystidia f Pileipellis
cell. Scale bars: a = 20 mm, b–e = 10 μm, f = 20 μm. Drawing by W.
Phonrob
Notes: Gerronema flavum is characterized by a small
omphaliod basidiomata that appears convex with depressed
center infundibuliform, radially fibrillose, glabrous, yellow
pileus, decurrent, subdistant lamellae, cylindrical, central,
yellowish white stipe, broadly ellipsoid basidiospores with
mean dimensions of 7.7 × 4.7 µm. The pileipellis is composed of hyaline with non-incrusted hyphae, the presence
of irregular cylindrical cheilo-, pleuro- and cualocystidia,
and sarcodimitic trama tissue. Gerronema kuruvense K.P.D.
Latha & Manim and G. subchrysophyllum (Murrill) Singer
are morphologically similar to a new species, namely Gerronema kuruvense, which was originally described from
tropical India. It is distinguished by forming orange yellow pileus, bigger basidiospores with mean dimensions
of 9.5 × 5.9 µm., an absence of cheilo- and pleurocystidia
and the presence of diverticulate caulocystidia (Latha et al.
2018). Gerronema subchrysophyllum, a North American
species, differs by having a larger basidiomata of nearly
double the size (pilei 4–21 mm wide and stipe 4–32 mm
long), simple clavate cheilocystidia and a lack a pleurocystidia (Singer 1970). The phylogenetic analyses inferred from
combined sequences (Fig. 179) confirmed that G. flavum is a
distinct species. This was further confirmed via an examination of other related morphological species and other taxa
within this genus.
Gerronema keralense K. P. D. Latha & Manim Phytotaxa
364 (1): 85–88 (2018).
211
Index Fungorum number: IF824928; Facesoffungi number: FoF10687; Fig. 181
Pileus 20 mm diam., infundibuliform, appressed-fibrillose, brownish orange to light brown (5C5-5D5); Lamellar
decurrent to deeply decurrent, distant (12) with 2–3 series
of lamellulae, greyish yellow (4B6) up to 1.5 mm.; Stipe
21 × 0.75 mm, central, cylindrical, pruinose, slightly broadened at base, solid, greyish yellow (4B6) overall. Context
thin, Odor and taste not distinctive.
Basidiospore
7–9 × (3–)4–5
μm
[x = 7.76 ± 0.78 × 4.4 ± 0.65, Q = 1.4–2.3, q = 1.79 ± 0.25,
n = 25, s = 1] ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia 25–34 × 5–7 μm, clavate, with 4 sterigmata, thin-walled, inamyloid. Cheilocystidia scattered,
27–31 × 5–8 µm, clavate, flexuose, irregular in shape, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Lamellar
trama subregular, hyphae 4–15 µm diam., smooth, hyaline,
inamyloid, thin-walled. Pileipellis cutis with pileocystidia,
3–12 µm, cylindrical, hyaline in KOH, inamyloid, inclusion cytoplasm turns to brown to light brown in Melzer's
reagent, thin-walled. Pileus trama sarcodimitic, subregular,
composed of 2 type of hyphae a) sarco-hyphae, elongate
fusoid cell 98–196 × 10–14 µm, hyaline, smooth, inamyloid,
thin-walled; b) generative hyphae 2–5 µm wide, cylindrical, branched, hyaline, smooth, inamyloid, thin-walled.
Stipitipellis cutis, hyphae 3–6 µm diam., parallel, cylindrical, pale yellow in KOH, smooth, inamyloid, slightly
thick-walled. Stipe trama sarcodimitic, parallel, composed
2 type of hyphae: a) sarco-hyphae, elongate fusoid cell
74–230 × 7–10 µm, hyaline, smooth, inamyloid, slightly
thick-walled; b) generative hyphae 3–7 µm wide, cylindrical,
branched, hyaline, smooth, inamyloid, slightly thick-walled.
Caulocystidia 28–47 × 7–9 µm, agglutinated, cylindrical to
clavate, hyaline in KOH, inamyloid, slightly thick-walled.
Clamp connections present in all tissues.
Habit, habitat and known distribution: Solitary on
decayed wood, known from topical India and Thailand.
Material examined: Thailand, Sukhothai Province, Si
Satchanalai National Park, Natural trial, 15 June 2019, collector N Wannathes, N Suwannarach J Kumla, S Khuna,
BKF10263.
GenBank numbers: BKF10263- MZ452107 (ITS),
MZ452144 (LSU).
Notes: Gerronema keralense is characterized by a
medium-sized omphaliod basidiomata, infundibuliform,
brownish orange to light brown pileus, decurrent to deeply
decurrent, distant, greyish yellow lamella, cylindrical,
central, greyish yellow stipe, ellipsoid basidiospores
with mean dimensions of 7.8 × 4.4 µm, pileipellis with
pileocystidia, non-incrusted hyphae, cytoplasmic inclusion that turned from brown to light brown in Melzer's
reagent, the presence of irregular cylindrical cheilo-,
cualocystidia and sarcodimitic trama tissue. Our Thai
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212
Fig. 182 Gerronema kuruvense (BKF10266), a Basidiomata, b
Basidiospores, c Basidium, d Cheilocystidia, e Caulocystidia f
Pileipellis cell. Scale bars: a = 20 mm, b–e = 10 μm, f = 20 μm. Drawing by W. Phonrob
description is consistent with Gerronema keralense that
was originally described from India, except for the type
specimen that formed an applanate with a slight central
depression pileus (Latha et al. 2018). Notably, these
variations may be caused by a mutuality of basidiomata.
Gerronema keralense is morphologically similar to G.
kuruvense K.P.D. Latha & Manim, but the latter differs
by having bigger basidiospores with mean dimensions of
9.1 × 4.9 µm, a cytoplasmic inclusion of pileocystidia that
never changes colour with Melzer's reagent and caulocystidia that are not agglutinated (Latha et al. 2018).
Gerronema kuruvense K. P. D. Latha & Manim. Phytotaxa
364 (1): 82–85 (2018).
Index Fungorum number: IF824927; Facesoffungi number: FoF10688; Figs. 177b, 182
Pileus 27 mm diam., convex with depress center,
appressed-fibrillose, pale yellow (3A3); Lamellar subdecurrent, distant (16) with 2–3 series of lamellulae, narrow,
light yellow (3A5) up to 2 mm.; Stipe 25 × 1 mm, central,
cylindrical, pubescent, slightly broadened at base, hollow,
yellowish white (3A2) overall. Context thin, Odor and taste
not distinctive.
Basidiospore 8–10(–11) × 4–5(–6) μm
[x = 9.08 ± 0.81 × 4.88 ± 0.43, Q = 1.6–2.25, q = 1.89 ± 0.25,
n = 25, s = 1] ellipsoid, smooth, hyaline, inamyloid, thinwalled. Basidia 22–36 × 7–8 μm, clavate, with 2 sterigmata, sometime with 3–4 sterigmata, thin-walled, inamyloid. Cheilocystidia scattered, 21–51 × 5–11 µm, clavate,
flexuose, irregular in shape, sometimes 2-celled, hyaline,
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inamyloid, thin-walled. Pleurocystidia absent. Lamellar trama interwoven, arranged in two directions, hyphae
3–14 µm diam., cylindrical, smooth, hyaline, inamyloid,
thin-walled. Pileipellis cutis with scattered ascending pileocystidia, 4–14 µm, cylindrical, hyaline in KOH, inamyloid,
thin-walled. Pileus trama sarcodimitic, subregular to interwoven, composed of 2 type of hyphae: (a) sarco-hyphae,
elongate fusoid cell 94–212 × 10–14 µm, hyaline, smooth,
inamyloid, thin-walled; (b) generative hyphae 3–7 µm wide,
cylindrical, branched, hyaline, smooth, inamyloid, thinwalled. Stipitipellis cutis, hyphae 3–8 µm diam., parallel,
cylindrical, hyaline in KOH, smooth, inamyloid, slightly
thick-walled (up to 0.5 µm). Stipe trama sarcodimitic, parallel, composed 2 type of hyphae: a) sarco-hyphae, elongate
fusoid cell 80–230(– over 250) × 8–15 µm, hyaline, smooth,
inamyloid, thick-walled (1–2 µm); b) generative hyphae
3–6 µm wide, cylindrical, branched, hyaline, smooth, inamyloid, thin-walled. Caulocystidia abundant, 28–47 × 7–9 µm,
cylindrical to clavate, irregular in shape, flexuose, hyaline
in KOH, inamyloid, slightly thick-walled (up to 0.5 µm).
Clamp connections present in all tissues.
Habit, habitat and known distribution: Solitary on
decayed wood, known from topical India and Thailand.
Material examined: Thailand, Sukhothai Province, Si
Satchanalai National Park, trial to Tad Duan waterfall, 30
Aug 2020, collector N Wannathes, N Suwannarach J Kumla,
S Khuna, W Phonrob, S Tabtan, BKF10266.
GenBank numbers: BKF10266-MZ452090 (ITS),
MZ452669(LSU).
Notes: Gerronema kuruvense is characterized by a
medium-sized omphaliod basidiomata that is convex with
a depressed center. It has a pale-yellow pileus that is subdecurrent, distant, with light yellow lamella, cylindrical,
central, yellowish white stipe, ellipsoid basidiospores with
mean dimensions of 9.1 × 4.9 µm. The pileipellis is composed of hyaline with pileocystidia, non-incrusted hyphae,
the presence of irregular cylindrical cualocystidia and seldomly present cheilocystidia and sarcodimitic trama tissue.
The Thai specimen is almost indistinguishable from Gerronema kuruvense, which was originally described from
topical India. The holotype forms a smaller size basidiomata
(pilei 4–11 mm wide and stipe 3–18 mm long) and lacks
cheilocystidia (Latha et al. 2018). Gerronema kuruvense is
morphologically similar to G. strombodes (Berk. & Mont.)
Singer, but differs by forming larger basidiomata (pilei
25–80 mm wide and stipe of 30–60 × 2–6.8 mm) greyish
white pileus, pileipellis lacking pileocystidia and a complete
lack of caulocystidia (Singer 1970).
Tricholomopsis Singer, Schweiz. Z. Pilzk. 17: 56 (1939).
Tricholomopsis was established to accommodate a group
of saprophytic tricholomatoid fungi which have a fibrillose
or squamulose pileus, inamyloid smooth basidiospores, and
Fungal Diversity (2022) 117:1–272
213
Fig. 183 Tricholomopsis lechatii (LIP0202264,
holotype). a, b Basidiocarp c
Basidiospores d Basidia e One
hymenophoral hyphid. f Cheilocystidia. g Pileus covering. h
Pileus margin, radial section.
Drawings and macrophotos by
R. Courtecuisse, microphotos
by P.-A. Moreau
a sterile lamella edge covered with large prominent cheilocystidia (Mao et al. 2021). There are 76 species epithets in
the index fungorum for this genus.
Tricholomopsis lechatii Courtec., S. Dumez, S. Welti &
P.-A. Moreau, sp. nov.
MycoBank number: MB843163, Facesoffungi number:
FoF13397; Fig. 183
Etymology: The species is warmly dedicated to our friend
Christian Lechat (1952–2022), who recently and suddenly
died. He was an eminent specialist of the Hypocreales
worldwide and a very enthusiastic and faithful member of
the numerous field trips organized in the Tropics by one of
us (RC). He was present when this species was collected.
Holotypus: 0202264 (LIP).
Pileus 10–14 mm diam, slightly convex or almost flat,
showing a little umbilic (not deep) in the centre or slightly
eccentric; shape regular, circular or almost so, sometimes
slightly elliptical when seen from above. Surface remarkably
dull, finely tomentose or even velvety under lens. Color very
special, dirty olivaceous with ochraceous hue at center but
more yellow at the margin, which is slightly enrolled or at
least very obtuse and scarcely pectinate at the extreme edge.
Lamellae adnate, rather crowded, with two rather regular
rows of smaller gills, thin. Color rather bright and deep,
13
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Fungal Diversity (2022) 117:1–272
Fig. 184 Phylogenetic analyses were conducted online at www.phylo
geny.fr (Dereeper et al. 2008). Multiple sequence alignments were
performed with MUSCLE v. 3.7 (Edgar 2004). Maximum likelihood
(ML) phylogenetic analysis was achieved with PhyML v. 3.0 (Guindon et al. 2010), using the GTR + I + Γ model of evolution and the
Shimodaira Hasegawa version of the approximate likelihood-ratio test
(SH-aLRT) of branch support (Anisimova et al. 2011). Phylogram
was built using TreeDyn 198.3 (Chevenet et al. 2006) and edited with
Inkscape 0.91 (https://inkscape.org/fr). Newly generated sequences
for this study are in bold. The tree is rooted by a sequence of Pluteus
atromarginatus (Pluteaceae)
mustard yellow. Edge rather thick, clearly bearing a row
of small, densely crowded, aggregates of crystals. Stipe up
to 15 × 2 mm, slightly excentrical, cylindraceous, more or
less narrowly fistulose, dull yellow, paler than the gills but
sometimes pale brownish toward mid-length, arising from
a thin greyish mycelial patch of subiculum (up to 35 mm
diam.), which sometimes climbs up to the stipe up to a few
millimeters, thus greyish at the basis. Context yellowish in
the stipe, deeper yellow in the cap. Smell none. Taste not
recorded.
Basidiospores 6.2–7.5 × 4.8–5 µm, rather broad, elliptical
to slightly tear-shaped, apex broader and generally clearly
rounded (not elongate nor conical). Apiculus small, faintly
distinct. Content with many droplets (sometimes a large one)
and confuse, cloudy or punctate around them. Wall smooth,
inamyloid, not cyanophilic. Basidia 20–28 × 5–8 µm,
2-spored (very few 1-spored and only one 3-spored seen)
with very long and sharp sterigmata (up to 10 µm), containing large droplets, the bigger ones at the top. Clamp present
at the base of all basidia. Subhymenium ramose, rather thick
with short, clamped hyphae up to 1.5–2.5 µm wide, somewhat wavy with some longer hyphae parallel to the hymenium. Hymenophoral trama parallel in a wavy arrangement,
weakly interwoven, made of clamped hyphae 4–8 µm wide.
Edge sterile; cheilocystidia very numerous and prominent,
cylindrical to slightly clavate, often regularly thickened
toward apex, 35–45 × 8–12 µm, thin-walled, hyaline; base
clamped, originated from simple or rarely forked hyphae.
Pleurocystidia absent; facial sterile cells present as sparse
slender “hyphids” cylindrical or slightly thickened at apex,
up to 4 µm wide. Pileipellis a trichoderm of more or less
straightly erected, unicellular or sometimes articulate elements, rather densely arranged, 30–70 × 7–15 µm, the apex
mostly rounded or sometimes irregular, rarely mucronate to
appendiculate or exceptionally forked; pigmentation epiparietal and vacuolar: wall yellowish, very finely incrusting
(minutely punctate or finely marked with transverse zebra
depending on the focus made on the cell wall) in all parts
13
Fungal Diversity (2022) 117:1–272
215
Fig. 185 Sporisorium anadelphiae-leptocomae (SOMF
30250, holotype). a, b Habit
(black arrows in a and b show
sori). c Spores in LM. d Spores
and sterile cells in LM. e–h
Spores in SEM. i Spores and
sterile cells in SEM. Scale bars:
a, b = 0.5 cm, c, d = 10 μm,
e–i = 5 μm
of the basidiome; vacuolar pigment abundant in subpellis,
also present in broad elements of suprapellis, brown-yellow.
Habitat: Saprobic on rotten wood of unidentified Angiosperm, wet tropical forest, along a wet depression. So far
only known from the type locality, French Guiana.
Material examined: French Guiana: Saül, Roche Bateau
trail, 23 Aug. 2018, R. Courtecuisse & C. Lechat, RC/
Guy18.018 (LIP 0202264, holotype).
GenBank numbers: LIP 0202264-OM793061 (ITS),
OM793062 (LSU).
Notes: Tricholomopsis lechatii phylogenetically belongs
to the T. aurea-complex, a pantropical group of collybioid
species so far rather poorly documented in molecular databases (Fig. 184). Tricholomopsis aurea was originally
described from DR Congo (as Marasmius aureus; Beeli,
1928), and recently transferred in the genus Tricholomopsis
by Desjardin and Perry (2017) based on recent collections
from São Tomé. Although the species is reported as common
in many tropical regions (GBIF data: https://www.gbif.org/
en/species/332554), only few sequences from the neotropics
are available, and the ITS marker indicates only faint differences between African and American collections (Desjardin
and Perry 2017, consider them as conspecific at this time).
The new species described here differs significantly from
T. aurea, by a pileipellis of trichodermioid structure, with a
well-differentiated subpellis and a distinct vacuolar pigment
(responsible of the grey-brown tomentum on pileus), whilst
T. aurea has a thin cutis-like structure, resulting in a glabrous, golden yellow surface with ± inflate terminal elements
with granular intracellular pigmentation (Pegler 1983).
Only few neotropical species of Tricholomopsis were
recorded in the Neotropics, and none can be attributed to
T. lechatii. Tricholomopsis tropica Dennis from Trinidad
is devoid of yellow tinge in context and displays abundant
hymenial gloeocystidia (Pegler 1983), what makes its classification in this genus somewhat doubtul; T. atrogrisea Pegler
from Martinique, possibly more related to T. lechatii, has
a gelatinized pileipellis made of adpressed hyphae; T. elegans Dennis (Dennis 1961) could be the closest relative of
T. lechatii but differs at least by smaller, subglobose spores
(4.5–5 µm long) and 4-spored basidia. No species described
so far display either the spectacular pale grey mycelial patch
observed in T. lechatii, nor its conspicuous 2-spored basidia
(Fig. 183).
Ustilaginomycotina Doweld.
We follow the latest treatment and updated account of
Ustilaginomycotina in Begerow and McTaggart (2018).
13
216
Fungal Diversity (2022) 117:1–272
Fig. 186 Most likely tree generated using maximum likelihood
analysis (RAxML 8.2.11, Stamatakis 2014) based on concatenated
MAFFT v7.450 (Katoh and Standley 2013) alignments of ITS and
LSU dataset. The tree is rooted with Mycosarcoma maydis Bref. and
M. mackinlayi (McTaggart & R.G. Shivas) McTaggart et al. Values
at nodes indicate bootstrap values inferred by 1000 replicates; only
values ≥ 60% are shown
Ustilaginomycetes R. Bauer et al.
The classification of the orders in Ustilaginomycetes follows Begerow and McTaggart (2018).
Sporisorium Ehrenb. ex Link, in Willdenow, Sp. pl., Edn 4
6(2): 86 (1825).
Sporisorium is a grass-infecting genus of smut fungi. It is
characterized by sori formed in different parts of the inflorescence or destroying the entire inflorescence, sometimes
also comprising the upper part of the stem. The sori are
initially enclosed by a thick, brownish peridium that later
ruptures irregularly exposing a single, stout columella surrounded by a mass of spores and sterile cells.
Until ten years ago, with 326 species Sporisorium was
perceived by far the most species-rich genus of smut fungi.
A new concept for Ustilago, Sporisorium, and Macalpinomyces was proposed by McTaggart et al. (2012a, b). In its
modern circumscription, Sporisorium comprises 198 species
(Vánky 2011, 2013; Denchev et al. 2012, 2016; McTaggart
Ustilaginales G. Winter.
There are seven families in this order: Anthracoideaceae
Denchev, Clintamraceae Vánky, Geminaginaceae Vánky,
Melanotaeniaceae Begerow et al., Pericladiaceae Vánky,
Ustilaginaceae Tul & C. Tul., and Websdaneaceae Vánky
(He et al. 2019; but excluding Cintractiellaceae Vánky, see
McTaggart et al. 2020).
Ustilaginaceae Tul & C. Tul. [as 'Ustilagineae'], Annls
Sci. Nat., Bot., sér. 3 7: 14 (1847).
Ustilaginaceae was introduced by Tulasne and Tulasne
(1847). Twenty-eight genera are currently recognized in this
family (He et al. 2019).
13
Fungal Diversity (2022) 117:1–272
et al. 2012b; Denchev and Denchev 2013, 2016; Wang et al.
2015).
During an examination of specimens of grasses in the
herbarium MA of the Real Jardín Botánico (Madrid, Spain),
a smut fungus belonging to Sporisorium was found on a
specimen of Anadelphia leptocoma (MA 690545) from Burkina Faso. Based on distinct morphology and phylogenetic
evidence (Fig. 186), this fungus is introduced here as a novel
species.
Sporisorium anadelphiae-leptocomae T. Denchev,
Denchev, Kemler, M.P. Martín & Begerow, sp. nov.
Index Fungorum number: IF558175; Facesoffungi number: FoF09652; Fig. 185.
Etymology: The specific epithet refers to the host species.
Holotype: SOMF 30250
Parasitic on Anadelphia leptocoma. Infection systemic.
Sori destroying the racemes, 5–10 mm long, fusiform, all
racemes of an infected plant affected; initially covered by a
thick, yellow–brown peridium which later ruptures irregularly, exposing a single columella, surrounded by a powdery, dark reddish-brown mass of spores and sterile cells.
Columella shorter than the sorus, flagelliform, apically
slightly branched, sometimes slightly flattened or with shallow longitudinal furrows. Sori at first completely enclosed
by the spatheoles, later more or less visible. Sterile cells
firmly packed in irregular groups, collapsed, 9–14(–17)
μm long, usually larger than the spores, hyaline to pale yellow; wall 0.6–1.0 μm thick. In SEM, smooth, sometimes
partially rugulose. Spores subglobose, slightly irregular,
broadly ellipsoidal, ellipsoidal or ovoid, (6.5–)7.5–9.5(–1
0.5) × (6–)7–8.5(–9.5) (8.6 ± 0.6 × 7.8 ± 0.6) μm (n = 300),
medium reddish brown; wall evenly thickened, 0.5–0.7 μm
thick, smooth. In SEM, minutely echinulate-verruculose,
ornaments up to 0.2 μm high, densely punctate between the
main ornaments.
Material examined: Burkina Faso, Cascades Region,
Léraba Province, Sindou Department, Tourni, dam, riverbed with seeping water, fallow fields, 10°46′ N, 05°09′
W, alt. 200–500 m, on Anadelphia leptocoma (Trin.) Pilg.
(Poaceae), 28 October 1997, S. Lægaard, H. Mipro & T.
Soberé, no. 18421 (SOMF 30250, holotype; MA 690545,
isotype).
GenBank numbers: MW599285(ITS), MW599284
(LSU).
Notes: Anadelphia Hack. is a small genus of Poaceae
Barnhart, tribe Andropogoneae Dumort., subtribe
Andropogoninae J. Presl (including 514 species in 25
genera) (Soreng et al. 2017). Anadelphia comprises 14
species (Clayton et al. 2015) characterized by compound
inflorescence composed of racemes. Anadelphia leptocoma is distributed in Tropical Africa (from Senegal to
Zambia), mainly in West Tropical Africa (Clayton et al.
217
2015; Poilecot et al. 2015; Ibrahim et al. 2018). It is a
characteristic grass of the savannahs (Poilecot et al. 2015).
Seven smut fungi have been previously reported on
hosts in Anadelphia and its closely related genera, Elymandra Stapf and Monocymbium Stapf: Anthracocystis
anadelphiae (Vienn.-Bourg.) McTaggart & R.G. Shivas,
Jamesdicksonia anadelphiae (Vienn.-Bourg.) Piątek, J.
anadelphiae-trichaetae T. Denchev & Denchev, Macalpinomyces elymandrae (Vienn.-Bourg.) Vánky, Sporisorium anadelphiae-trichaetae T. Denchev & Denchev,
S. monocymbii (Syd.) Vánky, and Tilletia elymandrae
Vienn.-Bourg. (Denchev and Denchev 2016). Sporisorium
anadelphiae-leptocomae can be easily distinguished from
S. anadelphiae-trichaetae by having (i) sori that destroy
the racemes entirely, while the sori of S. anadelphiaetrichaetae affect only the spikelets, and (ii) minutely
echinulate-verruculose spores with ornaments up to
0.2 μm in height, while S. anadelphiae-trichaetae possesses moderately echinulate spores, with spinules up to
0.7 μm in height. The sori of Sporisorium monocymbii,
destroying spikelets or groups of spikelets and forming a
very characteristic, strongly branched body (see Figs. 45,
47, 48 in Denchev and Denchev 2016), also differ from
those of S. anadelphiae-leptocomae. Additionally, Sporisorium monocymbii has minutely echinulate spores, with
spinules up to 0.3(–0.4) μm in height and larger sterile
cells, 9–19(–24) μm long vs 9–14(–17) μm long for S.
anadelphiae-leptocomae.
The phylogenetic analysis of Sporisorium, based on combined ITS and LSU sequences resulted in a similar topology
to previous analyses (McTaggart et al. 2012a). Sporisorium
anadelphiae-leptocomae formed a statistically very well
supported clade (100% bootstrap) together with S. queenslandicum Vánky et al., infecting Sehima nervosa (Roem. &
Schult.) Stapf, and S. exsertum (McAlpine) L. Guo, infecting
Themeda spp. Sister to this clade is a group containing S.
rarum R.G. Shivas et al., S. ryleyi Vánky & R.G. Shivas, S.
paspali (Speg.) Vánky, S. porosum (Langdon) McTaggart
& R.G. Shivas, S. trachypogonicola Vánky & C. Vánky, S.
vanderystii (Henn.) Langdon & Full., and S. wynaadense
(Sundaram) Vánky & R.G. Shivas. All of the species from
these two clades are parasitizing grasses from the tribe
Andropogoneae, with the exception of S. paspali, infecting
Paspalum spp., belonging to the tribe Paspaleae (Fig.*).
The smut fungi of Burkina Faso are very poorly known
with only six species reported from this country: Anthracocystis ehrenbergii (J.G. Kühn) McTaggart & R.G. Shivas, A. livingstoneana (Vánky) McTaggart & R.G. Shivas,
Moesziomyces penicillariae (Bref.) Vánky, Sporisorium
reilianum (J.G. Kühn) Langdon & Full., S. scitamineum
(Syd) M. Piepenbr. et al., and S. sorghi Ehrenb. ex Link
(Vánky et al. 2011; Piepenbring et al. 2020).
13
218
Fungal Diversity (2022) 117:1–272
Table 5 Updated list of fungal taxa published in the previous fungal diversity notes
FDN number
Species name
Status
Country/Region
Comment
Thailand
Placed in Caniaceae by Wijayawardene et al. (2020)
N/C
Fungal diversity notes 1–110 (Liu et al. 2015)
1
Amphibambusa
New genus
2
Amphibambusa bambusicola
New species
3
Amphisphaeria sorbi
New species
Italy
4
Atrotorquata spartii
New species
Italy
N/C
5
Oxydothis atypica
New species
Thailand
was introduced without molecular data, was not included in
Konta et al. 2016
6
Pestalotiopsis digitalis
New species
New zealand
N/C
7
Pestalotiopsis dracontomelon
New species
Thailand
N/C
8
Pestalotiopsis italiana
New species
Italy
N/C
9
Conicomyces pseudotransvaalensis
New species
Japan
N/C
10
Dinemasporium nelloi
New species
Italy
N/C
11
Diaporthe thunbergiicola
New species
Thailand
N/C
12
Diatrype palmicola
New species
Thailand
N/C
13
Phaeoisaria pseudoclematidis
New species
Thailand
N/C
14
Colletotrichum sedi
New species
Russia
N/C
15
Natantispora unipolaris
New species
Taiwan
N/C
16
Saagaromyces mangrovei
New species
Saudi arabia
N/C
17
Myrothecium macrosporum
New species
Thailand
Nom. illegit., Art. 53.1
18
Neogaeumannomyces
New genus
Thailand
N/C
19
Neogaeumannomyces bambusicola
New species
20
Meliola tamarindi
Reference specimen
Thailand
N/C
21
Hapalocystis berkeleyi
Reference specimen
Italy
N/C
22
Cytospora berberidis
New species
China
N/C
23
Cytospora sibiraeae
New species
China
N/C
24.*
Annulohypoxylon leptascum
New record
Brazil
N/C
25.*
Annulohypoxylon nitens
New sequence data
Thailand
N/C
26.*
Annulohypoxylon stygium
New sequence data
Sri Lanka
N/C
27.*
Annulohypoxylon thailandicum
New species
Thailand
N/C
28.*
Biscogniauxia marginata
New sequence data
France
N/C
29.*
Fasciatispora nypae
Reference specimen
Thailand
N/C
30.*
Hypoxylon fendleri
New sequence data
Venezuela
N/C
31.*
Hypoxylon lenormandii
New sequence data
Cuba
N/C
32.*
Hypoxylon monticulosum
New sequence data
Thailand
N/C
33.*
Flammeascoma
New genus
Thailand
N/C
34.*
Flammeascoma bambusae
New species
35.*
Palmiascoma
New genus
Thailand
N/C
36.*
Palmiascoma gregariascomum
New species
37.*
Chaetocapnodium
New genus
Thailand
Chaetocapnodium siamense Hongsanan & K.D. Hyde [as
'siamensis']
New species
38.*
Chaetocapnodium siamensis
39.*
Phragmocapnias philippinensis
New species
Philippines
Chaetocapnodium philippinense (Abdollahzadeh et al. 2020)
40.*
Brunneomycosphaerella
New genus
Italy
Placed in Mycosphaerellales genera incertae sedis by (Wijayawardene et al. 2022)
41.*
Brunneomycosphaerella laburni
New species
42.*
Dictyosporium aquaticum
New species
Egypt
N/C
43.*
Dictyosporium meiosporum
New species
Thailand
N/ C
44.*
Dictyosporium thailandicum
New species
Thailand
N/C
45.*
Didymella cirsii
New species
Italy
N/C
46.*
Microsphaeropsis olivacea
first report of the sexual
morph
Germany
N/C
47.*
Phoma medicaginis
Additional collection
Italy
Ascochyta medicaginicola (Chen et al. 2015)
48.*
Kalmusia italica
New species
Italy
N/C
49.*
Kalmusia spartii
New species
Italy
N/C
50.*
Montagnula graminicola
New species
Italy
N/C
51.*
Paraconiothyrium nelloi
New species
Italy
N/C
13
Fungal Diversity (2022) 117:1–272
219
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
52
Paraconiothyrium thysanolaenae
New species
Thailand
N/C
53
Paraphaeosphaeria spartii
New species
Italy
N/C
54
Pseudocamarosporium cotinae
New species
Russia
Pseudocamarosporium cotini Norph., Bulgakov & K.D. Hyde
[as 'cotinae']
55
Psiloglonium colihuae
Reference specimen
Thailand
N/C
56
Psiloglonium multiseptatum
New species
Thailand
N/C
57
Psiloglonium sasicola
Reference specimen
Thailand
N/C
58
Aliquandostipite manochii
New species
Thailand
Aliquandostipite manochiae Sri-indr., Boonyuen, Suetrong,
K.L. Pang & E.B.G. Jones [as 'manochii']
59
Keissleriella sparticola
New species
Italy
Keissleriella spartiicola Singtr. & K.D. Hyde [as 'sparticola']
60
Leptosphaeria doliolum
Additional collection
Italy
N/C
61
Leptosphaeria ebuli
New species
Italy
N/C
62
Paraleptosphaeria nitschkei
Reference specimen
Italy
N/C
63
Plenodomus agnitus
Reference specimen
Germany
N/C
64
Lophiostoma pseudodictyosporium
New species
Italy
N/C
65
Lophiostoma ravennicum
New species
Italy
N/C
66
Lophiotrema eburnoides
New species
Japan
Lophiotrema eburneoides Kaz. Tanaka, A. Hashim. & K. Hiray.
[as 'eburnoides']
67
Byssosphaeria musae
New species
Thailand
N/C
68
Pseudotrichia rubriostiolata
New species
Thailand
N/C
69
Pseudotrichia thailandica
New species
Thailand
N/C
70
Sarimanas
New genus
Japan
N/C
71
Sarimanas pseudofluviatile
New species
72
Sarimanas shirakamiense
New species
73
Neomicrothyrium
Valid publication of the
genus
Japan
N/C
Zeloasperisporium (Hongsanan et al. 2015)
74
Pallidocercospora acaciigena
New record
Thailand
75
Pseudocercospora tamarindi
New species
Thailand
N/C
N/C
76
Zasmidium musae
First report of the sexual
morph
Thailand
N/C
77
Paradictyoarthriniaceae
New family
Thailand
N/C
78
Paradictyoarthrinium tectonicola
New species
79
Allophaeosphaeria
New genus
Italy
Septoriella neodactylidis (Marin-Felix et al. 2019)
80
Allophaeosphaeria dactylidis
New species
81
Allophaeosphaeria muriformia
New species
Italy
Allophaeosphaeria muriformis (Marin-Felix et al. 2019)
82
Neosetophoma clematidis
New species
Italy
N/C
83
Neosetophoma italica
New species
Italy
N/C
84
Phaeosphaeria musae
New species
Thailand
Nom. inval., Art. 39.1 (Melbourne)
85
Wojnowicia dactylidicola
New species
Italy
Galiicola dactylidicola (Thambugala et al. 2017)
86
Wojnowicia dactylidis
New species
Italy
Wojnowiciella dactylidis (Hernandez-Restrepo et al. 2016)
87
Wojnowicia lonicerae
New species
Italy
Wojnowiciella lonicerae (Hernandez-Restrepo et al. 2016)
88
Splanchnonema pupula
Reference specimen
Italy
N/C
89
Alternaria cesenica
New species
Italy
N/C
90
Camarosporium aborescentis
New species
Russia
Camarosporium arborescentis (Wanasinghe et al. 2017)
91
Camarosporium aureum
New species
Russia
N/C
92
Camarosporium caraganicola
New species
Russia
Camarosporidiella caraganicola (Wanasinghe et al. 2017)
93
Multiseptospora
New genus
Thailand
Placed in Parabambusicolaceae by Wijayawardene et al. (2022)
Italy
N/C
94
Multiseptospora thailandica
New species
95
Saccotheciaceae
New family
96
Tothia spartii
New species
N/C
97
Chaetothyrium agathis
New species
Philippines
Chaetothyrium agathidis Hongsanan & K.D. Hyde [as 'agathis']
98
Trichomerium siamensis
New species
Thailand
Trichomerium siamense Hongsanan & K.D. Hyde [as 'siamensis']
99
Paralecia
New genus
Italy
Placed in Cladoniaceae by Wijayawardene et al. (2022)
100
Paralecia pratorum
New species
101
Lauriomyces synnematicus
New species
Thailand
N/C
13
220
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
102
Agaricus flavicentrus
New species
Thailand
N/C
103
Agaricus hanthanaensis
New species
Sri Lanka
N/C
104
Agaricus parvibicolor
New species
Thailand
N/C
105
Agaricus sodalis
New species
Thailand
N/C
106
Cantharellus luteostipitatus
New species
Madagascar
N/C
107
Lactarius atrobrunneus
New species
Thailand
N/C
108
Lactarius politus
New species
Thailand
N/C
109
Phylloporia dependens
New species
China
N/C
110
Russula cortinarioides
New species
US
N/C
Italy
Lophiostomataceae (Magaña-Dueñas et al. 2020)
Italy
Massariosphaeria grandispora (Magaña-Dueñas et al. 2020)
Fungal diversity notes 111–252 (Ariyawansa et al. 2015)
111
Pseudomassariosphaeria
New genus
112
Pseudomassariosphaeria bromicola
New species
113
Pseudomassariosphaeria grandispora
New combination
114
Flammeascoma lignicola
New species
Thailand
N/C
115
Ascocylindricaceae
New family
Saudi arabia
N/C
116
Ascocylindrica
New genus
117
Ascocylindrica marina
New species
118
Lembosia xyliae
New species
Thailand
N/C
119
Diplodia crataegicola
New species
Italy
N/C
120
Diplodia galiicola
New species
Italy
N/C
121
Caryosporaceae
New family
Thailand
N/C
122
Caryospora aquatica
New species
123
Cucurbitaria ephedricola
Reference specimen
Italy
N/C
124
Heracleicola
New genus
Italy
125
Heracleicola premilcurensis
New species
Ascochyta (Chen et al. 2017a, b)
Ascochyta premilcurensis (Chen et al. 2017a, b)
126
Neodidymella
New genus
127
Neodidymella thailandicum
New species
128
Austropleospora osteospermi
Type species description
129
Austropleospora archidendri
New combination
130
Pseudopithomyces
New genus
Reference specimens or changes in classification
131
Pseudopithomyces chartarum
New combination
N/C
132
Pseudopithomyces palmicola
New species
Thailand
N/C
133
Pseudopithomyces maydicus
New combination
Thailand
N/C
134
Pseudopithomyces sacchari
New combination
135
Floricola viticola
New species
Italy
Teichospora viticola (Jaklitsch et al. 2016)
136
Brunneoclavispora
New genus
Thailand
N/C
137
Brunneoclavispora bambusae
New species
138
Neolophiostoma
New genus
Thailand
N/C
139
Neolophiostoma pigmentatum
New species
140
Sulcosporium
New genus
Thailand
N/C
141
Sulcosporium thailandica
New species
142
Pseudoasteromassaria
New genus
143
Pseudoasteromassaria fagi
New species
144
Keissleriella dactylidicola
New species
145
Neomassariosphaeria
Changes in classification
146
Lophiostoma caulium
147
148
Thailand
N/C
Neodidymella thailandica Phook., R.H. Perera & K.D. Hyde [as
'thailandicum']
Australia
N/C
N/C
N/C
Sulcosporium thailandicum Phook. & K.D. Hyde [as 'thailandica']
Japan
N/C
N/C
Italy
N/C
Reference specimen
Italy
Sigarispora caulium (Thambugala et al. 2015)
Lophiohelichrysum
New genus
Italy
N/C
Lophiohelichrysum helichrysi
New species
149
Aquasubmersa japonica
New species
Japan
N/C
150
Pseudomonodictys
New genus
Thailand
Parabambusicolaceae (Wijayawardene et al. 2022)
151
Pseudomonodictys tectonae
New species
152
Byssosphaeria rhodomphala
Reference specimen
Thailand
N/C
13
Amniculicolaceae (Jones et al. 2015)
Fungal Diversity (2022) 117:1–272
221
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
153
Microthyrium buxicola
New species
Italy
N/C
154
Tumidispora
New genus
Thailand
N/C
155
Tumidispora shoreae
New species
156
Alloleptosphaeria clematidis
New species
Italy
N/C
157
Allophaeosphaeria cytisi
New species
Italy
Arezzomyces cytisi (Marin-Felix et al. 2019)
158
Allophaeosphaeria subcylindrospora
New species
Italy
Septoriella subcylindrospora (Marin-Felix et al. 2019)
159
Dematiopleospora luzulae
New species
Italy
N/C
160
Entodesmium artemisiae
New species
Russia
Ophiobolus artemisiae (Phookamsak et al. 2017)
161
Galiicola
New genus
Italy
N/C
162
Galiicola pseudophaeosphaeria
New species
163
Loratospora luzulae
New species
Italy
N/C
164
Nodulosphaeria senecionis
New species
Italy
N/C
165
Ophiosphaerella aquaticus
New species
Thailand
Ophiosphaerella aquatica Z.L. Luo, Hong Y. Su & K.D. Hyde
[as 'aquaticus']
166
Populocrescentia
New genus
Italy
N/C
167
Populocrescentia forlicesenensis
New species
168
Vagicola
New genus
169
Vagicola vagans
New combination
170
Alternaria ethzedia
Illustration of the sexual
morph
Italy
N/C
171
Elongatopedicellata
New genus
Thailand
N/C
172
Elongatopedicellata lignicola
New species
N/C
Septoriella vagans (Marin-Felix et al. 2019)
173
Roussoella magnatum
New species
Thailand
Roussoella magnata D.Q. Dai & K.D. Hyde [as 'magnatum']
174
Roussoella angustior
New species
Thailand
Roussoella angusta D.Q. Dai & K.D. Hyde [as 'angustior']
175
Shrungabeeja longiappendiculata
New species
Thailand
N/C
176
Massariosphaeria
Family placement
Cyclothyriellaceae (Wijayawardene et al. 2022)
177
Wicklowiaceae
New family
N/C
178
Mycocalicium hyaloparvicellulum
New species
Italy
179
Acarospora septentrionalis
New species
Iceland
N/C
180
Acarospora castaneocarpa
New species
Finland
N/C
181
Chapsa multicarpa
New species
Thailand
N/C
182
Fissurina carassensis
New species
Brazil
N/C
183
Sticta fuscotomentosa
New species
Colombia
N/C
184
Sticta subfilicinella
New species
Colombia
N/C
185
Helvella pseudolacunosa
New species
China
N/C
186
Helvella rugosa
New species
China
N/C
187
Dictyosporella
New genus
Egypt
N/C
188
Dictyosporella aquatica
New species
189
Chaetosphaeria rivularia
New species
France
Chaetosphaeria rivularis Réblová & J. Fourn. [as 'rivularia']
190
Beauveria gryllotalpidicola
New species
Thailand
N/C
191
Beauveria loeiensis
New species
Thailand
N/C
192
Seimatosporium sorbi
New species
Italy
Sporocadus sorbi (Liu et al. 2018)
193
Seimatosporium pseudorosarum
New species
Italy
N/C
194
Colletotrichum aciculare
New species
Thailand
Colletotrichum truncatum (Jayawardena et al. 2016)
195
Colletotrichum fusiforme
New species
Thailand
N/C
196
Colletotrichum hymenocallidicola
New species
Thailand
Colletotrichum orchidearum
Damm et al. (2019)
Taiwan
N/C
197
Tinhaudeus
New genus
198
Tinhaudeus formosanus
New species
N/C
199
Pestalotiopsis subshorea
New species
China
N/C
200
Pestalotiopsis dracaenea
New species
China
Pestalotiopsis dracaenae Yong Wang bis, Yu Song, K. Geng &
K.D. Hyde [as 'dracaenea']
201
Pestalotiopsis montellica
Changes in classification China
N/C
202
Phaeoacremonium tectonae
New species
Thailand
N/C
203
Cytospora parasitica
New species
Russia
N/C
N/C
13
222
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
204
Cytospora tanaitica
New species
Russia
N/C
205
Annulohypoxylon palmicola
New species
Thailand
N/C
206
Biscogniauxia effusa
New species
China
N/C
207
Nemania fusoidispora
New species
China
N/C
208
Agaricus pseudolangei
New species
Thailand
N/C
209
Agaricus haematinus
New species
Thailand
N/C
210
Agaricus atrodiscus
New species
Thailand
N/C
211
Agaricus exilissimus
New species
Thailand
N/C
212
Amanita melleialba
New species
China
N/C
213
Amanita pseudosychnopyramis
New species
China
N/C
214
Amanita subparvipantherina
New species
China
N/C
215
Entoloma calabrum
New species
Italy
N/C
216
Cora barbulata
New species
Costa rica
N/C
217
Dictyonema gomezianum
New species
Costa rica
N/C
218
Inocybe granulosa
New species
Sweden
Mallocybe granulosa (Matheny and Kudzma 2019)
219
Xerocomellus sarnarii
New species
Italy
N/C
220
Cantharellus eucalyptorum
New species
Madagascar
N/C
221
Cantharellus nigrescens
New species
Madagascar
N/C
222
Cantharellus tricolor
New species
Madagascar
N/C
223
Cantharellus variabilicolor
New species
Madagascar
N/C
224
Cortinarius alboamarescens
New species
Denmark
N/C
225
Cortinarius brunneoalbus
New species
USA
N/C
226
Cortinarius ochroamarus
New species
Finland
N/C
227
Cortinarius putorius
New species
USA
N/C
228
Cortinarius seidlii
New species
USA
Cortinarius seidliae Ammirati, Niskanen & Liimat. [as 'seidlii']
229
Dendrominiaceae
New family
N/C
230
Punctulariopsis cremeoalbida
New combination
N/C
231
Punctulariopsis efibulata
New combination
US
N/C
232
Hymenochaete micropora
New species
China
N/C
233
Hymenochaete subporioides
New species
China
234
Neoantrodiellaceae
New family
N/C
235
Neoantrodiella
New genus
N/C
236
Neoantrodiella gypsea
New combination
N/C
237
Neoantrodiella thujae
New combination
238
Xylodon ramicida
New species
Finland
N/C
239
Colospora
New genus
Indonesia
N/C
240
Colospora andalasii
New species
241
Russula guangxiensis
New species
China
N/C
242
Russula hakkae
New species
China
N/C
243
Tremella dirinariae
New species
US
N/C
244
Tremella graphidis
New species
US
N/C
245
Tremella pyrenulae
New species
US
N/C
246
Absidia caatinguensis
New species
Brazil
N/C
247
Absidia koreana
New species
Korea
N/C
248
Gongronella koreana
New species
Korea
N/C
249
Mortierella pisiformis
New species
Taiwan
N/C
250
Mortierella formosana
New species
Taiwan
N/C
251
Neocallimastix cameroonii
New species
UK
N/C
252
Piromyces irregularis
New species
Czech Republic
N/C
N/C
N/C
N/C
Fungal diversity notes 253–366 (Li et al. 2016)
253
Dothiorella rhamni
New species
Russia
254
Dothiorella vidmadera
Reference specimen
Italy
Dothiorella sarmentorum (Zhang et al. 2021)
255
Eutiarosporella dactylidis
New combination
Italy
N/C
256
Mucoharknessia anthoxanthi
New species
Italy
N/C
257
Neophaeocryptopus
New genus
13
Dothiora (Crous et al. 2018)
Fungal Diversity (2022) 117:1–272
223
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
258
Neophaeocryptopus cytisi
New species
Italy
Dothiora cytisi (Crous et al. 2018)
259
Saccothecium rubi
New species
Italy
N/C
260
Psiloglonium macrosporum
New species
Thailand
N/C
261
Pseudocamarosporium pini
New combination
Italy
N/C
262
Towyspora
New genus
UK
N/C
263
Towyspora aestuari
New species
264
Lindgomyces okinawaensis
New species
Japan
N/C
265
Lophiostoma pseudoarmatisporum
New species
Japan
N/C
266
Sigarispora ononidis
New species
Italy
N/C
267
Aposphaeria corallinolutea
New record
Thailand
N/C
268
Multilocularia
New genus
Thailand
N/C
269
Multilocularia bambusae
New species
270
Multiseptospora thysanolaenae
New species
Thailand
N/C
271
Parastagonospora cumpignensis
New species
Italy
Parastagonospora campignensis Tibpromma, Camporesi &
K.D. Hyde [as 'cumpignensis']
272
Comoclathris pimpinellae
New species
Russia
N/C
273
Angustospora
New genus
Egypt
N/C
274
Angustospora nilensis
New species
275
Polyplosphaeria thailandica
New species
Thailand
N/C
276
Longiostiolum
New genus
Thailand
Longiostiolaceae (Wijayawardene et al. 2022)
277
Longiostiolum tectonae
New species
278
Pseudodidymosphaeria phlei
New species
Italy
N/C
279
Clematidis
New genus
Italy
Pseudolophiotremataceae (Wijayawardene et al. 2022)
280
Clematidis italica
New species
281
Crassiparies
New genus
Japan
Neohendersoniaceae (Wijayawardene et al. 2022)
Saudi arabia
N/C
Thailand
N/C
N/C
282
Crassiparies quadrisporus
New species
283
Farasanispora
New genus
284
Farasanispora avicenniae
New species
285
Parameliola
New genus
286
Parameliola dimocarpi
New species
287
Parameliola acaciae
New species
Thailand
N/C
288
Kirschsteiniothelia tectonae
New species
Thailand
N/C
289
Ocellularia arachchigei
New species
Sri Lanka
N/C
290
Ocellularia ratnapurensis
New species
Sri Lanka
N/C
291
Rhabdodiscus albodenticulatus
New species
Sri Lanka
N/C
292
Pseudolachnella brevifusiformis
New species
Japan
N/C
293
Phragmoporthe conformis
reference species
Italy
N/C
294
Cytospora salicicola
New species
Russia
N/C
295
Colletotrichum menispermi
New species
Russia
N/C
296
Colletotrichum quinquefoliae
New species
Russia
N/C
297
Ochronectria thailandica
New species
Thailand
N/C
298
Moelleriella phukhiaoensis
New species
Thailand
N/C
299
Moelleriella pongdueatensis
New species
Thailand
N/C
300
Ophiocordyceps formosana
New record
China
Nom. inval., Arts 41.5, F.5.1 (Shenzhen)
301
Ophiocordyceps karstii
New species
China
N/C
302
Aniptodera aquibella
New species
Thailand
N/C
303
Humicola koreana
New species
Korea
N/C
304
Seimatosporium pseudocornii
New species
Italy
Seimatosporium pseudocorni Wijayaw., Camporesi & K.D.
Hyde [as 'pseudocornii']
305
Seimatosporium pseudorosae
New species
Italy
N/C
306
Cryptovalsa ampelina
reference specimen
Italy
N/C
307
Diatrype thailandica
New species
Thailand
Allodiatrype thailandica (Konta et al. 2020)
308
Annulohypoxylon albidiscum
New species
Thailand
N/C
309
Astrocystis thailandica
New species
Thailand
310
Camporesia
New genus
N/C
N/C
13
224
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
311
Camporesia sambuci
New species
Italy
N/C
312
Durotheca macrostroma
New species
Thailand
N/C
313
Halorosellinia rhizophorae
New species
Thailand
N/C
314
Hypoxylon lilloi
New species
Argentina
N/C
315
Rosellinia chiangmaiensis
New species
Thailand
N/C
316
Petrakia echinata
Reference specimen
Italy
N/C
317
Agaricus coccyginus
New species
China
N/C
318
Agaricus luteofibrillosus
New species
China
N/C
319
Clarkeinda trachodes
Reference specimen
Sri Lanka
N/C
320
Amanita atrobrunnea
New species
Thailand
N/C
321
Amanita digitosa
New species
Thailand
N/C
322
Amanita gleocystidiosa
New species
Thailand
N/C
323
Amanita pyriformis
New species
Thailand
N/C
324
Amanita strobilipes
New species
Thailand
N/C
325
Cortinarius albosericeus
New species
Canada
N/C
326
Cortinarius badioflavidus
New species
US
N/C
327
Cortinarius denigratus
New species
Canada
N/C
328
Cortinarius duboisensis
New species
US
N/C
329
Cortinarius fragrantissimus
New species
Canada
N/C
330
Cortinarius roseobasilis
New species
US
N/C
331
Cortinarius vinaceobrunneus
New species
Canada
N/C
332
Cortinarius vinaceogrisescens
New species
US
N/C
333
Cortinarius wahkiacus
New species
US
N/C
334
Musumecia alpina
New species
China
N/C
335
Musumecia sardoa
New species
Italy
N/C
336
Cyanoboletus hymenoglutinosus
New species
India
Nom. inval., Art. 40.7 (Shenzhen)
337
Leccinellum indoaurantiacum
New species
India
Nom. inval., Art. 40.7 (Shenzhen)
338
Galzinia longibasidia
New sequence data
Iran
N/C
339
Leptocorticium tenellum
New sequence data
Chile
N/C
340
Fomitiporia atlantica
New species
Brazil
N/C
341
Fomitiporia subtilissima
New species
Brazil
N/C
342
Inonotus shoreicola
New species
Thailand
N/C
343
Ganoderma wuzhishanensis
New species
China
Ganoderma wuzhishanense T.C. Wen, Hapuar. & K.D. Hyde
[as 'wuzhishanensis']
344
Dentocorticium ussuricum
New sequence data
Russia
N/C
345
Lentinus stuppeus
Reference specimen
Thailand
N/C
346
Bondarzewia tibetica
New species
China
N/C
347
Lactifluus armeniacus
New species
Thailand
N/C
348
Lactifluus ramipilosus
New species
Thailand
N/C
349
Russula amethystina
New species
China
N/C
350
Russula wangii
New species
China
N/C
351
Anaeromyces robustus
New species
N/C
352
Neocallimastix californiae
New species
N/C
353
Piromyces finnis
New species
354
Phytophthora estuarina
New species
Brazil
N/C
355
Phytophthora rhizophorae
New species
Brazil
N/C
356
Salispina
New genus
Brazil
Salispinaceae (Wijayawardene et al. 2022)
357
Salispina intermedia
New species
358
Salispina lobata
New combination
359
Salispina spinosa
New combination
360
Mortierella calciphila
New species
Poland
N/C
361
Absidia stercoraria
New species
Korea
N/C
362
Gongronella orasabula
New species
Korea
N/C
363
Mucor caatinguensis
New species
Brazil
N/C
364
Mucor koreanus
New species
Korea
N/C
13
N/C
Fungal Diversity (2022) 117:1–272
225
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
365
Mucor merdicola
New species
Brazil
N/C
366
Rhizopus koreanus
New species
Korea
N/C
Fungal diversity notes 367–490 (Hyde et al. 2016)
367
Asterina cynometrae
New species
Philippines
N/C
368
Dothiorella iranica
New host
Italy
N/C
369
Dothiorella sarmentorum
New host
Russia
N/C
370
Dothiorella vidmadera
New host
Russia
Dothiorella sarmentorum (Zhang et al. 2021)
371
Pallidocercospora thailandica
New combination
Thailand
N/C
372
Dothiora buxi
New species
Italy
N/C
373
Gloniopsis calami
New species
Thailand
N/C
374
Pseudocoleophoma typhicola
New species
UK
N/C
375
Pseudodictyosporium thailandica
New species
Thailand
N/C
376
Neomicrosphaeropsis cytisi
New species
Italy
N/C
377
Neomicrosphaeropsis cytisinus
New species
Italy
Neomicrosphaeropsis cytisina Tennakoon, Camporesi & K.D.
Hyde [as 'cytisinus']
378
Neomicrosphaeropsis minima
New species
Italy
N/C
379
Neodidymelliopsis ranunculi
New species
Italy
N/C
380
Platychora ulmi
Italy
N/C
381
Stagonosporopsis centaureae
New species
Italy
N/C
382
Montagnula cirsii
New species
Italy
N/C
383
Tremateia arundicola
New species
Italy
Tremateia arunicola Wanas., E.B.G. Jones & K.D. Hyde [as
'arundicola']
384
Tremateia guiyangensis
New species
UK
N/C
385
Lentithecium unicellulare
New species
China
N/C
386
Lentithecium voraginesporum
New species
Egypt
N/C
387
Leptosphaeria cirsii
New species
Saudi arabia
N/C
388
Leptosphaeria irregularis
New species
Italy
N/C
389
Lindgomycetaceae
New family
390
Arundellina typhae
New species
UK
N/C
391
Lindgomyces pseudomadisonensis
New species
UK
N/C
392
Vaginatispora fuckelii
New record
Japan
Neovaginatispora fuckelii (Hashimoto et al. 2018)
393
Hermatomyces mirum
New combination
China
Hermatomyces mirus (Starbäck) C.G. Lin, Bhat & K.D. Hyde
[as 'mirum']
N/C
394
Hermatomyces subiculosa
New species
Thailand
N/C
395
Lophiotrema bambusae
New species
Thailand
Atrocalyx bambusae (De Silva et al. 2018)
396
Lophiotrema fallopiae
New species
Japan
N/C
397
Neomassaria
New genus
Italy
N/C
398
Neomassaria fabacearum
New species
399
Stagonospora forlicesenensis
New species
Italy
N/C
400
Bertiella ellipsoidea
New species
Thailand
N/C
401
Occultibambusa aquatica
New species
Thailand
N/C
402
Occultibambusa chiangraiensis
New species
Thailand
N/C
403
Camarosporioides
New genus
Germany
N/C
404
Camarosporioides phragmitis
New species
405
Chaetosphaeronema achilleae
New species
Russia
N/C
406
Dematiopleospora alliariae
New species
Italy
Hawksworthiana alliariae (Wanasinghe et al. 2018)
407
Dematiopleospora cirsii
New species
Italy
N/C
408
Juncaceicola italica
New species
Italy
N/C
409
Leptospora rubella
Reference specimen
UK
N/C
410
Leptospora galii
New species
Italy
N/C
411
Leptospora thailandica
New species
Thailand
N/C
412
Muriphaeosphaeria ambrosiae
New species
Russia
N/C
413
Nodulosphaeria italica
New species
Italy
N/C
414
Poaceicola arundinis
New species
Italy
Septoriella arundinis (Marin-Felix et al. 2019)
415
Pseudophaeosphaeria
New genus
Italy
N/C
13
226
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
416
Pseudophaeosphaeria rubi
New species
417
Wojnowicia italica
418
Comoclathris italica
New species
Italy
N/C
New species
Italy
419
N/C
Neoroussoella lenispora
New species
China
N/C
420
Neotorula submersa
New species
China
N/C
421
Aquaphila albicans
Reference specimen
Thailand
N/C
422
Chlamydotubeufia helicospora
New species
Thailand
Dematiohelicomyces helicosporus (Lu et al. 2018)
423
Helicoma guttulatum
New species
Thailand
N/C
424
Neoacanthostigma septoconstrictum
Reference specimen
Thailand
N/C
425
Tubeufia hyalospora
New species
Thailand
N/C
426
Tubeufia roseohelicospora
New species
Thailand
N/C
427
Pleurotrema thailandica
New species
Thailand
N/C
428
Alloarthopyrenia
New genus
Italy
N/C
429
Alloarthopyrenia italica
New species
Thailand
Neomassarinaceae (Wijayawardene et al. 2022)
430
Neomassarina
New genus
431
Neomassarina thailandica
New species
432
Aspergillus koreanus
New species
Korea
N/C
433
Ceramothyrium menglunense
New species
China
N/C
434
Minimelanolocus submersus
New species
China
N/C
435
Trichomerium bambusae
New species
Thailand
N/C
436
Terriera thailandica
New species
Thailand
N/C
437
Helvella tinta
New species
China
N/C
438
Helvella floriforma
New species
China
N/C
439
Helvella oblongispora
New distribution record
China
N/C
440
Peziza fruticosa
New species
Italy
N/C
441
Coronophora myricoides
New species
China
N/C
442
Diaporthe aseana
New species
Thailand
N/C
443
Diaporthe eres
New host
Italy
N/C
444
Diaporthe foeniculina
New record
Italy
N/C
445
Diaporthe garethjonesii
New species
Thailand
N/C
446
Diaporthe siamensis
New host
Thailand
N/C
447
Cytospora cotini
New species
Russia
N/C
448
Colletotrichum insertae
New species
Russia
N/C
449
Blastophorum aquaticum
New species
China
Cylindrotrichum aquaticum (Luo et al. 2019)
450
Ophiocordyceps hemisphaerica
New species
Brazil
N/C
451
Ophiocordyceps lacrimoidis
New species
Brazil
N/C
452
Purpureocillium sodanum
New species
Iran
N/C
453
Alfaria spartii
New species
Italy
N/C
454
Emericellopsis persica
New species
Iran
N/C
455
Meliola citri-maximae
New species
Thailand
N/C
456
Meliola pseudosasae
Reference specimens
China
N/C
457
Arthrinium paraphaeospermum
New species
Thailand
N/C
458
Bartaliniaceae
New family
Published with the identifier 'Index Fungorum number:
IF511183', but this identifier was not issued for the name
published
Index Fungorum (2022a, b)
459
Neotruncatella endophytica
New species
Korea
Hymenopleella endophytica (Liu et al. 2019a, b, c, d)
460
Cainia globosa
New species
Italy
N/C
461
Ciliochorella phanericola
New species
Thailand
N/C
462
Neopestalotiopsis cocoës
New species
Thailand
N/C
463
Neopestalotiopsis musae
New species
Thailand
464
Pestalotiopsis sequoiae
New species
Italy
N/C
465
Anthostomella ravennica
New species
Italy
N/C
466
Anthostomella thailandica
New species
Thailand
N/C
467
Sporidesmium pyriformatum
New species
Thailand
N/C
13
Fungal Diversity (2022) 117:1–272
227
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
468
Sporidesmium aquaticivaginatum
New species
Thailand
N/C
469
Sporidesmium olivaceoconidium
New species
Thailand
N/C
470
Distoseptispora multiseptata
New species
Thailand
N/C
471
Distoseptispora tectonae
New species
Thailand
N/C
472
Distoseptispora tectonigena
New species
Thailand
N/C
473
Paracapsulospora
New genus
Thailand
N/C
474
Paracapsulospora metroxyli
New species
475
Clavulinopsis aurantiaca
New species
Brazil
N/C
476
Ramariopsis atlantica
New species
Brazil
N/C
477
Cortinarius fulvescens
neo-type
Canada
N/C
478
Cortinarius fulvescentoideus
New species
Finland
N/C
479
Cortinarius nymphatus
New species
Canada
N/C
480
Cortinarius pseudobulliardioides
New species
Finland
N/C
481
Cortinarius tenuifulvescens
New species
Canada
N/C
482
Rhodocybe indica
New species
India
N/C
483
Rhodocybe luteobrunnea
New species
India
N/C
484
Rhodocybe griseoaurantia
New species
India
N/C
485
Cyathus pyristriatus
New species
Thailand
N/C
486
Polyporus mangshanensis
New species
India
N/C
487
Russula indoalba
New species
India
N/C
488
Russula pseudoamoenicolor
New species
India
N/C
489
Lactarius dirkii
New species
Korea
N/C
490
Cunninghamella gigacellularis
New species
Brazil
N/C
N/C
Fungal diversity notes 491–602 (Tibpromma et al. 2017)
491
Morenoina calamicola
New species
Thailand
492
Barriopsis thailandica
New species
Thailand
N/C
493
Polythrincium trifolii
reference specimens
China
N/C
494
Dothiora buxi
asexual morph report
Russia
N/C
495
Hysterium centramurum
New species
Thailand
Ericboehmia centramura
Gardiennet et al. (2019)
496
Hysterobrevium mori
New host
Italy
N/C
497
Angustimassarina alni
New species
Germany
N/C
498
Angustimassarina arezzoensis
New species
Italy
N/C
499
Angustimassarina premilcurensis
New species
Italy
N/C
500
Angustimassarina italica
New species
Italy
N/C
501
Angustimassarina lonicerae
New species
Italy
N/C
502
Biatriosporaceae
New family
503
Berkleasmium ariense
New species
India
N/C
504
Platystomum rosae
New host
Italy
N/C
505
Sigarispora muriformis
New species
Italy
N/C
506
Vaginatispora appendiculata
New host
Thailand
N/C
507
Lophiotrema guttulata
New species
Thailand
Atrocalyx guttulata (De Silva et al. 2018)
508
Lophiotrema vagabundum
New host
Italy
N/C
509
Hermatomyces chiangmaiensis
New species
Thailand
N/C
510
Hermatomyces chromolaenae
New species
Thailand
N/C
511
Melanommataceae
New combination/family
512
Aposphaeria corallinolutea
Asexual morph report
Russia
N/C
513
Nigrograna cangshanensis
New species
China
N/C
514
Roussoellaceae
New family
515
Parathyridaria robiniae
New species
Italy
N/C
516
Paraphaeosphaeria viciae
New species
Italy
N/C
517
Pseudoasteromassaria spadicea
New species
Thailand
N/C
518
Keissleriella cirsii
New species
Russia
N/C
519
Pleurophoma italica
New species
Italy
N/C
520
Pseudodidymosphaeria phlei
New host
Italy
N/C
N/C
N/C
N/C
13
228
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
521
Inflatispora caryotae
New species
Thailand
N/C
522
Ascochyta italica
New species
Italy
N/C
523
Ascochyta medicaginicola
New host
Italy
N/C
524
Ascochyta rosae
New species
Italy
N/C
525
Calophoma petasitis
New species
Italy
N/C
526
Didysimulans
New genus
Italy
N/C
527
Didysimulans italica
New species
528
Didysimulans mezzanensis
New species
Italy
N/C
529
Nothophoma quercina
New host
Italy
N/C
530
Stagonosporopsis ailanthicola
New species
Italy
N/C
531
Paraleptosphaeria padi
New species
Russia
N/C
532
Subplenodomus galicola
New species
Italy
N/C
533
Entodesmium italica
New species
Italy
Ophiobolus italicus (Phookamsak et al. 2017)
534
Neosetophoma garethjonesii
New species
UK
N/C
535
Nodulosphaeria guttulatum
New species
Italy
N/C
536
Nodulosphaeria multiseptata
New species
Italy
N/C
537
Nodulosphaeria sambuci
New species
Italy
N/C
538
Phaeosphaeria calamicola
New species
Thailand
N/C
539
Phaeosphaeriopsis yuccae
New species
Russia
N/C
540
Camarosporium laburnicola
New species
Russia
Camarosporidiella laburnicola (Wanasinghe et al. 2017)
541
Camarosporium moricola
New species
Russia
Camarosporidiella moricola (Wanasinghe et al. 2017)
542
Penicillium punicae
New species
Korea
N/C
543
Exophiala italica
New species
Italy
N/C
544
Gongronella brasiliensis
New species
Brazil
N/C
545
Mucor stercorarius
New species
Korea
N/C
546
Gnomoniopsis sanguisorbae
reference specimen
Italy
N/C
547
Sillia italica
New species
Italy
N/C
548
Cytospora gelida
New species
Russia
N/C
549
Cytospora ceratosperma
New record
Russia
N/C
550
Hypocrella calendulina
Asexual morph reported
Thailand
N/C
551
Moelleriella thanathonensis
New species
Thailand
N/C
552
Myrothecium septentrionale
New species
Thailand
N/C
553
Colletotrichum sambucicola
New species
Italy
N/C
554
Myrmecridium fluviae
New species
Korea
N/C
555
Truncatella spartii
New host
Italy
Heterotruncatella spartii (Liu et al. 2019a, b, c, d)
556
Rosellinia mearnsii
New species
China
N/C
557
Evlachovaea indica
New species
India
N/C
558
Helvella costifera
reference specimen
China
N/C
559
Helvella crispoides
New species
Thailand
N/C
560
Cookeina tricholoma
Reference specimen
Sri Lanka
N/C Wijayawardene et al. (2022)
561
Amanita cornelii
New species
India
N/C
562
Amanita emodotrygon
New species
India
N/C
563
Lepiota cylindrocystidia
New species
Thailand
N/C
564
Lepiota flavocarpa
New species
Thailand
Nom. Inval., Art. F.5.1 (Shenzhen)
565
Lepiota maerimensis
New species
Thailand
N/C
566
Leucocoprinus cretaceus
New record
Sri Lanka
N/C
567
Entoloma magnum
New species
India
N/C
568
Inocybe brunneosquamulosa
New species
India
Pseudosperma brunneosquamulosum (Matheny and Kudzma
2019)
569
Inocybe luteobrunnea
New species
India
Pseudosperma luteobrunneum (Matheny and Kudzma 2019)
570
Inocybe rubrobrunnea
New species
India
N/C
571
Marasmius luculentus
New species
India
N/C
572
Favolaschia auriscalpium
New record
Sri Lanka
N/C
573
Favolaschia manipularis
New record
Sri Lanka
N/C
574
Cyptotrama asprata
New record
Thailand
N/C
13
Fungal Diversity (2022) 117:1–272
229
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
575
Austroboletus appendiculatus
New species
India
N/C
576
Boletellus emodensis
New record
Sri Lanka
N/C
577
Rubroboletus demonensis
New species
Italy
N/C
578
Rubroboletus eastwoodiae
New combination
579
Strobilomyces longistipitatus
New species
India
N/C
580
Clavulina grisea
New species
Brazil
N/C
581
Clavulina ossea
New species
Brazil
N/C
582
Clavulina paraincrustata
New species
Brazil
N/C
583
Fomitopsis flabellata
New species
Brazil
N/C
584
Fomitopsis roseoalba
New species
Brazil
N/C
585
Favolus gracilisporus
New species
Korea
N/C
586
Lentinus sajor-caju
New record
Sri Lanka
N/C
587
Lentinus squarrosulus
New record
Sri Lanka
N/C
588
Lentinus velutinus
New record
Sri Lanka
N/C
589
Panus subfasciatus
New species
Thailand
N/C
590
Polyporus brevibasidiosus
New species
Korea
N/C
591
Polyporus koreanus
New species
Korea
N/C
592
Polyporus orientivarius
New species
Korea
N/C
593
Polyporus parvovarius
New species
Korea
N/C
594
Polyporus subdictyopus
New species
Korea
N/C
595
Polyporus ulleungus
New species
Korea
N/C
596
Wolfiporia pseudococos
New species
China
N/C
597
Coprinopsis cerkezii
New species
Croatia
N/C
598
Russula yanheensis
New species
China
N/C
599
Russula virescens
New record
Sri Lanka
N/C
600
Dacrymyces chiangraiensis
New species
Thailand
N/C
601
Femsjonia monospora
New species
China
N/C
602
Tremella fuciformis
New record
Sri Lanka
N/C
China
N/C
N/C
Fungal diversity notes 603–708 (Hyde et al. 2017)
603
Jahnula guttulaspora
New species
604
Murispora rubicunda
Description of collection Germany
of type species of
genus
N/C
605
Angustimassarina coryli
New species
Italy
N/C
606
Sardiniella celtidis
New species
Italy
N/C
607
Leptoxyphium glochidion
New host
Thailand
N/C
608
Dictyocheirospora cheirospora
New species
China
N/C
609
Epicoccum cedri
New species
Italy
N/C
610
Epicoccum pruni
New species
Russia
N/C
611
Neodidymelliopsis moricola
New species
Russia
N/C
612
Pseudopithomyces kunmingnensis
New species
China
Pseudopithomyces kunmingensis Karun. & K.D. Hyde [as
'kunmingnensis'],
613
Dothiora coronillae
New species
Italy
N/C
614
Dothiora spartii
New species
Italy
N/C
615
Neophaeocryptopus spartii
New species
Italy
N/C
616
Dyfrolomyces phetchaburiensis
New species
Thailand
N/C
617
Fuscostagonosporaceae
New family
618
Fuscostagonospora cytisi
New species
Italy
N/C
N/C
619
Hermatomyces nabanheensis
New species
China
N/C
620
Hysterium rhizophorae
New species
Thailand
N/C
621
Rhytidhysteron
New record
622
Kirschsteiniothelia rostrata
New species
Thailand
N/C
623
Poaceascoma halophila
New species
Thailand
N/C
624
Setoseptoria scirpi
New species
UK
N/C
625
Alternariaster trigonosporus
New species
Russia
N/C
N/C
13
230
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
626
Hongkongmyces thailandica
New species
Thailand
N/C
627
Paucispora kunmingense
New species
China
N/C
628
Brooksia tropicalis
Description of type species of genus
Papua New Guinea
N/C
629
Microthyriaceae
New family
630
Dictyosporina
New genus
631
Dictyosporina ferruginea
New species
632
Lophium zalerioides
New species
Brazil
N/C
633
Nigrograna thymi
New species
Italy
N/C
634
Periconia aquatica
New species
Italy
N/C
635
Periconia minutissima
Reference specimen
China
N/C
636
Periconia submersa
New species
China
N/C
637
Acericola
New genus
Italy
N/C
638
Acericola italica
New species
639
Amarenomyces dactylidis
New species
Italy
N/C
640
Ophiosimulans plantaginis
New species
Italy
Paraophiobolus plantaginis (Qing Tian, Camporesi & K.D.
Hyde) Phookamsak Wanas. & K.D. Hyde
641
Phaeosphaeria acaciae
New species
China
N/C
642
Phaeopoacea muriformis
New species
China
N/C
643
Chaetothyrina artocarpi
New species
Thailand
N/C
644
Curvularia palmicola
New species
Thailand
N/C
645
Exserohilum turcicum
Description of type species of genus
Italy
N/C
646
Stemphylium vesicarium
New host
Italy
N/C
647
Carinispora nypae
New record
Brunei
N/C
648
Bryomyces scapaniae
Description of type species of genus
Austria
N/C
649
Immotthia hypoxylon
Description of type species of genus
US
Immotthia atrograna (Cooke & Ellis) M.E. Barr
Barr (1993)
650
Magnibotryascoma mali
New species
China
N/C
651
Bryochiton monascus
Description of type species of genus
Sweden
N/C
N/C
N/C
China
N/C
652
Tetraploa yakushimensis
New host
Thailand
653
Torula gaodangensis
New species
China
N/C
654
Chlamydotubeufia huaikangplaensis
New record
Thailand
N/C
655
Chlamydotubeufia krabiensis
New species
Thailand
N/C
656
Licopolia franciscana
Description of type species of genus
Brazil
N/C
657
Buelliella minimula
Description of type species of genus
US
N/C
658
Cocciscia hammeri
Description of type species of genus
Norway
N/C
659
Ramgea ozimecii
New species
Croatia
N/C
660
Otidea pruinosa
New species
China
N/C
661
Otidea stipitata
New species
China
N/C
662
Cytospora mali-sylvestris
New species
Russia
N/C
663
Diaporthe rhusicola
New combination
Italy
N/C
664
Diaporthe subclavata
New record
Thailand
N/C
665
Diatrypella vulgaris
New record
Thailand
Nom. inval., Art. 40.7 (Melbourne)
666
Endoxylina astroidea
Description of type species of genus
Sweden
Eutypa astroidea Rappaz. In: Mycol. helv. 2(3): 378. (1987)
N/C
667
Colletotrichum fioriniae
New host
Italy
668
Colletotrichum lauri
New species
Italy
N/C
669
Meliola clerodendri-infortunati
New species
Thailand
N/C
670
Ophiocordyceps cossidarum
New species
Thailand
N/C
671
Ophiocordyceps issidarum
New species
Thailand
N/C
13
Fungal Diversity (2022) 117:1–272
231
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
672
Longitudinalis
New genus
China
N/C
673
Longitudinalis nabanheensis
New species
674
Phaeoisaria microspora
New species
Thailand
N/C
675
Pleurothecium floriforme
New species
Thailand
N/C
676
Immersidiscosia eucalypti
New host
Italy
N/C
677
Seimatosporium italicum
New species
Italy
N/C
678
Grandibotrys hyalinus
New species
Thailand
N/C
679
Koorchalomella salmonispora
New species
Egypt
Ascomycota
Wijayawardene et al. (2022)
680
Xepicula leucotricha
New record
Thailand
Ascomycota
Wijayawardene et al. (2022)
681
Rizalia guianensis
New combination
British guiana
N/C
682
Astrocystis bambusicola
New species
Thailand
N/C
683
Collodiscula chiangraiensis
New species
Thailand
N/C
684
Fasciatispora calami
New species
Thailand
N/C
685
Vamsapriya breviconidiophora
New species
Thailand
N/C
686
Ascotrichella hawksworthii
Description of the type
species
Chile
N/C
687
Castellaniomyces
New genus
Italy
N/C
688
Castellaniomyces rosae
New species
689
Biciliopsis leptogiicola
Description of the type
species
Papua New Guinea
N/C
690
Agaricus purpurlesquameus
New species
Thailand
N/C
691
Agaricus rufusfibrillosus
New species
China
N/C
692
Tephrocybella constrictospora
New species
Italy
N/C
693
Steccherinum amapaense
New species
Brazil
N/C
694
Tyromyces amazonicus
New species
Brazil
N/C
695
Tyromyces angulatus
New species
Brazil
N/C
696
Lactifluus holophyllus
New species
Korea
N/C
697
Lactifluus luteolamellatus
New species
Korea
N/C
698
Lactifluus pseudohygrophoroides
New species
Korea
N/C
699
Russula benwooii
New species
US
N/C
700
Russula hypofragilis
New species
US
N/C
701
Russula obscurozelleri
New species
US
N/C
702
Russula parapallens
New species
US
N/C
703
Russula phoenicea
New species
Canada
N/C
704
Russula pseudopelargonia
New species
US
N/C
705
Russula pseudotsugarum
New species
US
N/C
706
Russula rhodocephala
New species
US
N/C
707
Russula salishensis
New species
US
N/C
708
Mortierella formicae
New species
Poland
N/C
N/C
N/C
Fungal diversity notes 709–839 (Wanasinghe et al. 2018)
709
Diplodia seriata
New host
Italy
710
Dothiorella iberica
New host
Italy
N/C
711
Lasiodiplodia theobromae
New host
UK
N/C
712
Neofusicoccum australe
New host
Thailand
N/C
713
Pseudocercospora rosae
New species
Uzbekistan
N/C
714
Endoconidioma rosae-hissaricae
New species
Italy
N/C
715
Angustimassarina quercicola
New host
Italy
N/C
716
Angustimassarina rosarum
New species
Uzbekistan
N/C
717
Astragalicola vasilyevae
New species
Italy
N/C
718
Epicoccum rosae
New species
Italy
N/C
719
Neoascochyta rosicola
New species
Italy
N/C
720
Paraconiothyrium rosae
New species
Italy
N/C
721
Paraphaeosphaeria michotii
New host
Italy
N/C
722
Paraphaeosphaeria rosae
New species
Sweden
N/C
13
232
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
723
Paraphaeosphaeria rosicola
New species
Italy
N/C
724
Pseudocamarosporium pteleae
New species
Russia
N/C
725
Pseudocamarosporium ulmiminoris
New species
Italy
N/C
726
Pseudopithomyces rosae
New species
Italy
N/C
727
Keissleriella rosacearum
New species
UK
N/C
728
Keissleriella rosarum
New species
Italy
Keissleriella rosacearum
729
Keissleriella rosae
New species
Italy
N/C
730
Keissleriella phragmiticola
New species
Italy
N/C
731
Murilentithecium rosae
New species
Italy
N/C
732
Phragmocamarosporium rosae
New species
UK
N/C
733
Pleurophoma pleurospora
New host
Uzbekistan
Dinemasporium pleurospora
Species Fungorum (2022a, b)
734
Setoseptoria arundelensis
New species
UK
N/C
735
Setoseptoria englandensis
New species
UK
N/C
736
Setoseptoria lulworthcovensis
New species
UK
N/C
737
Coelodictyosporium rosarum
New species
UK
N/C
738
Lophiostoma rosae
New species
Uzbekistan
N/C
739
Neopaucispora
New genus
Uzbekistan
N/C
740
Neopaucispora rosaecae
New species
741
Sigarispora caryophyllacearum
New species
Russia
N/C
742
Sigarispora caulium
New host
Italy
Sigarispora caulium (Fr.) Thambug., Wanas., Kaz. Tanaka &
K.D. Hyde, Index Fungorum 267: 1 (2015)
743
Sigarispora junci
New species
Italy
N/C
744
Sigarispora medicaginicola
New species
Russia
N/C
745
Sigarispora rosicola
New species
Russia
N/C
746
Sigarispora scrophulariae
New species
Russia
N/C
747
Sigarispora thymi
New species
Russia
N/C
748
Suttonomyces rosae
New species
Italy
N/C
749
Marjia
New genus
Uzbekistan
N/C
750
Marjia tianschanica
New species
751
Marjia uzbekistanica
New species
Uzbekistan
N/C
752
Melanocucurbitaria
New genus
Uzbekistan
N/C
753
Melanocucurbitaria uzbekistanica
New species
754
Melanodiplodia
New genus
Uzbekistan
N/C
755
Melanodiplodia tianschanica
New species
756
Monoseptella
New genus
Uzbekistan
N/C
757
Monoseptella rosae
New species
758
Muriformistrickeria rosae
New species
Italy
N/C
759
Muriformistrickeria rubi
New host
Sweden
N/C
760
Pseudostrickeria rosae
New species
Italy
N/C
761
Uzbekistanica
New genus
Uzbekistan
N/C
762
Uzbekistanica rosae-hissaricae
New species
763
Uzbekistanica yakutkhanika
New species
Uzbekistan
N/C
764
Xenomassariosphaeria
New genus
Italy
N/C
765
Xenomassariosphaeria rosae
New species
766
Bhatiellae
New genus
Italy
N/C
767
Bhatiellae rosae
New species
768
Dactylidina
New genus
Italy
N/C
769
Dactylidina dactylidis
New combination
Septoriella neodactylidis
Marlin-Felix et al. (2019a)
770
Dactylidina shoemakeri
New species
Italy
N/C
771
Dematiopleospora donetzica
New species
Russia
N/C
772
Dematiopleospora rosicola
New species
Italy
N/C
773
Dematiopleospora salsolae
New species
Uzbekistan
N/C
774
Embarria
New genus
Italy
N/C
13
Fungal Diversity (2022) 117:1–272
233
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
775
Embarria clematidis
New combination
776
Hawksworthiana
New genus
777
Hawksworthiana alliariae
New combination
Russia
Dlhawksworthia alliariae
Dlhawksworthia alliariae (Thambug., Camporesi & K.D.
Hyde) Wanas. & K.D. Hyde, in Wanasinghe et al., Index
Fungorum 357: 1 (2018)
778
Hawksworthiana clematidicola
New species
Italy
Dlhawksworthia clematidicola
Dlhawksworthia clematidicola (Thambug., Camporesi & K.D.
Hyde) Wanas. & K.D. Hyde, in Wanasinghe et al., Index
Fungorum 357: 1 (2018)
779
Hawksworthiana lonicerae
New species
Italy
Dlhawksworthia lonicerae
Dlhawksworthia lonicerae (Thambug., Camporesi & K.D.
Hyde) Wanas. & K.D. Hyde, in Wanasinghe et al., Index
Fungorum 357: 1 (2018)
780
Italica
New genus
Italy
N/C
781
Italica achilleae
New species
782
Italica luzulae
New combination
Italy
N/C
783
Neosetophoma rosarum
New species
Italy
N/C
784
Neosetophoma rosigena
New species
UK
N/C
785
Neosetophoma rosae
New species
Italy
N/C
786
Neostagonospora artemisiae
New species
Russia
N/C
787
Ophiobolus artemisiicola
New species
Russia
N/C
788
Poaceicola arundinis
sexual record
Italy
Septoriella arundinis
Marin-Felix et al. (2019a)
789
Poaceicola arundinicola
New species
Italy
Septoriella arundinicola
Marin-Felix et al. (2019a)
790
Poaceicola agrostina
New species
Italy
Septoriella agrostina
Marin-Felix et al. (2019a)
791
Poaceicola rosae
New species
Italy
Septoriella rosae
Marin-Felix et al. (2019a)
792
Populocrescentia ammophilae
New species
Italy
N/C
793
Populocrescentia rosae
New species
Uzbekistan
N/C
794
Sclerostagonospora rosicola
New species
Italy
N/C
795
Sclerostagonospora lathyri
New species
China
N/C
796
Sclerostagonospora rosae
New species
Italy
N/C
797
Wojnowicia rosicola
New species
Italy
Ascomycota
Wijayawardene et al. (2022)
798
Alternaria doliconidium
New species
Italy
N/C
799
Alternaria hampshirensis
New species
UK
N/C
800
Comoclathris rosae
New species
Italy
N/C
801
Comoclathris rosarum
New species
Italy
N/C
802
Comoclathris rosigena
New species
Italy
N/C
803
Pleospora rosae
New species
Italy
N/C
804
Pleospora rosae-caninae
New species
Italy
N/C
805
Sporormurispora
New genus
Russia
N/C
806
Sporormurispora atraphaxidis
New species
807
Sporormurispora pruni
New species
Uzbekistan
N/C
808
Teichospora rubriostiolata
New host
UK
Magnibotryascoma rubriostiolatum (Phukhamsakda et al.
2020)
809
Cycasicola
New genus
India
N/C
810
Cycasicola goaensis
New species
811
Neoconiothyrium
New genus
812
Neoconiothyrium rosae
New species
813
Parathyridaria rosae
New species
UK
N/C
814
Pararoussoella
New genus
UK
N/C
815
Pararoussoella rosarum
New species
Nom. illegit., Art. 53.1
Italy
N/C
Pseudoneoconiothyrium rosae
Pseudoneoconiothyrium rosae (Phukhams., Camporesi & K.D.
Hyde) Phukhams., Camporesi & K.D. Hyde, Index Fungorum 357: 1 (2018)
13
234
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
816
Penicillium aquaticum
New species
Korea
N/C
817
Penicillium acidum
New species
Korea
N/C
818
Amandinea punctata
New host
Sweden
N/C
819
Lecidella elaeochroma
New host
UK
N/C
820
Diaporthe eres
New host
UK
N/C
821
Diaporthe foeniculina
New host
Italy
N/C
822
Diaporthe rhusicola
New host
UK
N/C
823
Diaporthe rosae
New species
Thailand
N/C
824
Diaporthe rosicola
New species
UK
Diaporthe eres (Chaisiri et al. 2021)
825
Diaporthe rudis
New host
Italy
N/C
826
Bartalinia rosicola
New species
Italy
N/C
827
Broomella rosae
New species
Italy
N/C
828
Seiridium rosarum
New species
Italy
N/C
829
Seimatosporium rosicola
New species
Italy
Sporocadus rosigena (Liu et al. 2019a, b, c, d)
830
Seimatosporium rosigenum
New species
Italy
N/C
831
Paraxylaria
New genus
Uzbekistan
N/C
832
Paraxylaria rosacearum
New species
833
Castanediella camelliae
New species
Korea
N/C
834
Coniochaeta baysunika
New species
Uzbekistan
N/C
835
Coniochaeta rosae
New species
Uzbekistan
N/C
836
Backusella locustae
New species
Korea
N/C
837
Absidia jindoensis
New species
Korea
N/C
838
Mucor fluvius
New species
Korea
Mucor fluvii Hyang B. Lee, S.H. Lee & T.T.T. Nguyen [as
'fluvius']
839
Umbelopsis sinsidoensis
New species
Korea
N/C
Fungal diversity notes 840–928 (Tibpromma et al. 2018)
840
Cercospora capsici
New host
Thailand
N/C
841
Dictyocheirospora nabanheensis
New species
China
N/C
842
Dictyocheirospora pandanicola
New species
Thailand
N/C
843
Dictyocheirospora xishuangbannaensis
New species
China
N/C
844
Dictyosporium appendiculatum
New species
Thailand
N/C
845
Dictyosporium digitatum
New host
Hong Kong
N/C
846
Dictyosporium guttulatum
New species
Thailand
N/C
847
Dictyosporium hongkongensis
New species
Hong kong
N/C
848
Dictyosporium krabiense
New species
Thailand
N/C
849
Dictyosporium pandanicola
New species
Thailand
N/C
850
Deniquelata barringtoniae
New host
Thailand
N/C
851
Montagnula krabiensis
New species
Thailand
N/C
852
Pseudopithomyces pandanicola
New species
Thailand
N/C
853
Hermatomyces biconisporus
New species
China
N/C
854
Byssosphaeria siamensis
New host
Thailand
N/C
855
Neooccultibambusa thailandensis
New species
Thailand
N/C
856
Curvularia chonburiensis
New species
Thailand
N/C
857
Curvularia pandanicola
New species
Thailand
N/C
858
Curvularia thailandicum
New species
Thailand
Curvularia thailandica Tibpromma & K.D. Hyde [as 'thailandicum']
859
Curvularia xishuangbannaensis
New species
China
N/C
860
Roussoella solani
New host
China
Neoroussoella solani (Jayasiri et al. 2019)
861
Polyplosphaeria nabanheensis
New species
China
N/C
862
Polyplosphaeria pandanicola
New species
China
N/C
863
Polyplosphaeria xishuangbannaensis
New species
China
Ernakulamia xishuangbannaensis (Dong et al. 2020)
864
Rostriconidium pandanicola
New species
China
N/C
865
Torula chromolaenae
New host
China
N/C
866
Torula ficus
New host
Thailand
N/C
13
Fungal Diversity (2022) 117:1–272
235
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
867
Pseudoberkleasmium
New genus
China
N/C
868
Pseudoberkleasmium pandanicola
New species
869
Pseudochaetosphaeronema pandanicola
New species
Thailand
N/C
870
Lasiodiplodia chonburiensis
New species
Thailand
N/C
871
Lasiodiplodia hyalina
New host
Thailand
Lasiodiplodia thailandica (Zhang et al. 2021)
872
Lasiodiplodia pandanicola
New species
Thailand
Lasiodiplodia mahajangana (Zhang et al. 2021)
873
Lasiodiplodia pseudotheobromae
New host
Thailand
N/C
874
Neofusicoccum pandanicola
New species
China
Neofusicoccum parvum (Zhang et al. 2021)
875
Pseudofusicoccum adansoniae
New host
Thailand
N/C
876
Helicoma freycinetiae
New species
Thailand
N/C
877
Neohelicomyces pandanicola
New species
China
N/C
878
Tubeufia freycinetiae
New species
Thailand
N/C
879
Tubeufia inaequalis
New host
Thailand
N/C
880
Tubeufia pandanicola
New species
Thailand
N/C
881
Tubeufia parvispora
New species
Thailand
N/C
882
Yunnanomyces
New genus
China
N/C
883
Yunnanomyces pandanicola
New species
884
Stictis pandanicola
New species
China
Fitzroyomyces pandanicola (Wei et al. 2021)
885
Terriera pandanicola
New species
Thailand
N/C
886
Distoseptispora thailandica
New species
Thailand
N/C
887
Distoseptispora xishuangbannaensis
New species
China
N/C
888
Colletotrichum pandanicola
New species
Thailand
N/C
889
Malaysiascaceae
New family
890
Malaysiasca phaii
New host
Thailand
N/C
891
Acremoniisimulans
New genus
Thailand
N/C
892
Acremoniisimulans thailandensis
New species
893
Musicillium pandanicola
New species
Thailand
N/C
894
Clonostachys krabiensis
New species
Thailand
N/C
895
Lasionectria krabiense
New species
Thailand
N/C
896
Paracylindrocarpon nabanheensis
New species
China
N/C
897
Paracylindrocarpon pandanicola
New species
Hong Kong
N/C
898
Paracylindrocarpon xishuangbannaensis
New species
China
N/C
899
Cylindrocladiella xishuangbannaensis
New species
China
N/C
900
Pandanaceomyces
New genus
Thailand
N/C
901
Pandanaceomyces krabiensis
New species
902
Pseudoachroiostachys
New genus
Thailand
N/C
903
Pseudoachroiostachys krabiense
New species
904
Volutella krabiense
New species
Thailand
N/C
905
Volutella thailandensis
New species
Thailand
N/C
906
Pseudohyaloseta
New genus
Thailand
N/C
907
Pseudohyaloseta pandanicola
New species
908
Parasarcopodium hongkongensis
New species
Hong Kong
N/C
909
Pseudoornatispora
New genus
Thailand
N/C
910
Pseudoornatispora krabiense
New species
911
Sirastachys phangngaensis
New species
Thailand
N/C
912
Parascedosporium putredinis
New host
Thailand
N/C
913
Canalisporium krabiense
New species
Thailand
N/C
914
Canalisporium thailandensis
New species
Thailand
N/C
915
Dictyochaeta pandanicola
New species
China
N/C
916
Dictyochaeta siamensis
New host
Thailand
N/C
917
Menisporopsis pandanicola
New species
Hong Kong
N/C
918
Thozetella pandanicola
New species
Thailand
N/C
N/C
13
236
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
919
Chaetomium globosum
New record
Thailand
N/C
920
Beltrania krabiensis
New species
Thailand
N/C
921
Beltraniella pandanicola
New species
Thailand
N/C
922
Beltraniella thailandicus
New species
Thailand
N/C
923
Neopestalotiopsis chiangmaiensis
New species
Thailand
N/C
924
Neopestalotiopsis pandanicola
New species
China
N/C
925
Neopestalotiopsis phangngaensis
New species
Thailand
N/C
926
Pestalotiopsis krabiensis
New species
Thailand
N/C
927
Pestalotiopsis pandanicola
New species
Thailand
N/C
928
Vermiculariopsiella hongkongensis
New species
Hong Kong
N/C
N/C
Fungal diversity notes 929–1035 (Phookamsak et al. 2019)
929
Hyweljonesia indica
New species
India
930
Dictyocheirospora metroxylonis
New species
Thailand
N/C
931
Vicosamyces
New genus
Brazil
N/C
932
Vicosamyces venturisporus
New species
933
Keissleriella caraganae
New species
China
N/C
934
Murilentithecium lonicerae
New species
China
N/C
935
Phragmocamarosporium hederae
New host
China
Phragmocamarosporium hederae Wijayaw., R.K. Schumach. &
K.D. Hyde, Index Fungorum 370: 1 (2018)
936
Leptosphaeria urticae
New species
UK
N/C
937
Plenodomus artemisiae
New species
China
N/C
938
Plenodomus lijiangensis
New species
China
N/C
939
Plenodomus sinensis
New hosts and asexual
morph records
China
N/C
940
Sphaerellopsis isthmospora
New species
China
N/C
941
Sphaerellopsis paraphysata
New host
China
N/C
942
Lophiotrema mucilaginosis
New species
China
N/C
943
Brunneofusispora
New genus
China
N/C
944
Brunneofusispora sinensis
New species
945
Lonicericola
New genus
China
N/C
946
Lonicericola hyaloseptispora
New species
947
Parabambusicola thysanolaenae
New species
China
N/C
948
Paratrimmatostroma
New genus
China
N/C
949
Paratrimmatostroma kunmingensis
New species
950
Periconia cortaderiae
New host
China
N/C
951
Amarenomyces dactylidis
New host
China
N/C
952
Brunneomurispora
New genus
China
N/C
953
Brunneomurispora lonicerae
New species
954
Galiicola baoshanensis
New species
China
N/C
955
Neosetophoma lonicerae
New species
China
N/C
956
Ophiobolus malleolus
New species
Russia
N/C
957
Phaeosphaeria cycadis
New species
China
N/C
958
Wojnowiciella kunmingensis
New species
China
N/C
959
Wojnowiciella rosicola
New combination
India
N/C
960
Pseudoastrosphaeriellopsis
New genus
Thailand
N/C
961
Pseudoastrosphaeriellopsis
kaveriana
New species
962
Neoroussoella heveae
New species
Thailand
N/C
963
Neoroussoella leucaenae
New host
Thailand
N/C
964
Roussoella elaeicola
New species
China
N/C
965
Neobambusicola brunnea
New species
China
N/C
966
Liua
New genus
Brazil
N/C
967
Liua muriformis
New species
968
Lembosia brigadeirensis
New species
China
N/C
969
Dothiorella acericola
New species
Russia
N/C
13
Fungal Diversity (2022) 117:1–272
237
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
970
Dothiorella sarmentorum
New host
Thailand
N/C
971
Sphaeropsis eucalypticola
New host
India
N/C
972
Parazalerion
New genus
China
N/C
973
Parazalerion indica
New species
974
Muyocopron lithocarpi
New host
China
N/C
975
Pseudohelicomyces menglunicus
New species
Thailand
N/C
976
Cyphellophora filicis
New species
China
N/C
977
Capronia camelliae-yunnanensis
New species
China
N/C
978
Penicillium dokdoense
New species
Korea
N/C
979
Micropeltis phetchaburiensis
New species
Thailand
N/C
980
Velebitea
New genus
Croatia
N/C
981
Velebitea chrysotexta
New species
982
Sarcopeziza sicula
re-circumscribed
Italy
N/C
983
Conlarium thailandense
New species
Thailand
N/C
984
Cytospora ulmicola
New species
Russia
N/C
985
Septomelanconiella
New genus
Thailand
N/C
986
Septomelanconiella thailandica
New species
987
Pseudoplagiostoma mangiferae
New species
China
N/C
988
Coniella vitis
New host
Russia
N/C
989
Distoseptispora thysanolaenae
New species
China
N/C
990
Proliferophorum
New genus
Thailand
N/C
991
Proliferophorum thailandicum
New species
992
Plectosphaerella kunmingensis
New species
China
N/C
993
Leptobacillium leptobactrum
New record
India
N/C
994
Trichoderma koreanum
New species
Korea
N/C
995
Trichoderma pinicola
New species
Korea
N/C
996
Trichoderma rugulosum
New species
Korea
N/C
997
Emericellopsis koreana
New species
Korea
N/C
998
Canalisporium kenyense
Reference specimen
Thailand
N/C
999
Chaetosphaeria panamensis
New host
Taiwan
Paragaeumannomyces panamensis (Réblová et al. 2020)
1000
Thozetella lithocarpi
New species
Thailand
N/C
1001
Coniochaeta simbalensis
New species
India
N/C
1002
Tamsiniella labiosa
Reference specimen
China
N/C
1003
Zopfiella indica
New species
India
N/C
1004
Amphisphaeria mangrovei
New species
India
N/C
1005
Bartalinia kunmingensis
New species
China
N/C
1006
Robillarda mangiferae
New species
China
N/C
China
N/C
1007
Neoeutypella
New genus
1008
Neoeutypella baoshanensis
New species
1009
Peroneutypa mangrovei
New species
India
N/C
1010
Hypoxylon teeravasati
New species
India
N/C
1011
Agaricus memnonius
New species
China
N/C
1012
Agaricus langensis
New species
China
N/C
1013
Coprinus trigonosporus
New species
Saudi Arabia
N/C
1014
Amanita altipes
New record
Thailand
N/C
1015
Amanita flavoalba
New species
India
N/C
1016
Amanita melleialba
New record
Thailand
N/C
1017
Amanita subtropicana
New species
India
N/C
1018
Hygrocybe lucida
New species
India
N/C
1019
Marasmius indojasminodorus
New species
India
N/C
1020
Marasmiellus bicoloripes
New species
India
N/C
1021
Coprinopsis kubickae
New record
Croatia
N/C
1022
Baorangia major
New species
Thailand
N/C
1023
Baorangia rufomaculata
New combination
US
N/C
1024
Lanmaoa pallidorosea
New combination
US
N/C
13
238
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1025
Clavulina thindii
New species
India
N/C
1026
Phanerochaete australosanguinea
New species
Chile
N/C
1027
Lactarius olivaceopallidus
New species
India
N/C
1028
Lactifluus midnapurensis
New species
India
N/C
1029
Russula choptae
New species
India
N/C
1030
Russula uttarakhandia
New species
India
N/C
1031
Aleurodiscus patagonicus
New species
Chile
N/C
1032
Trechispora echinospora
New species
Equatorial Guinea
N/C
1033
Tremellochaete atlantica
New species
Brazil
N/C
1034
Dacrymyces invisibilis
New species
Chile
N/C
1035
Mucor orantomantidis
New species
Korea
N/C
Fungal diversity notes 1036–1150 (Hyde et al. 2019)
1036
Plectocarpon galapagoense
New species
Ecuador
N/C
1037
Caatingomyces
New genus
Brazil
N/C
1038
Caatingomyces brasiliensis
New species
1039
Amniculicola aquatica
New species
China
N/C
1040
Amniculicola guttulata
New species
China
N/C
1041
Murispora cicognanii
New record
China
N/C
1042
Angustimassarina sylvatica
New species
Italy
N/C
1043
Camarosporidiella mori
New species
Russia
N/C
1044
Protofenestella ulmi
New record
Russia
N/C
1045
Dictyocheirospora aquadulcis
New species
Thailand
1046
Dictyocheirospora rotunda
New geographical record China
1047
Dictyocheirospora taiwanense
New species
Taiwan
N/C
1048
Digitodesmium chiangmaiense
New species
Thailand
N/C
1049
Calophoma humuli
New species
Russia
N/C
1050
Neodidymelliopsis farokhinejadii
New host
Italy
N/C
1051
Neodidymelliopsis sambuci
New species
Italy
N/C
1052
Nothophoma quercina
New host
China
N/C
1053
Hermatomyces bauhiniae
New species
Thailand
N/C
1054
Vaginatispora palmae
New species
Thailand
N/C
1055
Vaginatispora armatispora
New record
Thailand
N/C
1056
Paramassaria
New genus
Thailand
N/C
1057
Paramassaria samaneae
New species
1058
Muriphaeosphaeria angustifoliae
New species
Uzbekistan
N/C
1059
Neosetophoma miscanthi
New species
Taiwan
N/C
1060
Neosetophoma salicis
New species
Uzbekistan
N/C
1061
Nodulosphaeria aquilegiae
New species
Italy
N/C
1062
Nodulosphaeria thalictri
New species
Italy
N/C
1063
Phaeopoacea asparagicola
New species
Ukraine
N/C
1064
Phaeosphaeria penniseti
New species
Taiwan
N/C
1065
Pseudoberkleasmiaceae
New family
1066
Pseudoberkleasmium chiangmaiense
New species
Thailand
N/C
1067
Pyrenochaetopsis sinensis
New species
China
N/C
1068
Tetraploa nagasakiensis
New geographical record China
N/C
1069
Torula breviconidiophora
New species
Thailand
N/C
1070
Torula polyseptata
New species
Thailand
N/C
1071
Falciformispora aquatica
New species
Thailand
N/C
1072
Acrogenospora sphaerocephala
Reference specimen
Thailand
N/C
1073
Acrogenospora thailandica
New species
Thailand
N/C
1074
Lembosia xyliae
New host
Thailand
N/C
1075
Morenoina palmicola
New host
Thailand
N/C
1076
Dothiorella plurivora
New record
China
N/C
1077
Dothiorella rhamni
New host
Russia
N/C
13
N/C
N/C
N/C
Fungal Diversity (2022) 117:1–272
239
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1078
Dothiorella styphnolobii
New species
Russia
N/C
1079
Dothiorella symphoricarposicola
New host
Italy
Dothiorella sarmentorum (Zhang et al. 2021)
1080
Lasiodiplodia iraniensis
New host
Thailand
N/C
1081
Jahnula queenslandica
New species
Australia
N/C
1082
Kamalomyces mangrovei
New species
Thailand
N/C
1083
Thysanorea uniseptata
New species
Thailand
N/C
1084
Penicillium circulare
New species
Korea
N/C
1085
Penicillium geumsanense
New species
Korea
N/C
1086
Penicillium mali
New species
Netherlands
N/C
1087
Penicillium psychrotrophicum
New species
Korea
N/C
1088
Penicillium wandoense
New species
Korea
N/C
1089
Prolixandromyces
syn. nov
N/C
1090
Prolixandromyces australis
New combination
N/C
1091
Prolixandromyces elongatus
New combination
N/C
1092
Prolixandromyces falcatus
New combination
N/C
1093
Prolixandromyces longispinae
New combination
N/C
1094
Prolixandromyces microveliae
New combination
N/C
1095
Prolixandromyces neoalardi
New combination
N/C
1096
Prolixandromyces polhemorum
New combination
N/C
1097. **
Prolixandromyces protuberans
New combination
N/C
1098. **
Prolixandromyces pseudoveliae
New combination
N/C
1099. **
Prolixandromyces tenuistipitis
New combination
N/C
1100
Prolixandromyces umbonatus
New combination
1101
Stigmatomyces chamaemyiae
New species
Malta
N/C
1102
Stigmatomyces cocksii
New species
Australia
N/C
1103
Stigmatomyces papei
New species
Tanzania
N/C
1104
Stigmatomyces tschirnhausii
New species
Kenya
N/C
1105
Stigmatomyces vikhrevii
New species
Malaysia
N/C
1106
Buellia viridula
New species
Thailand
N/C
1107
Lecidella yunnanensis
New species
China
N/C
1108
Micarea squamulosa
New species
Brazil
N/C
1109
Porina sorediata
New species
Brazil
N/C
1110
Cryptoschizotrema
New genus
Brazil
N/C
1111
Cryptoschizotrema cryptotrema
New combination
1112
Pilidium concavum
New geographical record Thailand
N/C
1113
Boubovia gelatinosa
New species
China
N/C
1114
Diaporthe italiana
New species
Italy
N/C
1115
Diaporthe rumicicola
New species
Italy
N/C
1116
Plagiostoma salicellum
New geographical record Italy
N/C
N/C
1117
Distoseptispora dehongensis
New species
China
N/C
1118
Distoseptispora palmarum
New species
Thailand
N/C
1119
Acrostalagmus annulatus
New geographical record Thailand
N/C
1120
Cordyceps succavus
New species
N/C
1121
Blackwellomyces lateris
New species
Thailand
N/C
1122
Sarocladium kiliense
New record
China
N/C
1123
Phaeoisaria siamensis
New species
Thailand
N/C
1124
Canalisporium dehongense
New species
China
N/C
1125
Sporoschisma chiangraiense
New species
Thailand
N/C
1126
Ellisembia aurea
New species
France
N/C
1127
Neolinocarpon rachidis
New record
Thailand
N/C
1128
Trochilispora
New genus
Brazil
N/C
1129
Trochilispora schefflerae
New species
1130
Pestalotiopsis aggestorum
New geographical record Thailand
N/C
1131
Castanediella meliponae
New species
Brazil
N/C
1132
Diatrypella delonicis
New species
Thailand
N/C
Thailand
13
240
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1133
Peroneutypa scoparia
New record
Thailand
N/C
1134
Fasciatispora arengae
New record
Thailand
N/C
1135
Cortinarius minusculus
New species
Finland
N/C
1136
Cortinarius subscotoides
New species
Finland
N/C
1137
Coprinopsis psammophila
New species
Libya
N/C
1138
Coprinopsis villosa
New geographical record Croatia
N/C
1139
Neoacladium
New genus
India
N/C
1140
Neoacladium indicum
New species
1141
Cantharellus goossensiae
New record
China
N/C
1142
Cantharellus brunneopallidus
New species
Madagascar
N/C
1143
Cantharellus griseotinctus
New species
Madagascar
N/C
1144
Fomitiporia carpinea
New species
China
N/C
1145
Fomitiporia lagerstroemiae
New species
Vietnam
N/C
1146
Grammothele aurantiaca
New species
Brazil
N/C
1147
Grammothele micropora
New species
Brazil
N/C
1148
Grammothele brasiliensis
New record
Brazil
N/C
1149
Russula prasina
New species
China
N/C
1150
Rhizophydium koreanum
New species
Korea
N/C
Fungal diversity notes 1151–1276 (Hyde et al. 2020a, b, c)
1151
Dissoconium eucalypti
New record of the sexual Italy
morph
1152
Pseudocercospora maetaengensis
New species
Thailand
N/C
1153
Hysterobrevium constrictum
New record
China
N/C
1154
Rhytidhysteron camporesii
New species
China
N/C
1155
Rhytidhysteron erioi
New species
Thailand
N/C
1156
Angustimassarina camporesii
New species
Italy
N/C
1157
Camarosporidiella camporesii
New species
Italy
N/C
1158
Foliophoma camporesii
New species
Italy
N/C
1159
Dendryphiella phitsanulokensis
New host
Thailand
N/C
1160
Dictyosporium muriformis
New species
China
N/C
1161
Ascochyta medicaginicola
New record
Italy
N/C
1162
Ascochyta pisi
New host
Italy
Didymella pisi (Chilvers et al. 2009)
1163
Didymella camporesii
New species
Italy
N/C
1164
Didymella macrostoma
New host
Italy
N/C
1165
Neodidymelliopsis camporesii
New species
Italy
N/C
1166
Neodidymelliopsis ranunculi
New host
Italy
N/C
1167
Nothophoma quercina
New host
Russia
N/C
1168
Xenodidymella camporesii
New species
Italy
N/C
1169
Kalmusia erioi
New species
Thailand
N/C
1170
Montagnula camporesii
New species
Italy
N/C
1171
Neokalmusia kunmingensis
New species
China
N/C
1172
Pseudocamarosporium camporesii
New species
Italy
N/C
1173
Tremateia lamiacearum
New species
China
N/C
1174
Tremateia camporesii
New species
Thailand
N/C
1175
Fuscostagonospora camporesii
New species
Italy
N/C
1176
Brunneoclavispora camporesii
New species
Thailand
N/C
1177
Keissleriella camporesiana
New species
Italy
N/C
1178
Keissleriella camporesii
New species
Italy
N/C
1179
Pseudomurilentithecium
New genus
Italy
N/C
1180
Pseudomurilentithecium camporesii
New species
1181
Plenodomus triseptatus
New species
Russia
N/C
1182
Neovaginatispora fuckelii
New host
China
N/C
1183
Pseudochaetosphaeronema kunmingense
New species
China
N/C
13
N/C
Fungal Diversity (2022) 117:1–272
241
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1184
Camposporium appendiculatum
New species
China
N/C
1185
Camposporium lycopodiellae
New combination
1186
Camposporium multiseptatum
New species
China
N/C
1187
Camposporium pellucidum
New record
China
N/C
1188
Camposporium septatum
New species
Thailand
N/C
1189
Uzbekistanica pruni
New species
Russia
N/C
1190
Occultibambusa bambusae
New host
Taiwan
N/C
1191
Paramonodictys
New genus
China
N/C
1192
Paramonodictys solitarius
New species
1193
Periconia palmicola
New species
Thailand
N/C
1194
Bhagirathimyces
New genus
India
N/C
1195
Bhagirathimyces himalayensis
New species
1196
Loratospora arezzoensis
New species
Italy
N/C
1197
Neosetophoma camporesii
New species
Italy
N/C
1198
Paraloratospora
New genus
Italy
N/C
1199
Paraloratospora camporesii
New species
1200
Paraloratospora gahniae
New combination
1201
Phaeosphaeria chinensis
New host
Taiwan
N/C
1202
Phaeosphaeriopsis pseudoagavacearum
New record of the sexual Italy
morph
N/C
1203
Septoriella camporesii
New species
Italy
N/C
1204
Wojnowiciella dactylidis
New host
Italy
N/C
1205
Ernakulamia tanakae
New species
India
N/C
1206
Pseudotetraploa rajmachiensis
New species
India
N/C
1207
Tetraploa dwibahubeeja
New species
India
N/C
1208
Tetraploa pseudoaristata
New species
India
N/C
1209
Tetraploa thrayabahubeeja
New species
India
N/C
1210
Tetraploa sasicola
New host
Taiwan
N/C
1211
Torula camporesii
New species
China
N/C
1212
Torula gaodangensis
New host
China
N/C
1213
Falciformispora uttaraditensis
New species
Thailand
N/C
1214
Diplodia mutila
New host
Italy
N/C
1215
Diplodia seriata
New host
Italy
N/C
1216
Setoapiospora
New genus
Thailand
N/C
1217
Setoapiospora thailandica
New species
1218
Camporesiomyces
New genus
China
N/C
1219
Camporesiomyces mali
New species
1220
Camporesiomyces patagoniensis
New combination
1221
Camporesiomyces vaccinii
New combination
1222
Verruconis mangrovei
New species
1223
Eriomycetaceae
New family
1224
Eriomyces
New genus
1225
Eriomyces heveae
New species
1226
Pyxine berteriana
1227
Heterosphaeria patella
1228
Rhexocercosporidium microsporum New combination
1229
Rhexocercosporidium senecionis
1230
1231
N/C
N/C
N/C
N/C
India
N/C
Thailand
N/C
New host
China
N/C
New host
Italy
N/C
New species
Italy
N/C
Srinivasanomyces
New genus
India
N/C
Srinivasanomyces kangrensis
New species
1232
Helvella subtinta
New species
China
N/C
1233
Wilcoxina verruculosa
New species
China
N/C
1234
Eriocamporesia
New genus
Thailand
N/C
1235
Eriocamporesia aurantia
New species
1236
Cytospora fusispora
New species
India
N/C
1237
Cytospora rosigena
New species
Russia
N/C
N/C
Cadophora microspora (Ekanayaka et al. 2019)
13
242
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1238
Diaporthe camporesii
New species
Italy
N/C
1239
Diaporthe cynaroidis
New record of the
sexual-asexual connection
Italy
N/C
1240
Diaporthe foeniculina
New host
Italy
N/C
1241
Diaporthe nigra
New species
Italy
N/C
1242
Neomyrmecridium guizhouense
New species
China
N/C
1243
Lanspora cylindrospora
New species
India
N/C
1244
Pseudoconlarium
New genus
China
N/C
1245
Pseudoconlarium punctiforme
New species
1246
Colletotrichum hederiicola
New species
Italy
N/C
1247
Acremonium chiangraiense
New species
Thailand
N/C
1248
Clonostachys eriocamporesiana
New species
Thailand
N/C
1249
Clonostachys eriocamporesii
New species
Thailand
N/C
1250
Mariannaea atlantica
New species
Brazil
N/C
1251
Conioscypha verrucosa
New species
China
N/C
1252
Neomonodictys
New genus
Thailand
N/C
1253
Neomonodictys muriformis
New species
1254
Canalisporium aquaticium
New species
Thailand
N/C
1255
Coniochaeta vineae
New species
China
N/C
1256
Pseudodactylaria camporesiana
New species
Thailand
N/C
1257
Neoleptosporella camporesiana
New species
Thailand
Nom. inval., Art. 35.1 (Shenzhen)
1258
Arthrinium marii
New host
Italy
N/C
1259
Pseudotruncatella camporesii
New species
Italy
N/C
1260
Pseudopestalotiopsis theae
New record
China
N/C
1261
Diatrypella yunnanensis
New species
China
N/C
1262
Fasciatisporaceae
New family
1263
Fasciatispora cocoes
New species
Thailand
N/C
1264
Astrocystis bambusicola
New record
China
N/C
1265
Melanographium phoenicis
New species
Thailand
N/C
1266
Xenoanthostomella
New genus
Thailand
N/C
1267
Xenoanthostomella chromolaenae
New species
1268
Diutina bernali
New species
Panama
N/C
1269
Diutina sipiczkii
New species
Panama
N/C
1270
Cortinarius ainsworthii
New species
Belgium
N/C
1271
Cortinarius aurae
New species
UK
N/C
1272
Cortinarius britannicus
New species
UK
N/C
1273
Cortinarius heatherae
New species
UK
N/C
1274
Cortinarius scoticus
New species
UK
N/C
1275
Cortinarius subsaniosus
New species
UK
N/C
1276
Adustochaete nivea
New species
Brazil
N/C
N/C
Fungal diversity notes 1276–1386 (Yuan et al. 2020)
1277
Umbelopsis heterosporus
New species
Brazil
N/C
1278
Aplosporella prunicola
New host
China
N/C
1279
Diplodia torilicola
New species
Italy
N/C
1280
Neodevriesia manglicola
New species
India
N/C
N/C
1281
Coniothyrium triseptatum
New species
China
1282
Neodidymelliopsis salviae
New species
Italy
N/C
1283
Neodidymelliopsis urticae
New species
Italy
N/C
1284
Magnopulchromyces
New genus
Brazil
N/C
1285
Magnopulchromyces scorpiophorus
New species
1286
Paradictyoarthrinium diffractum
New host
India
N/C
1287
Ophiosphaerella chiangraiensis
New species
Thailand
N/C
1288
Prosthemium betulinum
New record
Italy
N/C
13
Fungal Diversity (2022) 117:1–272
243
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1289
Neoroussoella magnoliae
New species
China
N/C
1290
Sporormurispora paulsenii
New species
Uzbekistan
Sporormurispora paulsenii D. Pem, Gafforov & K.D. Hyde,
Index Fungorum 468: 1 (2021)
1291
Helicoarctatus thailandicus
New species
Thailand
N/C
1292
Thaxteriellopsis obliqus
New species
India
N/C
1293
Scleroramularia vermispora
New species
China
N/C
1294
Graphis supracola
Thailand
N/C
1295
Podosphaera yulii
New species
China
N/C
1296
Golovinomyces monardae
New record
China
N/C
1297
Ionopezia
New genus
1298
Ionopezia gerardii
New combination
1299
Hansenopezia
New genus
1300
Hansenopezia retrocurvata
New combination
1301
Hansenopezia decora
New species
Croatia
N/C
1302
Neottiella gigaspora
New species
China
N/C
1303
Arthrinium sorghi
New species
Brazil
N/C
1304
Chloridium macrocladum
New combination
Taiwan
Nom. inval., Art. 41.5 (Shenzhen)
1305
Diaporthe pimpinellae
New species
Italy
N/C
1306
Moelleriella gracilispora
New species
China
N/C
1307
Trichoderma ceratophylletum
New species
China
N/C
1308
Hirsutella hongheensis
New species
China
N/C
1309
Colletotrichum parthenocissicola
New species
Russia
N/C
1310
Dictyosporella guizhouensis
New species
China
N/C
1311
Xylolentia reniformis
New species
China
N/C
1312
Catenuliconidia
New genus
China
N/C
1313
Catenuliconidia uniseptata
New species
1314
Lycoperdon lahorense
New species
Pakistan
N/C
1315
Lycoperdon pseudocurtisii
New species
Pakistan
N/C
1316
Cortinarius indorusseus
New species
India
N/C
1317
Cortinarius paurigarhwalensis
New species
India
N/C
1318
Cortinarius sinensis
New species
China
N/C
1319
Cortinarius subsanguineus
New species
China
N/C
1320
Cortinarius xiaojinensis
New species
China
N/C
1321
Humidicutis brunneovinacea
New species
Mexico
N/C
1322
Amyloceraceomyces
New genus
China
N/C
1323
Amyloceraceomyces angustisporus
New species
1324
Amylocorticium ellipsosporum
New species
China
N/C
1325
Clavulina sphaeropedunculata
New species
Mexico
N/C
1326
Lentaria gossypina
New species
Mexico
N/C
1327
Lentaria variabilis
New species
Mexico
N/C
1328
Fuscoporia licnoides
New combination
Brazil
N/C
1329
Fuscoporia marquesiana
New species
Brazil
N/C
1330
Fuscoporia scruposa
New combination
Brazil
N/C
1331
Fuscoporia semiarida
New species
Brazil
N/C
1332
Rigidoporus juniperinus
New species
Uzbekistan
N/C
1333
Rhodofomitopsis pseudofeei
New species
Australia
N/C
1334
Rhodofomitopsis monomitica
New combination
1335
Rhodofomitopsis oleracea
New combination
1336
Antrodiella descendena
New combination
Malaysia
N/C
1337
Tyromyces minutulus
New species
China
N/C
1338
Russula benghalensis
New species
India
N/C
1339
Tomentella asiae-orientalis
New species
China
N/C
1340
Tomentella atrobadia
New species
China
N/C
1341
Tomentella atrocastanea
New species
China
N/C
Published 9 July 2020; see Ionopezia Matočec, I. Kušan &
Jadan 2020 (published 31st Oct. 2020)
Croatia
N/C
N/C
N/C
N/C
13
244
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1342
Tomentella aureomarginata
New species
China
N/C
1343
Tomentella brevis
New species
China
N/C
1344
Tomentella brunneoflava
New species
China
N/C
1345
Tomentella brunneogrisea
New species
China
N/C
1346
Tomentella capitatocystidiata
New species
China
N/C
1347
Tomentella changbaiensis
New species
China
N/C
1348
Tomentella citrinocystidiata
New species
China
N/C
1349
Tomentella coffeae
New species
China
N/C
1350
Tomentella conclusa
New species
China
N/C
1351
Tomentella cystidiata
New species
China
N/C
1352
Tomentella dimidiata
New species
China
N/C
1353
Tomentella duplexa
New species
China
N/C
1354
Tomentella efibulata
New species
China
N/C
1355
Tomentella efibulis
New species
China
N/C
1356
Tomentella farinosa
New species
China
N/C
1357
Tomentella flavidobadia
New species
China
N/C
1358
Tomentella fuscocrustosa
New species
China
N/C
1359
Tomentella fuscofarinosa
New species
China
N/C
1360
Tomentella fuscogranulosa
New species
China
N/C
1361
Tomentella fuscopelliculosa
New species
China
N/C
1362
Tomentella globospora
New species
China
N/C
1363
Tomentella gloeocystidiata
New species
China
N/C
1364
Tomentella griseocastanea
New species
China
N/C
1365
Tomentella griseofusca
New species
China
N/C
1366
Tomentella griseomarginata
New species
China
N/C
1367
Tomentella inconspicua
New species
China
N/C
1368
Tomentella incrustata
New species
China
N/C
1369
Tomentella interrupta
New species
China
N/C
1370
Tomentella liaoningensis
New species
China
N/C
1371
Tomentella longiaculeifera
New species
China
N/C
1372
Tomentella longiechinuli
New species
China
N/C
1373
Tomentella megaspora
New species
China
N/C
1374
Tomentella olivacea
New species
China
N/C
1375
Tomentella olivaceobrunnea
New species
China
N/C
1376
Tomentella pallidobrunnea
New species
China
N/C
1377
Tomentella pallidomarginata
New species
China
N/C
1378
Tomentella parvispora
New species
China
N/C
1379
Tomentella pertenuis
New species
China
N/C
1380
Tomentella qingyuanensis
New species
China
N/C
1381
Tomentella segregata
New species
China
N/C
1382
Tomentella separata
New species
China
N/C
1383
Tomentella stipitata
New species
China
N/C
1384
Tomentella storea
New species
China
N/C
1385
Scytinopogon minisporus
New species
Mexico
N/C
1386
Phaeotremella yunnanensis
New species
China
N/C
Fungal diversity notes 1387–1577 (Boonmee et al. 2021)
1387
Diplodia alanphillipsii
New species
Iran
N/C
1388
Chaetoscutula juniperi
New record
Spain
N/C
1389
Dothiora coronicola
New species
Italy
1390
Dyfrolomyces distoseptatus
New geographical record Thailand
N/C
1391
Aliquandostipite khaoyaiensis
New record
Thailand
N/C
1392
Minutisphaera thailandensis
New species
Thailand
N/C
1393
Acrocalymma fici
New habitat and geographical record
Thailand
N/C
1394
Dictyosporium pandanicola
New record
Thailand
N/C
13
N/C
Fungal Diversity (2022) 117:1–272
245
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1395
Camarosporidiella laburni
New record
Uzbekistan
N/C
1396
Cryptocoryneum rosae
New species
China
N/C
1397
Didymella azollae
New species
Iran
N/C
1398
Montagnula thailandica
New record
Thailand
N/C
1399
Paraconiothyrium ajrekarii
New species
India
N/C
1400
Spegazzinia camelliae
New record
Thailand
N/C
1401
Hermatomyces nabanheensis
New record
China
N/C
1402
Hermatomyces sphaericoides
New record
Thailand
N/C
1403
Poaceascoma taiwanense
New geographical record Thailand
N/C
1404
Hongkongmyces kokensis
New species
Thailand
N/C
1405
Xenovaginatispora
New genus
Thailand
N/C
1406
Xenovaginatispora phichaiensis
New species
1407
Longipedicellata aquatica
New record
Thailand
N/C
1408
Longipedicellata megafusiformis
New species
Thailand
N/C
1409
Submersispora variabilis
New record
Thailand
N/C
1410
Lophiostoma caudatum
New host
Uzbekistan
Sigarispora caudata
Thambugala et al. (2015)
1411
Lophiostoma clematidis-vitalbae
New host
Uzbekistan
N/C
1412
Vaginatispora nypae
New record
Thailand
N/C
1413
Lophiotrema hydei
New host
China
N/C
1414
Lophiotrema lincangensis
New species
China
N/C
1415
Lophiotrema neoarundinariae
New host
China
N/C
1416
Pseudochaetosphaeronema chiangraiense
New species
Thailand
N/C
1417
Helminthosporium chiangraiense
New species
Thailand
N/C
1418
Nigrograna jinghongensis
New species
China
N/C
1419
Seriascoma honghense
New species
China
N/C
1420
Paradictyoarthrinium diffractum
New record
Thailand
N/C
1421
Pleopunctum thailandicum
New species
Thailand
N/C
1422
Phaeosphaeriopsis aloes
New host
China
N/C
1423
Comoclathris permunda
Reference specimen
Uzbekistan
N/C
1424
Lepidosphaeria strobelii
New species
India
N/C
1425
Ernakulamia tanakae
New record
Thailand
N/C
1426
Thyridaria aureobrunnea
New species
Thailand
N/C
1427
Cylindrotorula
New genus
1428
Cylindrotorula indica
New species
India
N/C
1429
Dendryphion hydei
New record
China
N/C
1430
Torula lancangjiangensis
New species
China
N/C
1431
Torula mackenziei
New record
China
N/C
1432
Wicklowia fusiformispora
New species
Thailand
N/C
1433
Wicklowia submersa
New geographical record Thailand
N/C
1434
Helicosporium luteosporum
New record
Thailand
N/C
1435
Helicosporium sexuale
New species
Thailand
N/C
1436
Neohelicosporium irregulare
New record
Thailand
N/C
1437
Neohelicosporium parvisporum
New record
Thailand
N/C
1438
Tubeufia chiangmaiensis
New record
Thailand
N/C
1439
Tubeufia longihelicospora
New species
Thailand
N/C
1440
Tubeufia roseohelicospora
New record
Thailand
N/C
1441
Megacapitula villosa
New geographical record Thailand
N/C
1442
Cladophialophora abundans
New geographical record Thailand
N/C
1443
Cladophialophora aquatica
New species
Thailand
N/C
1444
Aspergillus lannaensis
New species
Thailand
N/C
1445
Pseudobactrodesmium stilboideum
New combination
Thailand
N/C
1446
Hydrophilomyces hydraenae
New species
US
N/C
1447
Laboulbenia divisa
New species
Costa Rica
N/C
N/C
13
246
Fungal Diversity (2022) 117:1–272
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1448
Laboulbenia triarthronis
New species
US
N/C
1449
Mimeomyces digitatus
New species
Ecuador
N/C
1450
Synandromyces makranczyi
New species
Peru
N/C
1451
Erysiphe salicicola
New species
Korea
N/C
1452
Scolecoleotia
New genus
Italy
N/C
1453
Scolecoleotia eriocamporesi
New species
1454
Coryneum fagi
New species
China
N/C
1455
Diaporthe chamaeropicola
New species
Portugal
N/C
1456
Diaporthe foeniculina
New record
Portugal
N/C
1457
Diaporthe pseudophoenicicola
New record
Portugal
N/C
1458
Diaporthe pyracanthae
New record
Portugal
N/C
1459
Phaeocytostroma yomense
New species
Thailand
N/C
1460
Parafuscosporella nilotica
New species
Egypt
N/C
1461
Fusarium atrovinosum
New record
Australia
N/C
1462
Fusarium clavum
New record
Australia
N/C
1463
Fusarium queenslandicum
New species
Australia
N/C
1464
Mariannaea camelliae
New species
Thailand
N/C
1465
Thyronectria caudata
New record
Uzbekistan
N/C
1466
Phaeoisaria aquatica
New record
Thailand
N/C
1467
Phaeoisaria synnematicus
New species
India
N/C
1468
Pleurotheciella dimorphospora
New species
China
N/C
1469
Pseudodactylaria albicolonia
New species
Thailand
N/C
1470
Canalisporium caribense
New record
Thailand
N/C
1471
Rhexoacrodictys nigrospora
New species
Thailand
N/C
1472
Cercophora dulciaquae
New species
Thailand
N/C
1473
Khaleijomyces umikazeanus
New species
Japan
N/C
1474
Eutypa flavovirens
New record
Italy
N/C
1475
Eutypa lata
New record
Italy
N/C
1476
Xylaria apiospora
New species
India
N/C
1477
Xylaria haemorrhoidalis
New record
India
N/C
1478
Melanographium smilacis
New species
Thailand
N/C
1479
Chlorophyllum demangei
New record
Laos
N/C
1480
Chlorophyllum globosum
New record
Laos
N/C
1481
Chlorophyllum hortense
New record
Laos
N/C
1482
Micropsalliota globocystis
New record
Laos
N/C
1483
Micropsalliota gracilis
New record
Laos
N/C
1484
Xanthagaricus necopinatus
New record
Laos
N/C
1485
Saproamanita manicata
New record
Thailand
Amanita manicata
1486
Cortinarius alutarius
New species
Poland
N/C
1487
Cortinarius mammillatus
New species
Poland
N/C
1488
Cortinarius quercoflocculosus
New species
Poland
N/C
1489
Laccaria populina
New species
Italy
N/C
1490
Hygrocybe boertmannii
New species
India
N/C
1491
Marasmius benghalensis
New species
India
N/C
1492
Marasmius jinfoshanensis
New species
China
N/C
1493
Marasmius subtropicus
New species
India
N/C
1494
Cruentomycena uttarakhandina
New species
India
N/C
1495
Cyathus uniperidiolus
New species
India
N/C
1496
Marasmiellus palmivorus
New record
Thailand
N/C
1497
Coprinellus punjabensis
New species
Pakistan
N/C
1498
Geastrum gorgonicum
New species
Cape
N/C
1499
Geastrum hansagiense
New species
Hungary
N/C
1500
Hyphodontia yunnanensis
New species
China
N/C
1501
Odontia huanrenensis
New species
China
Nom. inval., Art. 40.7 (Shenzhen)
1502
Odontia parvispina
New species
China
N/C
1503
Hyphoderma australosetigerum
New species
Chile
N/C
13
Fungal Diversity (2022) 117:1–272
247
Table 5 (continued)
FDN number
Species name
Status
Country/Region
Comment
1504
Efibula rodriguezarmasiae
New species
Spain
N/C
1505
Phanerochaete hainanensis
New species
China
N/C
1506
Favolus septatus
New record
India
N/C
1507
Lactarius pallidozonarius
New species
China
N/C
1508
Russula paravioleipes
New species
China
N/C
1509
Microbotryum polycnemoides
New species
Turkey
N/C
1510
Mortierella solitaria
New species
Austria
Nom. inval., Art. 40.7 (Shenzhen)
1511
Mucor harpali
New species
Korea
N/C
*is FDN number that appear different in text than FDN number of itself in Table of Contents in the publication
N/C-no change
Key to the smut fungi on Anadelphia, Elymandra, and
Monocymbium (modified after Denchev and Denchev
2016).
1 Sori in leaves and spatheoles………… ………
……………………………………………………………2
1*Sori in racemes, spikelets or ovaries………………
……………………………………………………………3.
2 Sori form non-erumpent streaks or patches on leaves
and spatheoles. Spores (9–)9.5–15.5(–18.5) μm long,
spore wall 1.2–3.0(–3.8) μm thick. [On Anadelphia
pumila]………………………………………………………
…………………………… Jamesdicksonia anadelphiae.
2*Sori form erumpent streaks on leaves and spatheoles. Spores (11.5–)13–23.5(–26.5) μm long, spore
wall (2.0–)2.5–7.0(–9.0) μm thick. [On Anadelphia
trichaeta]………Jamesdicksonia anadelphiae-trichaetae.
3 Sori in some ovaries of an inflorescence. [On Elymandra androphila]………………………………………4.
3*Sori in racemes or spikelets…………………………5.
4 Spores 10.5–15 μm long; spore walls echinate, with
1–1.5 μm high spines……Macalpinomyces elymandrae.
4* Spores 17–27 μm long; spore walls with apically flattened or rounded, densely spaced projections, 1–2.5(–3)
μm high………………………………Tilletia elymandrae.
5 Spore balls present. Spores dimorphic. Outer spores
(9.5–)10.5–15(–16) μm long. Sterile cells absent. [On Anadelphia pumila]……………Anthracocystis anadelphiae.
5* Spore balls absent. Spores not dimorphic, smaller.
Sterile cells present………………………………………6.
6 Sori destroying all racemes. Spores minutely
echinulate-verruculose, with ornaments up to 0.2 μm
in height. [On Anadelphia leptocoma]……… ………
……………Sporisorium anadelphiae-leptocomae.
6* Sori in spikelets or groups of spikelets. Spore ornamentation echinulate, with higher ornaments……………7.
7 Spore wall moderately echinulate, spinules up to
0.7 μm high. Spores (8.5–)9–11.5(–12.5) (10.5 ± 0.7)
μm long. [On Anadelphia trichaeta]……… ………
……………………Sporisorium anadelphiae-trichaetae.
7*Spore wall minutely echinulate, spinules up to
0.3(–0.4) μm high. Spores (7–)7.5–10.5(–11.5) (9.1 ± 0.6)
μm long. [On Monocymbium ceresiiforme]………………
………………………Sporisorium monocymbii. (Table 5)
Acknowledgements We thank Prof. Shaun Pennycook for revising the
Latin names. Ruvishika S. Jayawardena thanks the National Research
Council of Thailand (NRCT) grant “Biodiversity, taxonomy, phylogeny and evolution of Colletotrichum in northern Thailand” (grant no.
NRCT-TRG010), Thailand Science Research and Innovation (TSRI)
grant “Biodiversity, taxonomy, phylogeny and evolution of Colletotrichum on Avacado, Citrus, Durian and Mango in northern Thailand” (grant no. 652A01003) and Mae Fah Luang University grant
“Identification of fungicolous fungi in northern Thailand” (grant no.
651B01010) grants for funding this research. Kevin D. Hyde thanks
the National Research Council of Thailand (NRCT) grant “Total fungal
diversity in a given forest area with implications towards species numbers, chemical diversity and biotechnology” (grant no. N42A650547),
Thailand Science Research and Innovation (TSRI) grant “Macrofungi
diversity research from the Lancang-Mekong Watershed and Surrounding areas” (grant no. DBG6280009) and the Basic Research
Fund (Grant No. 652A01001), entitled “Studies of fungi associated
with Asteraceae and the discovery of biological properties”. K.D.
Hyde also thanks the visiting professorship offered by Chiang Mai
University. Nopparat Wannathes, Wiphawanee Phonrob, Nakarin
Suwannarach and Jaturong Kumla thank the Plant Genetic Conservation Project under the Royal initiative of Her Royal Highness Princess
Maha Chakri Sirindhorn, Pibulsongkram Rajabhat University, Thailand. The field collection in Khao Yai National Park is under permit
number 0907.4/13702 and in Si Satchanalai Nationa Park is under
permit numbers 0907.4/13696 and 0907.4/15381. André Aptroot,
Maria F. Souza, Lidiane A. dos Santos and Adriano A. Spielmann
thank the colleagues at the Botanical Laboratory of the UFMS for
organizing the excursion, Sandro Pereira for guidance in the field,
Wellington Fava for extracting DNA and PCR, and Edna Maria Facincani for information about geology. This study was financed in part
by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—Brasil (CAPES)—Finance Code 001 who provided a visiting
professorship to the first author. Many thanks to Wellington Fava for
generating the sequences. Jadson D.P. Bezerra and colleagues thank
the CNPq, CAPES (finance code 001) and FACEPE for financial support and scholarship. The study of Cvetomir M. Denchev and Teodor
T. Denchev received support from the SYNTHESYS Project http://
www. synth esys. info/, financed by European Community Research
Infrastructure Action under the FP7 “Capacities” Program at the Royal
Botanic Garden, Madrid (Grant no. ES-TAF-6618). The research of
C.C. Chen is supported by the Ministry of Science and Technology of
13
248
ROC (Taiwan) (Grant No. MOST 107-2621-B178-002-MY3). Chun
Y. Deng acknowledges the Science and Technology Support Project
of Guizhou Province (Project no. 20192451-2) for research support.
Yusufjon Gafforov acknowledges the Ministry of Innovative Development of the Republic of Uzbekistan (nos. AL 2021090820, MUK-202146) and the State Scientific and Technical Program of Uzbekistan for
research support. Sugantha Gunaseelan and Kezhocuyi Kezo thank
DST, SERB-EMR (EMR/2016/003078) for the financial assistance;
Kezhocuyi Kezo acknowledges SEED-DST (SP/YO/007/2008) grant.
Mark S. Calabon is grateful to the Mushroom Research Foundation and
Department of Science and Technology-Science Education Institute
(Philippines). E.B. Gareth Jones is supported under the Distinguished
Scientist Fellowship Program (DSFP), King Saud University, Kingdom
of Saudi Arabia. André L. Firmino, Vanessa P. Abreu and Olinto L.
Pereira thank the CNPq, FAPEMIG and CAPES (Finance Code 001)
for the financial support the guava producer of the city of Piraúba,
for supplying fundamental materials for the accomplishment of this
work. W. Rossi and M. Leonardi wish to thank the Italian entomologists who supplied them with the insects bearing the new species of
Laboulbeniales: A. Bordoni, P. Magrini and the late A. Vigna Taglianti;
they also wish to thank S. Santamaria for the photographs with DIC
optics. V.V. Sarma would like to thank the Ministry of Earth Sciences
(MOES), Govt. of India (MOES/36/OOIS/Extra/40/2014/PC-IV) for
funding this work. He also would like to thank the Tamil Nadu Forest Department and District Forest Office, Tiruvarur, Tamil Nadu for
providing permission to collect samples. Department of Biotechnology, Pondicherry University is thanked for providing the facilities.
B. Devadatha would like to thank the Ministry of Earth Sciences,
Govt. of India for providing a fellowship. V.V. Sarma would like to
thank SERB, Department of Science and Technology, Government of
India, for funding a project (SERB/SB/SO/PS/18/2014 dt.19.5.2015)
the Department of Biotechnology, Pondicherry University for facilities; forest departments of Andaman and Nicobar Islands and Tamil
Nadu, India are thanked for providing permission to collect samples.
M Niranjan thanks SERB, Govt. of India for a fellowship. Archana
Singh is thankful to the University Grants Commission (UGC) (grant
no. F.15-1/2016-17/PDFWM-2015-17-UTT-32206 (SA-II), New Delhi,
India for financial support. The author also thanks the Head, CAS in
Botany, Banaras Hindu University, Varanasi for providing laboratory
facilities; Central Instrumental facility (CIF), Indian Institute of Technology (IIT), Banaras Hindu University, Varanasi for SEM analysis and
Curator, NFCCI, Agharkar Research Institute, Pune for herbarium and
culture deposition and providing accession number. Thatsanee Luangharn thanks Thailand Science Research and Innovation (TSRI) for the
Fundamental Fund “Taxonomy, phylogeny, screening of biologically
active secondary metabolite and cultivation of Ganoderma species”
(Grant No. 652A16009) and Mae Fah Luang University grant “Testing
for optimal condition and cultivation Phallus indusiatus in northern,
Thailand” (Grant No. 651A16028). Naritsada Thongklang thanks Thailand Science Research and Innovation (TSRI) grant entitled “Valueadded products from wild Thai Auricularia to use as a new nutraceutical” (Grant No. 652A01007). Ausana Mapook thanks the Mae Fah
Luang University Fund (Grant No. 651A16029), entitled “Taxonomy,
phylogeny, risk assessment, and potential impact of fungi on Siam
weed in northern Thailand”.The work of Timur S. Bulgakov was done
within the framework of the State Task of the Federal State Budgetary
Institution “Federal Research Center” “Subtropical Scientific Center
of the Russian Academy of Sciences”, the research theme No FGRW2022-0006. Putarak Chomnunti thanks the National Research Council
of Thailand (NRCT) grant “A polyphasic taxonomy and bioactive compound of fungi on Musa spp.” (Grant No. N41A640165) for funding
this research and thanks Reinventing University 2021 for supporting
the research assistant. Hai-Sheng Yuan thanks the National Natural
Science Foundation of China (Grant Nos. 31970017 and U2102220)
for financial support. Saisamorn Lumyong also thanks Chiang Mai
University, Thailand for financial and facility support. Authors thank
13
Fungal Diversity (2022) 117:1–272
Mae Fah Luang University grant entitled “Taxonomy, phylogeny of
micro and macro fungi in Mae Fah Luang University premises” (Grant
No. 6431600). Huang Zhang would like to thank the National Natural
Science Foundation of China (Project ID: NSF 31500017).
Funding Funding was provided by the National Research Council of
Thailand (Grant Nos. NRCT-TRG010, N42A650547, N41A640165),
Ministry of Science and Technology of Thailand (Grant Nos.
652A01003, DBG6280009, 652A01001, 652A16009, 662A01014),
Mae Fah Luang University (Grant Nos. 651B01010, 651A16029,
651A16028, 6431600), Ministry of Higher Education, Science,
Research and Innovation, Thailand, Reinventing University System
Project (652A16049), Thailand Research Grant (DBG6180015),
European Community Research Infrastructure Action (Grant No.
ES-TAF-6618), Ministry of Science and Technology, Taiwan (Grant
No. MOST 107-2621-B178- 002-MY3), Science and Technology
Program of Guizhou Province (Grant No. 20192451-2), Ministry of
Innovative Development of the Republic of Uzbekistan (Grant Nos. AL
2021090820, MUK-2021-46), Ministry of Earth Sciences (Grant No.
MOES/36/OOIS/Extra/40/2014/ PC-IV), University Grants Commission (Grant No. F.15-1/2016-17/PDFWM-2015-17-UTT-32206), Ministry of Higher Education, Science, Research and Innovation, Thailand
(Grant No. 652A16049), Coordenação de Aperfeiçoamento de Pessoal
de Nível Superior - Brasil (CAPES - Finance Code 001), SYNTHESYS
Project http://www.synthesys.info/, financed by European Community
Research Infrastructure Action under the FP7 “Capacities” Program
at the Royal Botanic Garden, Madrid (Grant no. ES-TAF-6618), DST,
SERB-EMR (EMR/2016/003078), SEED-DST (SP/YO/007/2008),
Ministry of Earth Sciences (MOES), Govt. of India (MOES/36/
OOIS/Extra/40/2014/PC-IV) Department of Science and Technology, Government of India (SERB/SB/SO/PS/18/2014 dt.19.5.2015),
University Grants Commission (UGC) (grant no. F.15-1/2016-17/
PDFWM-2015-17-UTT-32206 (SA-II), Scientific Center of the Russian Academy of Sciences", the research theme No FGRW-2022-0006,
Key Laboratory of Green Prevention and Control on Fruits and Vegetables in South China, Guangdong (KA21031C502), Zhongkai University of Agriculture and Engineering, Guangzhou, Guangdong, China
(KA22016B746), National Natural Science Foundation of China (grant
nos. 31970017, 31500017 and U2102220).
Declarations
Conflict of interest Authors declare that they have no conflict of interest.
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Authors and Affiliations
Ruvishika S. Jayawardena1,2 · Kevin D. Hyde1,2,3 · Song Wang1 · Ya‑Ru Sun1,2,4 · Nakarin Suwannarach5,6 ·
Phongeun Sysouphanthong1,2,7 · Mohamed A. Abdel‑Wahab8 · Faten A. Abdel‑Aziz8 ·
Pranami D. Abeywickrama1,2,9 · Vanessa P. Abreu10 · Alireza Armand1,2 · André Aptroot11 · Dan‑Feng Bao1,12,13 ·
Dominik Begerow14 · Jean‑Michel Bellanger15 · Jadson D. P. Bezerra16 · Digvijayini Bundhun1,2 · Mark S. Calabon1,17 ·
Ting Cao18,19 · Taimy Cantillo20 · João L. V. R. Carvalho21 · Napalai Chaiwan1,2 · Che‑Chih Chen22 ·
Régis Courtecuisse23 · Bao‑Kai Cui24 · Ulrike Damm25 · Cvetomir M. Denchev26,27 · Teodor T. Denchev26,27 ·
Chun Y. Deng28 · Bandarupalli Devadatha29,46 · Nimali I. de Silva5,6 · Lidiane A. dos Santos30 · Nawal K. Dubey31 ·
13
Fungal Diversity (2022) 117:1–272
271
Sylvain Dumez23 · Himashi S. Ferdinandez32 · André L. Firmino33 · Yusufjon Gafforov34,35 · Achala J. Gajanayake1,2 ·
Deecksha Gomdola1,2 · Sugantha Gunaseelan36 · Shucheng‑He1,2,37 · Zin H. Htet1,2 · Malarvizhi Kaliyaperumal36 ·
Martin Kemler14 · Kezhocuyi Kezo36 · Nuwan D. Kularathnage1,2,3,61 · Marco Leonardi38 · Ji‑Peng Li28 ·
Chunfang Liao1,2,3 · Shun Liu24 · Michael Loizides40 · Thatsanee Luangharn1 · Jian Ma1,2,41 · Hugo Madrid42 ·
S. Mahadevakumar43,62 · Sajeewa S. N. Maharachchikumbura44 · Dimuthu S. Manamgoda32 · María P. Martín45 ·
Niranjan Mekala46,47 · Pierre‑Arthur Moreau23 · Yan‑Hong Mu18,19 · Pasouvang Pahoua1 · Dhandevi Pem1,2 ·
Olinto L. Pereira10 · Wiphawanee Phonrob48 · Chayanard Phukhamsakda1,39 · Mubashar Raza49 ·
Guang‑Cong Ren1,2 · Andrea C. Rinaldi50 · Walter Rossi38 · Binu C. Samarakoon1,2 · Milan C. Samarakoon13 ·
Vemuri V. Sarma51 · Indunil C. Senanayake3,61 · Archana Singh31 · Maria F. Souza11 · Cristina M. Souza‑Motta21 ·
Adriano A. Spielmann11 · Wenxin Su39 · Xia Tang1,2,63 · XingGuo Tian1,2,41,52 · Kasun M. Thambugala53 ·
Naritsada Thongklang1,2 · Danushka S. Tennakoon5,6 · Nopparat Wannathes48 · DingPeng Wei1,13,37 ·
Stéphane Welti23 · Subodini N. Wijesinghe1,2 · Hongde Yang1,2,37 · Yunhui Yang1,2,3 · Hai‑Sheng Yuan18 ·
Huang Zhang37 · Jingyi Zhang1,2,41 · Abhaya Balasuriya1 · Chitrabhanu S. Bhunjun1,2 · Timur S. Bulgakov54 ·
Lei Cai49 · Erio Camporesi55,56,57 · Putarak Chomnunti2 · Y. S. Deepika58 · Mingkwan Doilom3 · Wei‑Jun Duan59,64 ·
Shi‑Ling Han49 · Naruemon Huanraluek1 · E. B. Gareth Jones60 · N. Lakshmidevi58 · Yu Li39 · Saisamorn Lumyong5,6 ·
Zong‑Long Luo41 · Surapong Khuna5,6 · Jaturong Kumla5,6 · Ishara S. Manawasinghe3 · Ausana Mapook1 ·
Wilawan Punyaboon1 · Saowaluck Tibpromma52 · Yong‑Zhong Lu41 · JiYe Yan9 · Yong Wang4
* Kevin D. Hyde
kdhyde3@gmail.com
14
Institute of Plant Science and Microbiology, Universität
Hamburg, Organismic Botany and Mycology,
Ohnhorststr. 18, 22609 Hamburg, Germany
1
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
15
CEFE, CNRS, Univ. Montpellier, EPHE, IRD, INSERM,
1919, Route de Mende, 34293 Montpellier Cedex 5, France
2
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
16
3
Innovative Institute for Plant Health, Zhongkai University
of Agriculture and Engineering, Guangzhou 510225,
P.R. China
Setor de Micologia, Departamento de Biociências e
Tecnologia, Instituto de Patologia Tropical e Saúde Pública,
Universidade Federal de Goiás, Rua 235, S/N, Setor
Universitário, Goiânia, GO CEP: 74605-050, Brazil
17
Division of Biological Sciences, College of Arts
and Sciences, University of the Philippines Visayas,
5023 Miagao, Iloilo, Philippines
18
CAS Key Laboratory of Forest Ecology and Management,
Institute of Applied Ecology, Chinese Academy of Sciences,
Shenyang 110164, China
19
University of the Chinese Academy of Sciences,
Beijing 100049, China
20
Departamento de Ciências Biológicas, Universidade Estadual
de Feira de Santana, Av. Transnordestina, S/N – Novo
Horizonte, Feira de Santana, BA 44036-900, Brazil
21
Departamento de Micologia Prof. Chaves Batista,
Universidade Federal de Pernambuco, Av. Prof. Moraes
Rego, S/N, Centro de Biociências, Cidade Universitária,
Recife, PE CEP: 50670-901, Brazil
22
Biodiversity Research Center, Academia Sinica, 128
Academia Road, Sec. 2, Nankang 11529, Taipei, Taiwan
23
Faculty of Pharmacy of Lille, EA 4515 (LGCgE), Univ Lille,
59000 Lille, France
24
Institute of Microbiology, School of Ecology and Nature
Conservation, Beijing Forestry University, Beijing 100083,
China
25
Senckenberg Museum of Natural History Görlitz, PF 300
154, 02806 Görlitz, Germany
26
Institute of Biodiversity and Ecosystem Research, Bulgarian
Academy of Sciences, 2 Gagarin St., 1113 Sofia, Bulgaria
4
Department of Plant Pathology, College of Agriculture,
Guizhou University, Guiyang 550025, Guizhou, China
5
Research Center of Microbial Diversity and Sustainable
Utilization, Chiang Mai University, Chiang Mai 50200,
Thailand
6
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
7
Biotechnology and Ecology Institute, Ministry of Agriculture
and Forestry, P.O.Box: 811, Vientiane Capital, Lao PDR
8
Department of Botany and Microbiology, Faculty of Science,
Sohag University, Sohag 82524, Egypt
9
10
11
Beijing Key Laboratory of Environment-Friendly
Management on Fruit Diseases and Pests in North China,
Institute of Plant Protection, Beijing Academy of Agriculture
and Forestry Sciences, Beijing 100097, China
Departamento de Fitopatologia, Universidade Federal de
Viçosa, Viçosa, Minas Gerais, Brazil
Laboratório de Botânica/Liquenologia, Instituto de
Biociências, Universidade Federal de Mato Grosso
do Sul, Avenida Costa e Silva S/N, Bairro Universitário,
Campo Grande, Mato Grosso do Sul CEP 79070-900, Brazil
12
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, Yunnan, China
13
Department of Entomology and Plant Pathology, Faculty
of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
13
272
Fungal Diversity (2022) 117:1–272
27
IUCN SSC Rusts and Smuts Specialist Group, Sofia,
Bulgaria
28
Guizhou Institute of Biology, Guizhou Academy of Sciences,
Shanxi Road No. 1, Yunyan District, Guiyang 550001, China
29
30
31
Virus Diagnostic and Research Lab, Sri
Venkateswara Institute of Medical Sciences, Tirupati,
Andhra Pradesh 517501, India
Departamento de Micologia, Universidade Federal de
Pernambuco, Recife, Pernambuco, Brazil
Center of Advanced Study in Botany, Institute of Sciences,
Banaras Hindu University, Varanasi, Uttar Pradesh 221005,
India
48
Microbiology Program, Faculty of Science and Technology,
Pibulsongkram Rajabhat University, Phitsanulok 65000,
Thailand
49
State Key Laboratory of Mycology, Institute
of Microbiology, Chinese Academy of Sciences, No. 3, 1st
Beichen West Rd., Chaoyang District, Beijing 100101, China
50
Department of Biomedical Sciences, University of Cagliari,
Cittadella Universitaria, 09042 Monserrato, Italy
51
Department of Biotechnology, School of Life
Sciences, Pondicherry University, R.V. Nagar, Kalapet,
Pondicherry 605014, India
52
Center for Yunnan Plateau Biological Resources Protection
and Utilization, Yunnan Engineering Research Center
of Fruit Wine, College of Biological Resource and Food
Engineering, Qujing Normal University, Qujing 655011,
Yunnan, China
32
Department of Botany, Faculty of Applied Sciences,
University of Sri Jayewardenepura, Nugegoda, Sri Lanka
33
Universidade Federal de Uberlândia, Instituto de Ciências
Agrárias, Monte Carmelo, Minas Gerais, Brazil
53
34
Laboratory of Mycology, Institute of Botany, Academy
of Sciences of Republic of Uzbekistan, 32 Durmon Yuli
Street, Tashkent, Uzbekistan 100125
Generics and Molecular Biology Unit, Faculty
of Applied Sciences, University of Sri Jayewardenepura,
Gangodawila 10250, Nugegoda, Sri Lanka
54
35
AKFA University, 264 Milliy Bog Street, Tashkent,
Uzbekistan 111221
36
Centre for Advanced Studies in Botany, University
of Madras, Guindy Campus, Chennai 600025, India
Department of Plant Protection, Federal Research Centre
the Subtropical Scientific Centre of the Russian Academy
of Sciences, Jana Fabriciusa Str. 2/28, Krasnodar Region,
Sochi, Russia 354002
55
A.M.B, Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314,
25121 Brescia, Italy
56
A.M.B. Gruppo, Micologico Forlivese ‘‘Antonio Cicognani’’,
via Roma 18, 47121 Forlì, Italy
57
Società per gli Studi Naturalistici Della Romagna, C.P. 143,
48012 Bagnacavallo, RA, Italy
37
Shandong Provincial Key Laboratory of Water and Soil
Conservation and Environmental Protection, College
of Resources and Environment, Linyi University, West Side
of North Section of Industrial Avenue, Linyi 276000, China
38
University of L’Aquila Dept. MeSVA, sect. Environmental
Sciences via Vetoio, 67100 Coppito, AQ, Italy
58
39
Internationally Cooperative Research Center of China
for New Germplasm Breeding of Edible Mushroom, Jilin
Agricultural University 38, Changchun 130118, China
Department of Studies in Botany, University of Mysore,
Manasagangotri, Mysuru, Karnataka 570006, India
59
40
Ningbo Academy of Inspection and Quarantine, Ningbo,
Zhejiang 315012, PR China
Limassol, Cyprus
60
41
School of Food and Pharmaceutical Engineering, Guizhou
Institute of Technology, Guiyang 550003, China
Department of Botany and Microbiology, College of Science,
King Saud University, P.O. Box 2455, Riyadh 11451,
Saudi Arabia
42
Departamento de Tecnología Médica, Facultad de Ciencias
de la Salud, Universidad de Tarapacá, Sede Iquique, Av. Luis
Emilio Recabarren, 2477 Iquique, Chile
61
43
Forest Pathology Department, KSCSTE-Kerala Forest
Research Institute, Peechi, Thrissur, Kerala 680653, India
Key Laboratory of Green Prevention and Control on Fruits
and Vegetables in South China, Ministry of Agriculture
and Rural Affairs, Zhongkai University of Agriculture
and Engineering, Guangdong 510225, China
62
School of Life Science and Technology, University
of Electronic Science and Technology of China,
Chengdu 611731, China
Botanical Survey of India, Andaman and Nicobar Regional
Centre, Haddo, Port Blair, South Andaman 744102, India
63
Engineering and Research Center for Southwest
Biopharmaceutical Resource of National Education
Ministry of China, Guizhou University, Guiyang 550025,
Guizhou Province, China
64
Ningbo Customs District, Ningbo 315012, Zhejiang,
PR China
44
45
Real Jardín Botánico, RJB-CSIC, Plaza de Murillo 2,
28014 Madrid, Spain
46
Department of Biotechnology, Pondicherry University,
Kalapet, Pondicheryy 605014, India
47
Department of Botany, Rajiv Gandhi University, Rono Hills,
Doimukh, Papum Pare, Itanagar, Arunachal Pradesh 791112,
India
13