Fungal Diversity (2019) 96:1–242
https://doi.org/10.1007/s13225-019-00429-2
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Fungal diversity notes 1036–1150: taxonomic and phylogenetic
contributions on genera and species of fungal taxa
Kevin D. Hyde1,2,3,4,5,6,71 • Danushka S. Tennakoon1,2,3,6,7 • Rajesh Jeewon8 • D. Jayarama Bhat9,10 •
Sajeewa S. N. Maharachchikumbura11 • Walter Rossi12 • Marco Leonardi12 • Hyang Burm Lee13 •
Hye Yeon Mun14 • Jos Houbraken15 • Thuong T. T. Nguyen13 • Sun Jeong Jeon13 • Jens Christian Frisvad16 •
Dhanushka N. Wanasinghe1,3,4,71 • Robert Lücking17 • André Aptroot18 • Marcela E. S. Cáceres19 •
Samantha C. Karunarathna1,4,5,71 • Sinang Hongsanan3,20 • Rungtiwa Phookamsak1,3,4,5,71 • Nimali I. de Silva1,3,5 •
Kasun M. Thambugala21 • Ruvishika S. Jayawardena3 • Indunil C. Senanayake3,20 • Saranyaphat Boonmee3 •
Jie Chen22 • Zong-Long Luo23 • Chayanard Phukhamsakda2,3 • Olinto L. Pereira24 • Vanessa P. Abreu25 •
André Wilson Campos Rosado24 • Buyck Bart26 • Emile Randrianjohany27 • Valérie Hofstetter28 •
Tatiana B. Gibertoni29 • Adriene Mayra da Silva Soares30 • Helio Longoni Plautz Jr.31 • Helen Maria Pontes Sotão30 •
William Kalhy Silva Xavier32 • Jadson Diogo Pereira Bezerra33 • Thays Gabrielle Lins de Oliveira33 •
Cristina Maria de Souza-Motta33 • Oliane Maria Correia Magalhães33 • Digvijayini Bundhun3,34 •
Dulanjalee Harishchandra2,3,35 • Ishara S. Manawasinghe2,3,35 • Wei Dong3,6,34,36 • Sheng-Nan Zhang3,34 •
Dan-Feng Bao3,23,34 • Milan C. Samarakoon3,5,37 • Dhandevi Pem2,3,6,20 • Anuruddha Karunarathna1,3,7,34 •
Chuan-Gen Lin2,3,6 • Jing Yang2,3,6,37 • Rekhani H. Perera2,3,6,37 • Vinit Kumar3,34 • Shi-Ke Huang1,2,3,6 •
Monika C. Dayarathne1,2,3,6 • Anusha H. Ekanayaka1,2,3 • Subashini C. Jayasiri1,3 • Yuanpin Xiao2,3,6,38 •
Sirinapa Konta1,2,3,6 • Tuula Niskanen39 • Kare Liimatainen39 • Yu-Cheng Dai40 • Xiao-Hong Ji40 •
Xue-Mei Tian41 • Armin Mešić42 • Sanjay K. Singh43 • Kunthida Phutthacharoen2,3,6 • Lei Cai4 • Touny Sorvongxay3 •
Vinodhini Thiyagaraja1,3,6,34 • Chada Norphanphoun2,3,6,7,38 • Napalai Chaiwan1,2,3,6 • Yong-Zhong Lu3,6,38 •
Hong-Bo Jiang1,2,3,6 • Jin-Feng Zhang3,37 • Pranami D. Abeywickrama2,3,35 • Janith V. S. Aluthmuhandiram2,3,35 •
Rashika S. Brahmanage2,3,35 • Ming Zeng1,2,3,6 • Thilini Chethana2,3,35 • Deping Wei1,3,34 • Martina Réblová45 •
Jacques Fournier46 • Jana Nekvindová47 • Renan do Nascimento Barbosa48 • José Ewerton Felinto dos Santos33 •
Neiva Tinti de Oliveira33 • Guo-Jie Li44 • Damien Ertz49,50 • Qiu-Ju Shang2,3,37 • Alan J. L. Phillips51 •
Chang-Hsin Kuo7 • Erio Camporesi52,53,54 • Timur S. Bulgakov55 • Saisamorn Lumyong3,5,68,69 • E. B. Gareth Jones3,56 •
Putarak Chomnunti2,3 • Eleni Gentekaki2,3 • Frank Bungartz57,58,59 • Xiang-Yu Zeng3,38 • Sally Fryar60 •
Zdenko Tkalčec42 • Junmin Liang44 • Guangshuo Li44,61 • Ting-Chi Wen38,62 • Paras Nath Singh43 •
Yusufjon Gafforov63,64,70 • Itthayakorn Promputtha5,72 • Erandi Yasanthika2,3 • Ishani D. Goonasekara1,2,3 •
Rui-Lin Zhao44 • Qi Zhao1 • Paul M. Kirk65 • Jian-Kui Liu37,66 • JiYe Yan35 • Peter E. Mortimer1,71 •
Jianchu Xu1,4,71 • Mingkwan Doilom1,3,4,5,67,71
Received: 28 March 2019 / Accepted: 16 May 2019 / Published online: 24 June 2019
School of Science 2019
Abstract
This article is the tenth series of the Fungal Diversity Notes, where 114 taxa distributed in three phyla, ten classes, 30
orders and 53 families are described and illustrated. Taxa described in the present study include one new family (viz.
Pseudoberkleasmiaceae in Dothideomycetes), five new genera (Caatingomyces, Cryptoschizotrema, Neoacladium, Paramassaria and Trochilispora) and 71 new species, (viz. Acrogenospora thailandica, Amniculicola aquatica, A. guttulata,
Angustimassarina sylvatica, Blackwellomyces lateris, Boubovia gelatinosa, Buellia viridula, Caatingomyces brasiliensis,
Calophoma humuli, Camarosporidiella mori, Canalisporium dehongense, Cantharellus brunneopallidus, C. griseotinctus,
Castanediella meliponae, Coprinopsis psammophila, Cordyceps succavus, Cortinarius minusculus, C. subscotoides,
Diaporthe italiana, D. rumicicola, Diatrypella delonicis,
Dictyocheirospora aquadulcis, D. taiwanense, Digi& Mingkwan Doilom
todesmium chiangmaiense, Distoseptispora dehongensis,
j_hammochi@hotmail.com
D. palmarum, Dothiorella styphnolobii, Ellisembia aurea,
Extended author information available on the last page of the article
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Fungal Diversity (2019) 96:1–242
Falciformispora aquatic, Fomitiporia carpinea, F. lagerstroemiae, Grammothele aurantiaca, G. micropora, Hermatomyces bauhiniae, Jahnula queenslandica, Kamalomyces mangrovei, Lecidella yunnanensis, Micarea squamulosa, Muriphaeosphaeria angustifoliae, Neoacladium indicum, Neodidymelliopsis sambuci, Neosetophoma miscanthi, N. salicis,
Nodulosphaeria aquilegiae, N. thalictri, Paramassaria samaneae, Penicillium circulare, P. geumsanense, P. mali-pumilae,
P. psychrotrophicum, P. wandoense, Phaeoisaria siamensis, Phaeopoacea asparagicola, Phaeosphaeria penniseti, Plectocarpon galapagoense, Porina sorediata, Pseudoberkleasmium chiangmaiense, Pyrenochaetopsis sinensis, Rhizophydium
koreanum, Russula prasina, Sporoschisma chiangraiense, Stigmatomyces chamaemyiae, S. cocksii, S. papei, S. tschirnhausii, S. vikhrevii, Thysanorea uniseptata, Torula breviconidiophora, T. polyseptata, Trochilispora schefflerae and
Vaginatispora palmae). Further, twelve new combinations (viz. Cryptoschizotrema cryptotrema, Prolixandromyces australi, P. elongatus, P. falcatus, P. longispinae, P. microveliae, P. neoalardi, P. polhemorum, P. protuberans, P. pseudoveliae, P. tenuistipitis and P. umbonatus), an epitype is chosen for Cantharellus goossensiae, a reference specimen for
Acrogenospora sphaerocephala and new synonym Prolixandromyces are designated. Twenty-four new records on new
hosts and new geographical distributions are also reported (i.e. Acrostalagmus annulatus, Cantharellus goossensiae,
Coprinopsis villosa, Dothiorella plurivora, Dothiorella rhamni, Dothiorella symphoricarposicola, Dictyocheirospora
rotunda, Fasciatispora arengae, Grammothele brasiliensis, Lasiodiplodia iraniensis, Lembosia xyliae, Morenoina
palmicola, Murispora cicognanii, Neodidymelliopsis farokhinejadii, Neolinocarpon rachidis, Nothophoma quercina,
Peroneutypa scoparia, Pestalotiopsis aggestorum, Pilidium concavum, Plagiostoma salicellum, Protofenestella ulmi,
Sarocladium kiliense, Tetraploa nagasakiensis and Vaginatispora armatispora).
Keywords 71 new taxa Ascomycota Basidiomycota Dothideomycetes Eurotiomycetes Lecanoromycetes
Leotiomycetes Pezizomycetes Phylogeny Taxonomy
Table of contents
Pleosporomycetidae C.L. Schoch, Spatafora, Crous &
Shoemaker
Ascomycota R.H. Whittaker
Pleosporales Luttr. ex M.E. Barr
Amniculicolaceae Y. Zhang ter, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde
1039. Amniculicola aquatica Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov. (contribution by Zong-Long Luo/Dhanushka
Wanasinghe)
1040. Amniculicola guttulata Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov. (contribution by Zong-Long Luo/Dhanushka
Wanasinghe)
1041. Murispora cicognanii Wanas., Camporesi, E.B.G.
Jones & K.D. Hyde, Cryptog. Mycol. 36: 437 (2015), new
record from freshwater habitat (contribution by ZongLong Luo/Dhanushka Wanasinghe)
Arthoniomycetes O.E. Erikss. & Wink
Arthoniomycetidae P.M. Kirk, P. Cannon, Minter &
Stalpers
Arthoniales Henssen ex D. Hawksw. & O.E. Erikss
Lecanographaceae Ertz, Tehler, G. Thor & Frisch (contribution by Damien Ertz/Frank Bungartz)
1036. Plectocarpon galapagoense Ertz & Bungartz, sp. nov
Dothideomycetes O.E. Erikss. & Winka
Dothideomycetidae P.M. Kirk, P.F. Cannon, J.C. David &
Stalpers
Capnodiales Woron
Teratosphaeriaceae Crous & U. Braun
1037. Caatingomyces T.G.L. Oliveira, C.M. Souza-Motta,
O.M.C. Magalhães & J.D.P. Bezerra, gen. nov. (contribution by Jadson Bezerra/Thays Gabrielle Lins de Oliveira/
Cristina Maria de Souza-Motta/Oliane Maria Correia
Magalhães)
1038. Caatingomyces brasiliensis T.G.L. Oliveira, C.M.
Souza-Motta, O.M.C. Magalhães & J.D.P. Bezerra, sp.
nov. (contribution by Jadson Bezerra/Thays Gabrielle Lins
de Oliveira/Cristina Maria de Souza-Motta/Oliane Maria
Correia Magalhães)
123
Amorosiaceae Thambug. & K.D. Hyde
1042. Angustimassarina sylvatica N.I. de Silva, Camporesi & K.D. Hyde, sp. nov. (contribution by Nimali I. de
Silva/Kasun M. Thambugala/Ruvishika S. Jayawardena)
Camarosporidiellaceae Wanas., Wijayaw., Crous & K.D.
Hyde
1043. Camarosporidiella mori Phutthacharoen, T.S. Bulgakov and K.D. Hyde, sp. nov. (contribution by Kunthida
Phutthacharoen/Dhanushka Wanasinghe)
Fungal Diversity (2019) 96:1–242
Cucurbitariaceae G. Winter
1044. Protofenestella ulmi Jaklitsch & Voglmayr, Stud.
Mycol. 90: 111 (2017), new record (contribution by
Digvijayini Bundhun/Dhanushka Wanasinghe/Rajesh
Jeewon)
Dictyosporiaceae Boonmee & K.D. Hyde
1045. Dictyocheirospora aquadulcis Sorvongxay, S.
Boonmee & K.D Hyde, sp. nov. (contribution by Touny
sorvongxay/Saranyaphat Boonmee)
1046. Dictyocheirospora rotunda D’souza, J. Bhat & K.D.
Hyde, Fungal Divers. 80: 465 (2016), new geographical
record (contribution by Shi-Ke Huang/Saranyaphat
Boonmee)
1047. Dictyocheirospora taiwanense Tennakoon, C.H.
Kuo & K.D. Hyde, sp. nov. (contribution by Danushka S.
Tennakoon/Saranyaphat Boonmee)
1048. Digitodesmium chiangmaiense Shang & K.D. Hyde,
sp. nov. (contribution by Qiu-Ju Shang/Saranyaphat
Boonmee)
Didymellaceae Gruyter, Aveskamp & Verkley
1049. Calophoma humuli Thiyagaraja, Bulgakov & K.D.
Hyde, sp. nov. (contribution by Vinodhini Thiyagaraja/
Dhanushka Wanasinghe)
1050. Neodidymelliopsis farokhinejadii Ahmadp. &
Mehr.-Koushk., Sydowia 69:175 (2017), new host record
(contribution by Dulanjalee Harishchandra/Dhanushka
Wanasinghe)
1051. Neodidymelliopsis sambuci Manawas., Camporesi &
K.D. Hyde, sp. nov. (contribution by Ishara Manawasinghe/Dhanushka Wanasinghe)
1052. Nothophoma quercina (Syd. & P. Syd.) Q. Chen &
L. Cai, Stud. Mycol. 82: 213 (2015), new host record
(contribution by Dulanjalee Harishchandra/Dhanushka
Wanasinghe)
Hermatomycetaceae Locq.
1053. Hermatomyces bauhiniae Phukhams., D.J. Bhat &
K.D. Hyde, sp. nov. (contribution by Chayanard
Phukhamsakda/D. Jayarama Bhat)
Lophiostomataceae K. Hiray. & Kaz. Tanaka
1054. Vaginatispora palmae S.N. Zhang, J.K. Liu & K.D.
Hyde, sp. nov. (contribution by Shengnan Zhang/Jian-Kui
Liu/Kasun M. Thambugala)
1055. Vaginatispora armatispora (K.D. Hyde, Vrijmoed,
Chinnaraj & E.B.G. Jones) Wanas., E.B.G. Jones & K.D.
Hyde, Studies in Fungi 1: 62 (2016), new record (contribution by Dan-Feng Bao/Kasun M. Thambugala)
Massariaceae Nitschke
1056. Paramassaria Samarak., & K.D. Hyde, gen. nov.
(contribution by Milan C. Samarakoon/Dhanushka
Wanasinghe)
3
1057. Paramassaria samaneae Samarak & K.D. Hyde, sp.
nov. (contribution by Milan C. Samarakoon/Dhanushka
Wanasinghe)
Phaeosphaeriaceae M.E. Barr
1058. Muriphaeosphaeria angustifoliae D. Pem, Gafforov
& K.D. Hyde, sp. nov. (contribution by Dhandevi Pem/
Rajesh Jeewon)
1059. Neosetophoma miscanthi A. Karunarathna, C.H.
Kuo & K.D. Hyde, sp. nov. (contribution by Anuruddha
Karunarathna/CH Kuo/Rungtiwa Phookamsak)
1060. Neosetophoma salicis Norph., Gafforov, T.C. Wen
& K.D. Hyde, sp. nov. (contribution by Chada Norphanphoun/Y. Gafforov/Sinang Hongsanan)
1061. Nodulosphaeria aquilegiae Chaiwan, Camporesi &
K.D. Hyde, sp. nov. (contribution by Napalai Chaiwan/
Dhanushka Wanasinghe/Ruvishika S. Jayawardena)
1062. Nodulosphaeria thalictri D. Pem, Camporesi & K.D.
Hyde, sp. nov. (contribution by Dhandevi Pem/Rajesh
Jeewon)
1063. Phaeopoacea asparagicola Phukhams., A. Akulov
& K.D. Hyde, sp. nov. (contribution by Chayanard
Phukhamsakda/A. Akulov)
1064. Phaeosphaeria penniseti A. Karunarathna, C.H. Kuo
& K.D. Hyde, sp. nov. (contribution by Anuruddha Karunarathna/C.H Kuo/Rungtiwa Phookamsak)
Pseudoberkleasmiaceae Phukhams. & K.D. Hyde
1065. Pseudoberkleasmiaceae Phukhams. & K.D. Hyde,
fam. nov. (contribution by Chayanard Phukhamsakda)
1066. Pseudoberkleasmium chiangmaiense Y.Z. Lu &
K.D. Hyde, sp. nov. (contribution by Yong-Zhong Lu)
Pyrenochaetopsidaceae Valenzuela-Lopez, Crous, Cano,
Guarro & Stchigel
1067. Pyrenochaetopsis sinensis G.S. Li, J. M. Liang & L.
Cai, sp. nov. (contribution by Lei Cai/Junmin Liang/
Guangshuo Li)
Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray.
1068. Tetraploa nagasakiensis (Kaz. Tanaka & K. Hirayama) Kaz. Tanaka & K. Hirayama, Fungal Divers. 63:
253 (2013), new geographical record (contribution by
Hong-Bo Jiang/Rungtiwa Phookamsak)
Torulaceae Corda
1069. Torula breviconidiophora C.G. Lin & K.D. Hyde,
sp. nov. (contribution by Chuan-Gen Lin/Jian-Kui Liu)
1070. Torula polyseptata C.G. Lin & K.D. Hyde, sp. nov.
(contribution by Chuan-Gen Lin/Jian-Kui Liu)
Trematosphaeriaceae K.D. Hyde, Y. Zhang ter, Suetrong
& E.B.G. Jones
1071. Falciformispora aquatica D.F. Bao, K.D. Hyde &
H.Y. Su, sp. nov. (contribution by Dan-Feng Bao/Zonglong
Liu)
123
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Minutisphaerales Raja, Oberlies, Shearer & A.N. Mill.
Acrogenosporaceae Jayasiri & K.D. Hyde
1072. Acrogenospora sphaerocephala (Berk. & Broome)
M.B. Ellis, Dematiaceous Hyphomycetes: 114 (1971),
reference specimen (contribution by Jing Yang/Subhashini
Chathumini/Jian-Kui Liu)
1073. Acrogenospora thailandica J. Yang & K.D. Hyde,
sp. nov. (contribution by Jing Yang/Subhashini Chathumini/Jian-Kui Liu)
Dothideomycetes orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf. (= Lembosiaceae Hosag.)
1074. Lembosia xyliae X.Y. Zeng, T.C. Wen & K.D.
Hyde, Fungal Divers. (2015), new host record (contribution by Xiang-Yu Zeng/Sinang Hongsanan)
1075. Morenoina palmicola J. Fröhl., K.D. Hyde & Joanne
E. Taylor, Fungal Divers. 3:89 (2000), new host record
(contribution by Sirirnapa Konta/Sinang Hongsanan)
Botryosphaeriales C.L. Schoch, Crous & Shoemaker
Botryosphaeriaceae Theiss. & P. Syd.
1076. Dothiorella plurivora Abdollahz., Javadi & A.J.L.
Phillips, Persoonia 32:9 (2014), new record (contribution
by Shi-Ke Huang/Pranami D. Abeywickrama/Jian-Kui Liu/
A.J.L. Phillips)
1077. Dothiorella rhamni Wanas., Bulgakov, E.B.G. Jones
& K.D. Hyde, Fungal Divers. 78: 253 (2016), new host
record (contribution by Janith Vishwakirthi/Pranami D.
Abeywickrama/Jian-Kui Liu/A.J.L. Phillips)
1078. Dothiorella styphnolobii Brahmanage, Bulgakov &
K.D. Hyde, sp. nov. (contribution by Rashika Brahmanage/
Pranami D. Abeywickrama/Timur S. Bulgakov/Jian-Kui
Liu/A.J.L. Phillips)
1079. Dothiorella symphoricarposicola W.J. Li, J.K. Liu
& K. D. Hyde, Cyptogamie Mycol 35: 265 (2015), new
host record (contribution by Thilini Chethana/Pranami D.
Abeywickrama/Jian-Kui Liu/A.J.L. Phillips)
1080. Lasiodiplodia iraniensis Abdollahz., Zare & A.J.L.
Phillips, Persoonia 25: 8 (2010), new host, geographical
record (contribution by Hansika Perera/Vinith Kumar/JianKui Liu/A.J.L. Phillips)
Jahnulales K.L. Pang, Abdel-Wahab, El-Shar., E.B.G.
Jones & Sivichai
Aliquandostipitaceae Inderb.
1081. Jahnula queenslandica Dayar., Fryar & K.D. Hyde,
sp. nov. (contribution by Monika Dayarathne/Sally Fryar/
Kevin D. Hyde)
Tubeufiales Boonmee, K.D Hyde
Tubeufiaceae M.E. Barr
1082. Kamalomyces mangrovei Dayar., & K.D. Hyde, sp.
nov. (contribution by Monika Dayarathne/Kevin D. Hyde)
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Fungal Diversity (2019) 96:1–242
Eurotiomycetes O.E. Erikss. & Winka
Chaetothyriomycetidae Doweld
Chaetothyriales M.E. Barr
Herpotrichiellaceae Munk
1083. Thysanorea uniseptata N.G. Liu & K.D. Hyde, sp.
nov. (contribution by Ningguo Liu/Jian-Kui Liu)
Eurotiomycetidae Geiser & Lutzoni
Eurotiales G.W. Martin ex Benny & Kimbr.
Aspergillaceae Link (contribution by Hyang Burn Lee,
Thuong T.T. Nguyen, Hye Yeon Mun, Jos Houbraken/Jens
Christian Frisvad/Paul M. Kirk)
1084. Penicillium circulare Hyang B. Lee, P.M. Kirk &
T.T.T. Nguyen, sp. nov.
1085. Penicillium geumsanense Hyang B. Lee, P.M. Kirk
& T.T.T. Nguyen, sp. nov.
1086. Penicillium mali-pumilae Hyang B. Lee, T.T.T.
Nguyen & Houbraken, sp. nov.
1087. Penicillium psychrotrophicum Hyang B. Lee, H.Y.
Mun, J.C. Frisvad & Houbraken, sp. nov.
1088. Penicillium wandoense Hyang B. Lee, P.M. Kirk &
T.T.T. Nguyen, sp. nov.
Laboulbeniomycetes Engl.
Laboulbeniales Lindau
Laboulbeniaceae G. Winter (contribution by Walter
Rossi/Marco Leonardi)
1089. Prolixandromyces R.K. Benjamin 1970, Aliso 7:
174 = Monandromyces R. K. Benjamin 1999, Aliso 18: 72,
syn. nov.
1090. Prolixandromyces australis (R.K. Benjamin) W.
Rossi, comb. nov.
1091. Prolixandromyces elongatus (R.K. Benjamin) W.
Rossi, comb. nov.
1092. Prolixandromyces falcatus (R.K. Benjamin) W.
Rossi, comb. nov.
1093. Prolixandromyces longispinae (R.K. Benjamin) W.
Rossi, comb. nov.
1094. Prolixandromyces microveliae (Thaxter) W. Rossi,
comb. nov.
1095. Prolixandromyces neoalardi (R.K. Benjamin) W.
Rossi, comb. nov.
1096. Prolixandromyces polhemorum (R.K. Benjamin)
W. Rossi, comb. nov.
1197. Prolixandromyces protuberans (R.K. Benjamin) W.
Rossi, comb. nov.
1198. Prolixandromyces pseudoveliae (R.K. Benjamin)
W. Rossi, comb. nov.
1199. Prolixandromyces tenuistipitis (R.K. Benjamin) W.
Rossi, comb. nov.
1100. Prolixandromyces umbonatus (R.K. Benjamin) W.
Rossi, comb. nov.
Fungal Diversity (2019) 96:1–242
1101. Stigmatomyces chamaemyiae W. Rossi & M. Leonardi sp. nov.
1102. Stigmatomyces cocksii W. Rossi & M. Leonardi, sp.
nov.
1103. Stigmatomyces papei W. Rossi & M. Leonardi, sp.
nov.
1104. Stigmatomyces tschirnhausii W. Rossi & M. Leonardi, sp. nov.
1105. Stigmatomyces vikhrevii W. Rossi & M. Leonardi,
sp. nov.
Lecanoromycetes O.E. Erikss. & Winka
Lecanoromycetidae P.M. Kirk et al. ex Miadl.
Caliciales Bessey
Caliciaceae Chevall.
1106. Buellia viridula Ekanayaka, & K.D. Hyde, sp. nov.
(contribution by Hasini Ekanayaka)
Lecanorales Nannf.
Lecanoraceae Körb.
1107. Lecidella yunnanensis Ekanayaka, & K.D. Hyde, sp.
nov. (contribution by Hasini Ekanayaka)
Pilocarpaceae Zahlbr.
1108. Micarea squamulosa Aptroot, Lücking & M.
Cáceres, sp. nov. (contribution by André Aptroot/Robert
Lücking/Marcela E. S. Cáceres)
Ostropomycetidae V. Reeb, Lutzoni & Cl. Roux
Ostropales Nannf.
Porinaceae Rchb.
1109. Porina sorediata Aptroot, Lücking & M. Cáceres,
sp. nov. (contribution by André Aptroot/Robert Lücking/
Marcela E.S. Cáceres)
Graphidaceae Dumort.
1110. Cryptoschizotrema Aptroot, Lücking & M. Cáceres,
gen. nov. (contribution by André Aptroot/Robert Lücking/
Marcela E.S. Cáceres)
1111. Cryptoschizotrema cryptotrema (Nyl.) Aptroot,
Lücking & M. Cáceres, comb. nov. (contribution by André
Aptroot/Robert Lücking/Marcela E.S. Cáceres)
Leotiomycetes O.E. Erikss. & Winka
Helotiales Nannf.
Chaetomellaceae Baral, P.R. Johnst. & Rossman
1112. Pilidium concavum (Desm.) Höhn., Sber. Akad.
Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 148 (1915),
new geographical record (contribution by Milan Samarakoon/Jian-Kui Liu)
Pezizomycetes O.E. Erikss. & Winka
Pezizales J. Schröt.
Ascodesmidaceae J. Schröt.
5
1113. Boubovia gelatinosa M. Zeng, Q. Zhao & K.D.
Hyde, sp. nov. (contribution by Ming Zeng/Hasini
Ekanayaka)
Sordariomycetes O.E. Erikss. & Winka
Diaporthomycetidae Senan., Maharachch. & K.D. Hyde
Diaporthales Nannf.
Diaporthaceae Höhn. ex Wehm.
1114. Diaporthe italiana Chethana, Camporesi & K.
D. Hyde, sp. nov. (contribution by Thilini Chethna/Chinthani Senanayaka)
1115. Diaporthe rumicicola Manawas., Camporesi & K.D.
Hyde, sp. nov. (contribution by Ishara Manawasinghe/
Chinthani Senanayaka)
Gnomoniaceae G. Winter
1116. Plagiostoma salicellum (Fr.) Sogonov, Stud. Mycol.
62: 73 (2008), new geographical record (contribution by
Digvijayini
Bundhun/Chinthani
Senanayaka/Rajesh
Jeewon)
Diaporthomycetidae, families incertae sedis
Distoseptisporaceae K.D. Hyde & McKenzie
1117. Distoseptispora dehongensis W. Dong, H. Zhang &
K.D. Hyde, sp. nov. (contribution by Wei Dong/Mingkwan
Doilom)
1118. Distoseptispora palmarum S.N. Zhang, K.D. Hyde
& J.K. Liu, sp. nov. (contribution by Shengnan Zhang/JianKui Liu)
Hypocreomycetidae O.E. Erikss. & Winka
Glomerellales Chadef. ex Réblová, W. Gams & Seifert
Plectosphaerellaceae W. Gams, Summerb. & Zare
1119. Acrostalagmus annulatus (Berk. & M.A. Curtis)
Seifert, Stud. Mycol. 68: 186 (2011), new geographical
record (contribution by Subhashini Chathumini/Hansika
Perera/Ruvishika S. Jayawardena)
Hypocreales Lindau
Cordycipitaceae Kreisel ex G.H. Sung, J.M. Sung, HywelJones & Spatafora
1120. Cordyceps succavus Y.P. Xiao, T.C. Wen & K.D.
Hyde, sp. nov. (contribution by Yuanpin xiao/Ting-Chi
Wen/Sinang Hongsanan)
1121. Blackwellomyces lateris Y.P. Xiao, T.C. Wen &
K.D. Hyde, sp. nov. (contribution by Yuanpin xiao/TingChi Wen/Sinang Hongsanan)
Hypocreales genera incertae sedis
1122. Sarocladium kiliense (Grütz) Summerb., Stud.
Mycol. 68: 158 (2011), new record (contribution by Deping Wei/Dhanushka Wanasinghe)
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
123
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Fungal Diversity (2019) 96:1–242
1123. Phaeoisaria siamensis Jayasiri & K.D. Hyde, sp.
nov. (contribution by Subashini Chathumini)
Savoryellales Boonyuen, Suetrong, Sivichai, K.L. Pang &
E.B.G. Jones
Savoryellaceae Jaklitsch & Réblová
1124. Canalisporium dehongense W. Dong, H. Zhang &
K.D. Hyde, sp. nov. (contribution by Wei Dong/Mingkwan
Doilom)
Sordariomycetidae O.E. Erikss. & Winka
Chaetosphaeriales Huhndorf, A.N. Mill.
Fernández
&
F.A.
Chaetosphaeriaceae Réblová, M.E. Barr & Samuels
1125. Sporoschisma chiangraiense N.G. Liu & K.D.
Hyde, sp. nov. (contribution by Ningguo Liu/Jian-Kui Liu)
1126. Ellisembia aurea Réblová & J. Fourn., sp. nov.
(contribution by Réblová Martina/Jacques Fournier/Jana
Nekvindová)
Linocarpaceae Konta & K.D. Hyde
1127. Neolinocarpon rachidis Konta & K.D. Hyde,
Mycosphere 8: 1968 (2017), new record (contribution by
Sirinapa Konta/Saranyaphat Boonmee)
Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Amphisphaeriaceae G. Winter
1128. Trochilispora V.P. Abreu, A.W.C. Rosado & O.L.
Pereira, gen. nov. (contribution by Olinto Pereira/Abreu
VP/Rosado AWC)
1129. Trochilispora schefflerae V.P. Abreu, A.W.C
Rosado & O.L. Pereira, sp. nov. (contribution by Olinto
Pereira/Abreu VP/Rosado A.W.C)
Sporocadaceae Corda.
1130. Pestalotiopsis aggestorum F. Liu & L. Cai, Nature
Scientific Reports 7: 870 (2017), new geographical record
(contribution
by
Nimali
I.
De
silva/Sajeewa
Maharachchikumbura)
Xylariaceae Tul. & C. Tul.
1134. Fasciatispora arengae Konta & K. D. Hyde,
Mycosphere 9: 725 (2018), new record (contribution by
Sirinapa Konta/Saranyaphat Boonmee)
Basidiomycota R.T. Moore
Agaricomycetes Doweld
Agaricomycetidae Parmasto
Agaricales Underw.
Cortinariaceae R. Heim ex Pouzar (contribution by Tuula
Niskanen/Kare Liimatainen)
1135. Cortinarius minusculus Liimat. & Niskanen, sp.
nov.
1136. Cortinarius subscotoides Niskanen & Liimat., sp.
nov.
Psathyrellaceae Vilgalys et al.
1137. Coprinopsis psammophila Mešić & Tkalčec, sp.
nov. (contribution by Armin Mešić/Zdenko Tkalčec)
1138. Coprinopsis villosa L. Nagy, Desjardin, Vágvölgyi
& Papp, Mycologia 105(1): 120 (2013), new geographical
record (contribution by Armin Mešić/Zdenko Tkalčec)
Cantharellales Gäum.
Botryobasidiaceae Jülich (contribution by Sanjay K.
Singh/Paras Nath Singh)
1139. Neoacladium P.N. Singh & S.K. Singh, gen. nov.
1140. Neoacladium indicum P.N. Singh & S.K. Singh, sp.
nov.
Hydnaceae Chevall (contribution by Bart Buyck/E. Randrianjohany/V. Hofstetter)
1141. Cantharellus goossensiae (Beeli) Heinem., Bull.
Jard. Bot. Etat Brux. 28: 406 (1958), epitypification and
new records
1142. Cantharellus brunneopallidus Buyck, Randrianjohany & V. Hofst., sp. nov.
1143. Cantharellus griseotinctus Buyck, Randrianjohany
& V. Hofst., sp. nov.
Xylariales Nannf.
Castanediellaceae Hern.-Restr., Guarro & Crous
1131. Castanediella meliponae J.E. Felinto-Santos, R.N.
Barbosa & N.T. Oliveira, sp. nov. (contribution by Renan
do Nascimento Barbosa/José Ewerton Felinto dos Santos/
Neiva Tinti de Oliveira)
Hymenochaetales Oberw.
Hymenochaetaceae Donk (contribution by Yu-Cheng Dai/
Xiao-Hong Ji/Xue-Mei Tian)
1144. Fomitiporia carpinea X.H. Ji, X.M. Tian & Y.C.
Dai, sp. nov.
1145. Fomitiporia lagerstroemiae X.H. Ji, X.M. Tian &
Y.C. Dai, sp. nov.
Diatrypaceae Nitschke
1132. Diatrypella delonicis R.H. Perera & K.D. Hyde, sp.
nov. (contribution by Hansika Perera/Monika Dayarathna)
1133. Peroneutypa scoparia Carmarán & A.I. Romero,
Fungal Divers. 23: 84 (2006), new record from mangrove
habitat
(contribution
by Vinith
Kumar/Monika
Dayarathna)
Polyporales Gäum.
Grammotheleaceae Jülich (contribution by Tatiana B.
Gibertoni/A.M.S. Soares/Helio Longoni Plautz Jr/Helen
Maria Pontes Sotão/William Kalhy Silva Xavier)
1146. Grammothele aurantiaca A.M.S. Soares, sp. nov.
1147. Grammothele micropora A.M.S. Soares & W.K.S.
Xavier, sp. nov.
123
Fungal Diversity (2019) 96:1–242
1148. Grammothele brasiliensis Ryvarden, Syn. Fung. 33:
38 (2015), new record
Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C.
David
Russulaceae Lotsy (contribution by Guo-Jie Li/Rui-Lin
Zhao)
1149. Russula prasina G.J. Li & R.L. Zhao, sp. nov.
Chytridiomycota Arx
Rhizophydiomycetes Tedersoo, Koljalg
Rhizophydiales Letcher
Rhizophydiaceae Werderm. (contribution by Hyang Burn
Lee/Sun Jeong Jeon/Thuong T.T. Nguyen)
1150. Rhizophydium koreanum Hyang B. Lee, S.J. Jeon,
T.T.T. Nguyen, sp. nov.
Introduction
Fungi play vital roles in all ecosystems, as decomposers,
epiphytes, endophytes, other symbionts of plants, as well as
animal and plant pathogens (de Silva et al. 2016; Liu et al.
2017; Hyde et al. 2018a). They are hyper-diverse and
heterogeneous group of organisms that represent a large
and diverse component of microbial diversity (Hawksworth
2001; Hernández-Restrepo et al. 2017). Fungi show a great
variation in morphology, reproduction, life cycles and
modes of dispersal (Promputtha et al. 2007; Lofgren et al.
2018). They exhibit different lifestyles, including being
endophytes, biotrophs, hemi-biotrophs, necrotrophs and
saprotrophs (de Silva et al. 2016). Fungal communities are
a major component of soil biomass; they accelerate rock
weathering and decay dead plant material (Kendrick 2000;
Finlay 2008). They are an essential part of the ecosystem
because they play an important role in the decomposition
of organic materials (Soares et al. 2017; Asplund et al.
2018).
The diversity of fungi has been estimated to range
between 2.2 to 3.8 million (Hawksworth and Lücking
2017) however, only 120,000 species are presently known.
Hence, many species are still undescribed. One reason
behind this is due to the fact that many common fungi are
cosmopolitan, having a wider geographical distribution
than plants and other organisms. The fungi of many continents besides Europe have been poorly studied and in
many cases European names have been given to nonEuropean taxa without in depth analysis. Fungal inventories in many countries and in their infancy. This was shown
in a recent study on the fungi of northern Thailand where
up to 96% of species in a genus of mushrooms were found
to be new to science (Hyde et al. 2018b). Despite all
challenges, researchers have established reliable ways to
resolve fungal taxa at the ordinal, familial and generic, as
7
well as species levels (Hyde et al. 2013, 2017a, b, 2018b;
Ariyawansa et al. 2015; Wanasinghe et al. 2018a;
Phookamsak et al. 2019). Major results include 16 phyla
being accepted in the kingdom fungi (Tedersoo et al. 2018;
Wijayawardene et al. 2018a), an outline of the Ascomycota
provided by Wijayawardene et al. (2018a) and an outline of
the basal fungi by Wijayawardene et al. (2018b). In addition, Fungal Diversity notes (Ariyawansa et al. 2015; Hyde
et al. 2016, 2018a; Wanasinghe et al. 2018a; Phookamsak
et al. 2019), Fungal Planet (Crous et al. 2014) and Mycosphere notes series (Hyde et al. 2017b; Thambugala et al.
2017b; Jayawardena et al. 2018) have provided outlets for
introducing the numerous new species of fungi.
More than 1000 entries in Fungal Diversity notes
introducing new taxa, reference specimens, new data, and
other taxonomic contributions have been published with
phylogenetic analyses. This is the tenth in the series of
Fungal Diversity notes with more than 100 entries mainly
collected from China, Italy, Russia, Thailand, plus some
other Asian and European countries. The outcome of the
series provides a stable taxonomy and phylogeny that can
provide definitive classifications for researchers who need
to accurately identify fungi in their research work.
Materials and methods
Materials and methods follow the previous fungal diversity
notes (Hyde et al. 2016; Tibpromma et al. 2017; Wanasinghe et al. 2018a; Phookamsak et al. 2019).
Ascomycota R.H. Whittaker
Notes: We follow the latest treatments and updated
accounts of Ascomycota in Wijayawardene et al.
(2017a, 2018a).
Class Arthoniomycetes O.E. Erikss. & Wink
Arthoniomycetidae P.M. Kirk, P. Cannon, Minter &
Stalpers
Arthoniales Henssen ex D. Hawksw. & O.E. Erikss.
Notes: The order Arthoniales is one of the main lichenized groups of the Ascomycota and belongs to the class
Arthoniomycetes O.E. Erikss. & Winka in the Pezizomycotina O.E. Erikss. & Winka. The order currently includes
the families Andreiomycetaceae B.P. Hodk. & Lendemer,
Arthoniaceae Rchb. ex Rchb., Chrysotrichaceae Zahlbr.,
Lecanographaceae Ertz et al., Opegraphaceae Körb. ex
Stizenb., Roccellaceae Chevall. and Roccellographaceae,
and about 1500 accepted species (Lücking et al. 2017).
Most species of Arthoniales form lichen symbioses, mainly
with trentepohlioid algae, but 285 species are obligately
lichenicolous (non-lichenized) (Diederich et al. 2018) and a
few are considered as being doubtfully or not lichenized.
123
8
Lecanographaceae Ertz et al.
Notes: The family Lecanographaceae was recognized in
Ertz and Tehler (2011) and a formal description was given
in Frisch et al. (2014) based on the type genus Lecanographa Egea & Torrente, along with Alyxoria Ach. ex
Gray, Phacographa Hafellner, Plectocarpon Fée and
Zwackhia Körb. Some Opegrapha species (viz. O. brevis
and O. celtidicola) were placed in this family with
molecular evidence but will have to be combined in
another genus because the type of Opegrapha (O. vulgata)
belongs to the Opegraphaceae.
Plectocarpon Fée
Notes: The genus Plectocarpon Fée was introduced in
1825 as a replacement for Delisea Fée, a later homonym of
Delisea Lamouroux (Rhodophyta). Plectocarpon was first
established for a lichen (now Pseudocyphellaria glabra)
because Fée did not realized that the ascomata were those
of a lichenicolous fungus. The lichenicolous habit was
recognized 159 years later and the name Plectocarpon
resurrected for a genus of lichenicolous fungi (Hawksworth
and Galloway 1984; Galloway 2006). A world monograph
of the genus accepted 32 species, 15 newly described (Ertz
et al. 2005), while a few other species were added in recent
years. The genus was placed in the Lecanographaceae with
molecular evidence using Plectocarpon lichenum (Sommerf.) D. Hawksw. and P. nephromeum (Norman) R. Sant.
(Frisch et al. 2014), but the generic type still needs to be
sequenced.
Plectocarpon galapagoense Ertz & Bungartz, sp. nov.
Index Fungorum number: IF556360; Facesoffungi
number: FoF05977; Fig. 1
Etymology: Referring to the occurrence in the Galapagos
Islands.
Holotype: CDS 33415
Ascomata lichenicolous, infecting and developing inside
the host ascomata and thallus of Sarcographa tricosa sensu
lato, at first immersed, later bursting through the host
ascomata and thallus, black, epruinose, star-shaped
becoming rounded, 1–2 mm diam., not gall-inducing, not
constricted at the base, not producing necrotic areas; surface of lichenicolous ascomata plane to slightly convex,
distinctly lirellate-labyrinthiform with slit-like hymenial
disc, epruinose, typically delimited by remnants of lirellae
and thallus cortex from the host, in particular along the
ascomatal margins of the lichenicolous fungus. Stroma c.
90–130 lm thick; sterile stromatic hyphae dark brown to
carbonized above, 15–40 lm thick, poorly developed
between the loculi, absent below, K–. Hymenium hyaline,
not inspersed with oil droplets, 65–100 lm high; hymenial
gel I ? directly reddish or I ? blue turning reddish,
K/I ? blue; epihymenium hyaline to pale brown, K–,
I ? blue turning reddish with some parts remaining blue.
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Subhymenium hyaline or pale brown, K–, I ? blue with
some parts turning reddish, 10–20 lm high. Paraphysoids
sparingly branched in the hymenium, richly branched in the
epihymenium, 1.5 lm thick, with broadened tips,
2.5–3 lm. Asci subcylindrical to narrowly clavate, with a
narrow ocular chamber, with a tiny K/I ? blue ring in
mature asci, (4–)6(–8)-spored, 55–70 9 11–14 lm. Ascospores fusiform, (2–)3-septate, septation starts with one
median septum, the second cell often slightly larger, hyaline, becoming dark brown granulose when overmature,
(15–)16.5–20.0(–22) 9 (4.5–)4.5–5.5(–6) lm (n = 50), l/b
ratio 3.5–4.0; perispore hyaline, 1–1.5(–2) lm thick.
Conidiomata sphaerical, immersed between the lirellae:
wall dark brown, K–, c. 4–10 lm thick; conidia bacilliform, 4–5 9 1 lm.
Material examined: ECUADOR, Galapagos Islands,
Pinta Island, just below the highest point of the island, 0
350 200 N, 90 450 1300 W, 615 m elev., forest of Zanthoxylum fagara with abundant ferns in the understory,
lichenicolous on Sarcographa tricosa sensu lato growing
on twigs and branches of Chiococca alba, 26 February
2007, F. Bungartz 5759 A (CDS 33415, holotype); F.
Bungartz 5755 A (CDS 33411, paratype); F. Bungartz
5760 A (CDS 33416, paratype).
Distribution and ecology: Described as new from
Galapagos and probably endemic; this lichenicolous fungus
seems to be weakly parasitic, as no necrotic areas are
visible on the hosts ascomata and thallus of Sarcographa
tricosa sensu lato However, as the lichenicolous fungus
frequently occur on the acomata of the host, it should incur
a significant damage and deprive the host of some of its
reproductive structures. The host lichen belongs to S. trigosa sensu lato (identification by Robert Lücking): excipulum very thin, c. 8–25 lm laterally; hymenium not
inspersed, of c. 100–125 lm high; hypothecium hyaline, c.
12–25 lm; ascospores 5-septate, with thick septa, hyaline
becoming greyish brown, c. 20–28 9 7–8 lm.
Notes: In the key to Plectocarpon provided by Ertz et al.
(2005), the new species would key out close to Plectocarpon macaronesiae Diederich, Etayo & Sérusiaux, but
that species is endemic to Macaronesia and differs from P.
galapagoense by smaller ascomata (up to 1.1 mm diam.),
with a shiny rough surface, K ? green intensifying sterile
stromatic hyphae and a different host genus (Lobaria).
Among the recently described Plectocarpon species, P.
dirinariae Ertz & van den Boom is the most similar to P.
galapagoense but it differs by strongly convex and entirely
non-carbonized ascomata and a different host genus
(Dirinaria) (Ertz & van den Boom 2012). Plectocarpon
aequatoriale Etayo, described from Ecuador, has notably
strongly convex ascomata, longer ascospores and a different host genus (Sticta) (Etayo 2017). Opegrapha plectocarpoidea Diederich is lichenicolous on Phaeographis and
Fungal Diversity (2019) 96:1–242
Fig. 1 Plectocarpon galapagoense (Bungartz 5759A, holotype). a–d
Black ascomata emerging from the lirellate ascomata and the thallus
of the host lichen (= Sarcographa tricosa sensu lato, with paler,
distinctly pruinose disc). a, b Overview of the infected host lichen. c,
d Detail of lichenicolous ascomata. c The left part of the lichenicolous
9
ascoma has been removed for microscopical examination. d The
lichenicolous ascoma can be seen on the left, the still intact part of an
ascoma of the host on the right. e Section through an ascoma of
Plectocarpon, in water. f Ascus in KOH. g Ascospores in KOH. Scale
bars: a, b = 2.5 mm, c, d = 500 lm, e = 100 lm, f, g = 10 lm
123
10
reminds of a species of Plectocarpon (Aptroot et al. 1997).
That species, known from high altitude in Papua New
Guinea, differs from Plectocarpon galapagoense by
indistinctly lirellate, almost rounded ascomata with
8-spored asci and a dark brown exciple that continues
below the hymenium.
Class Dothideomycetes O.E. Erikss. & Winka
Notes: Dothideomycetes is the largest class of
Ascomycota. Liu et al. (2017) provided an updated multilocus phylogeny of the class Dothideomycetes with
molecular dating evidence to unravel the evolutionary
relationships and strengthen the ranking among taxa. In this
study, we follow the classifications in the studies of Hyde
et al. (2013), Wijayawardene et al. (2014) and Liu et al.
(2017). The subclasses of Dothideomycetes and their
families in each order are listed in alphabetical order.
Dothideomycetidae P.M. Kirk, P.F. Cannon, J.C. David &
Stalpers
Notes: Dothideomycetidae was introduced by Kirk et al.
(2001) and later validated by Schoch et al. (2006).
According to the present status, Dothideomycetidae comprises the orders Capnodiales, Dothideales and Myriangiales which lack pseudoparaphyses and periphyses (Hyde
et al. 2013).
Capnodiales Woron.
Notes: See Chomnunti et al. (2011) and Hyde et al.
(2013) for details.
Teratosphaeriaceae Crous & U. Braun
Notes: The family Teratosphaeriaceae (Capnodiales,
Dothideomycetes) represents complex cryptic taxa that are
characterized by globular ascomata with brown walls of
textura angularis, located in a stroma of pseudoparenchymatous cells (Crous et al. 2007). Teratosphaeriaceae
comprises 59 quite diverse genera (Wijayawardene et al.
2018a), accommodating endophytic, saprobic, phytopathogenic, and opportunistic species (Crous et al. 2009;
Quaedvlieg et al. 2014). We introduce a new genus
Caatingomyces within Teratosphaeriaceae based on phylogenetic analyses of ITS and LSU rDNA sequence data,
morphology and ecology. In this paper we introduce a new
monotypic genus.
Caatingomyces T.G.L. Oliveira, C.M. Souza-Motta,
O.M.C. Magalhães & J.D.P. Bezerra, gen. nov.
MycoBank number: MB827888; Facesoffungi number:
FoF05818
Etymology: Named after its occurrence in the Brazilian
tropical dry forest, Caatinga.
Endophytic in plants. Sexual morph Undetermined.
Asexual morph Mycelium superficial or immersed, hyaline
to medium brown, septate, branched, hyphae thin-walled,
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Fungal Diversity (2019) 96:1–242
broad and darker when adhered to the conidiomata. Conidiomata pycnidial, superficial or immersed, isolated or
clustered, globose to subovoid, medium brown when young
and dark at maturity, pseudoparenchymatous, glabrose,
thin-walled. Neck absent or poorly developed, conidia
released by irregular rupture. Conidiogenous cells phialidic, slightly tapered and truncated at the apex with robust
base, sometimes branched, hyaline to medium brown,
formed from the internal cells of the pycnidial wall.
Conidia hyaline, aseptate, ellipsoid, obtuse apex with
truncated base, smooth, thin-walled.
Type: Caatingomyces brasiliensis T.G.L. Oliveira, C.M.
Souza-Motta, O.M.C. Magalhães & J.D.P. Bezerra.
Caatingomyces brasiliensis T.G.L. Oliveira, C.M. SouzaMotta, O.M.C. Magalhães & J.D.P. Bezerra, sp. nov.
MycoBank number: MB827889; Facesoffungi number:
FoF05817; Fig. 2
Etymology: Named after its occurrence in Brazil.
Holotype: URM 91831
Endophyte of branches of Poincianella pyramidalis.
Sexual morph Undetermined. Asexual morph Mycelium
superficial or immersed, hyaline to medium brown, septate,
branched, hyphae thin-walled, broad and darker when
linked to the conidiomata. Conidiomata pycnidial (28.6–)
31.2–52 (–67.6) 9 (23.4–) 26–31.2 (– 52) lm, superficial
or immersed in the culture medium, isolated or clustered,
globose to subovoid, medium brown when young and dark
at maturity, pseudoparenchymatous, glabrous, thin-walled.
Rostrate ostiole absent or poorly developed, conidia
released by irregular rupture. Conidiogenous cells (3–)
3.5–5 (–6.5) 9 (3–) 4–5.5 lm, phialidic, slightly tapered
and truncated at apex with robust base, sometimes branched, hyaline to medium brown, arising from the internal
cells of the pycnidial wall. Conidia (3.2–)
4.2–5.3 9 2.1 lm, hyaline, aseptate, ellipsoid, apex obtuse
with truncated base, smooth and thin-walled.
Culture characteristics: Colonies reaching 3 to 3.5 cm in
diameter after 15 days at 25 C in the dark. On OA spreading, erumpent, aerial mycelium sparse with smooth margins;
surface grey-olivaceous to white, reverse black. On PDA
spreading, erumpent, aerial mycelium sparse with smooth
and uneven margins; surface yellowish-brown to pale brown
with presence of yellowish exudate, reverse pale brown
centre to black at the extremities. On MEA spreading, initially creamy, erumpent, surface yellowish-whitish with
smooth margins and presence of exudate, reverse beige.
Material examined: BRAZIL, Paraı́ba State, Santa
Teresinha municipally, Tamanduá farm, 071.524S,
03723.518 W, as endophyte from branches of Poincianella pyramidalis (Fabaceae), May 2013, J.D.P. Bezerra
(URM 91831, holotype), ex-type living culture (URM
7916); as endophyte from branches of Poincianella
Fungal Diversity (2019) 96:1–242
11
Fig. 2 Caatingomyces brasiliensis (URM 91831, holotype).
a Colony on MEA after 7 days. b Colony on PDA after 7 days.
c Colony on AO after 7 days. d Detail of colony on PDA after 7 days.
e Pycnidia. f Detail of pycnidia. g Detail of pycnidial wall. h–l
Conidiogenous cells and conidia. m Conidia. Scale bars: e = 50 lm,
f = 25 lm, g–m = 10 lm
pyramidalis (Fabaceae), May 2013, J.D.P. Bezerra, living
culture (URM 7917).
GenBank numbers: ITS: MH929437, MH929438, LSU:
MH929439, MH929440, RPB2: MH929441, MH929442,
TEF1-a: MH929443, MH929444.
Notes: Maximum likelihood analysis using the alignment of LSU rDNA sequences as presented by Quaedvlieg
et al. (2014), demonstrated relationships of the new genus
with Readeriella Syd. & P. Syd. in Teratosphaeriaceae
(data not shown). In the phylogenetic analyses (Fig. 3)
using combined LSU and ITS sequence data of Teratosphaeriaceae, the new genus Caatingomyces is related to
Readeriella species, but was placed in a distinct clade with
high support. The TEF1-a and RPB2 sequences of C.
brasiliensis were not included in the combined analyses,
but are available in GenBank for further verification of its
relationships and placement in Teratosphaeriaceae and
affiliated families.
Caatingomyces brasiliensis mainly differs from Readeriella species that have a thick conidiomata wall, the
presence of single and circular ostiole, by the shape of the
conidiogenous cells producing three collarettes and deltoid,
thick-walled and guttulate conidia with 3 lateral obtuse
projections (Sutton 1980). Furthermore, Caatingomyces
also differs from species of other genera in Teratosphaeriaceae, such as Parapenidiella Crous & Summerell which has conidia arranged in branched acropetal
chains, pigmented ramoconidia, and erect and macronematous conidiophores (Crous et al. 2012); from Eupenidiella Quaedvl. & Crous by erect and solitary, terminally
penicillate conidiophores and subcylindrical conidia
arranged in branched chains and secondary conidia
(Quaedvlieg et al. 2014).
Subclass Pleosporomycetidae C.L. Schoch et al.
Pleosporales Luttr. ex M.E. Barr
Notes: Pleosporales was established by Luttrell (1955)
to accommodate members of Dothideomycetes having
perithecioid ascomata with pseudoparaphyses amongst the
asci (Zhang et al. 2009a). Pleosporales members are highly
diverse and can be endophytes or epiphytes, parasitices,
lichenicolous or saprobes in terrestrial or aquatic
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Fig. 3 Phylogram generated from Bayesian inference analysis based
on combined LSU and ITS sequence data from 45 representative
members of Teratosphaeriaceae. Related sequences are taken from
Quaedvlieg et al. (2014). Forty-eight strains are included in the
combined gene analyses comprising 1326 characters after alignment
(708 characters for LSU, 618 characters for ITS). Parastagonospora
nodorum (CBS 110109) (Phaeosphaeriaceae, Pleosporales) is used as
the outgroup taxon. Analyses of each single gene were performed and
the topology of each tree had clade stability. The tree topology of the
Bayesian analysis was similar to the maximum likelihood analysis. In
the Bayesian inference analysis presented, the estimated base
frequencies for the LSU region were as follows: A = 0.2445,
C = 0.2356, G = 0.3014, T = 0.2186; substitution rates AC = 0.5614,
AG = 1.2118, AT = 0.8832, CG = 0.6931, CT = 5.0186, GT =
1.0000; range distribution parameter a = 0.6982. The estimated
base frequencies for the ITS region were as follows: A = 0.1855,
C = 0.3132, G = 0.2366, T = 0.2647; substitution rates AC = 2.1196,
AG = 3.1193, AT = 2.6297, CG = 2.0375, CT = 5.3598, GT =
1.0000; range distribution parameter a = 0.4064. The best ranking
RaxML tree with a final probability value was of - 8679.44. The
matrix presented 1063 distinct alignment patterns, with 35.42% of
characters or indeterminate intervals. Branches with later Bayesian
probabilities (BYPP) equal to or greater than 0.95 and bootstrap
support values for maximum likelihood (ML) analysis greater than 70
are placed above or below nodes. Ex-type strains are in bold and
black. The newly generated sequences are indicated in bold and blue
environments or even occur on animal dung (Zhang et al.
2009a). Among the orders of Dothideomycetes, Pleosporales is the largest and most diverse order containing more
than 75 families (Wijayawardene et al. 2018a).
pseudoparaphyses and short-pedicellate asci, bearing hyaline, reddish-brown or pale, 1- to multi-septate or muriform
ascospores, generally with a hyaline gelatinous sheath’’
(Zhang et al. 2008).
Amniculicolaceae Y. Zhang ter, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde
Notes: The family Amniculicolaceae was introduced by
Zhang et al. (2009a) to describe various freshwater taxa
from Europe and later accepted by Shearer et al. (2009)
with a well-supported phylogeny. The family is characterized by ‘‘ascomata with a rough black surface, usually
staining
the
woody
substrate
purple,
narrow
Amniculicola Y. Zhang ter & K.D. Hyde
Notes: Amniculicola was introduced to accommodate A.
lignicola, which was isolated from submerged wood in
freshwater in France (Zhang et al. 2008). This genus is
characterised by ascomata with slit-like ostioles, thin,
branching and anastomosing pseudoparaphyses, cylindrical
asci, and hyaline, 1–3-septate ascospores. Presently, four
species, viz. Amniculicola immersa, A. lignicola, A.
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longissima, A. parva are accepted in Amniculicola and all
species were collected from freshwater habitats.
Amniculicola aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF555506; Facesoffungi
number: FoF05308; Fig. 4
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–1324
Saprobic on decaying wood submerged in freshwater.
Sexual morph Ascomata 260–330 lm high, 250–320 lm
diam., solitary, scattered, or in small groups, superficial,
coriaceous, with basal wall remaining immersed in host
tissue, globose to subglobose, black, often laterally flattened, with a flattened base not easily removed from the
substrate, mostly bearing remnants of wood fibres. Peridium 35–50 lm thick, 2-layered, outer layer composed of
small, dark brown to black, heavily pigmented, thickwalled cells of textura angularis, cells at apex smaller and
walls thicker, inner layer composed of hyaline, thin-walled
cells of textura angularis. Hamathecium of dense, very
long trabeculate pseudoparaphyses. Asci 110–130(–
152) 9 10–13 lm (
x = 120 9 11.5 lm, n = 10), 8spored, bitunicate, fissitunicate, cylindrical, with a short,
narrowed, twisted, bifurcate pedicel. Ascospores 24–
32 9 6–8 lm (
x = 28 9 7 lm, n = 25), obliquely uniseriate and partially overlapping, fusiform, with narrowly
rounded to acute ends, hyaline, uniseptate, deeply constricted at the septum, sometimes curved, smooth, with 4–6
prominent guttules, and a wide gelatinous sheath. Asexual
morph Undetermined.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in Erhai Lake, March 2015,
Z.L. Luo, S-318 (MFLU 18-1324, holotype; HKAS 92579,
isotype), ex-type living culture (MFLUCC 16-1123);
saprobic on decaying wood submerged in Erhai Lake,
March 2015, H.Y. Su, S-467 (MFLU 18-1325, paratype;
HKAS 92559), ex-paratype living culture (MFLUCC
16-0915).
GenBank numbers: LSU: MK106096, TEF1-a:
MK109800 (MFLUCC 16-1123); LSU: MK106097, SSU:
MK106108, TEF1-a: MK109801 (MFLUCC 16-0915).
Notes: Amniculicola aquatica mostly resembles A.
parva in having superficial, coriaceous ascomata of similar
size, cylindrical asci with a short, narrowed, twisted,
bifurcate pedicel and fusiform, hyaline ascospores with a
wide gelatinous sheath. However, A. aquatica differs from
A. parva has a thicker peridium, and uniseptate, wider
ascospores (6–8 vs. 4.5–6.5 lm) (Zhang et al. 2009b).
Phylogenetic analysis based on LSU, SSU and TEF1-a
sequence data showed that A. aquatica is a distinct taxon in
the genus Amniculicola (Fig. 7).
13
Amniculicola guttulata Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF555507; Facesoffungi
number: FoF05309; Fig. 5
Etymology: Referring to the ascospores with large
guttules.
Holotype: MFLU 18–1326
Saprobic on decaying wood submerged in freshwater
habitats. Sexual morph Ascomata 290–320 lm high, 280–
300 lm diam., solitary, scattered, or in small groups,
superficial, coriaceous, with basal wall remaining
immersed in host tissue, globose to subglobose, black,
often laterally flattened, with a flattened base not easily
removed from the substrate, mostly bearing remnants of
wood fibres. Peridium 27–35 lm thick, 2-layered, outer
layer composed of small, brown to dark brown, thickwalled cells of textura angularis, cells at apex smaller and
walls thicker, inner layer composed of hyaline, thin-walled
cells of textura angularis. Hamathecium of dense, very
long trabeculate pseudoparaphyses. Asci 113–127 9 9–
11 lm (
x = 120 9 10 lm, n = 10), 8-spored, bitunicate,
fissitunicate, cylindrical, with a short, narrowed, twisted,
bifurcate
pedicel.
Ascospores
23–27 9 5–7 lm
(
x = 25 9 6 lm, n = 25), obliquely uniseriate and partially overlapping, fusiform, with narrowly rounded to
acute ends, hyaline, uniseptate, deeply constricted at the
septum, sometimes curved, smooth, with 4–6 prominent
guttules, and gelatinous sheath. Asexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in Dulong River, May 2015,
Z.L. Luo, S-428 (MFLU 18-1326, holotype; HKAS 92685,
isotype), ex-type living culture (MFLUCC 16-0907);
saprobic on decaying wood submerged in Dulong River,
May 2015, Z.L. Luo, S-516, living culture MFLUCC
16-1297; saprobic on decaying wood submerged in
Nujiang River, May 2015, Z.L. Luo, S-538 (MFLU
18-1327, paratype), ex-paratype living culture (DLUCC
0538).
GenBank numbers: LSU: MK106098, TEF1-a:
MK109802 (MFLUCC 16-0907); LSU MK106099
(MFLUCC 16-1297); LSU: MK106100, SSU: MK106109,
TEF1-a: MK109803 (DLUCC 0538).
Notes: The new species Amniculicola guttulata resembles A. aquatica in having superficial, coriaceous, globose
to subglobose ascomata, cylindrical asci of similar size and
with a short, narrowed, twisted, bifurcate pedicel, and
fusiform, hyaline, uniseptate ascospores deeply constricted
at the septum. However, A. guttulata differs from A.
aquatica has a thinner peridium and smaller ascospores
(23–27 9 5–7 vs. 24–32 9 6–8 lm). Phylogenetic analysis showed that isolates of A. guttulata formed a separate
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Fig. 4 Amniculicola aquatica (MFLU 18–1324, holotype). a Appearance of ascomata on host substrate. b Section through ascoma. c Structure
of peridium. d Pseudoparaphyses. e, f Asci. g–k Ascospores. Scale bars: b = 150 lm, e, f = 40 lm, c, d = 20 lm, g–k = 15 lm
clade in this genus with high support (98 ML/0.99
Bayesian).
Murispora Y. Zhang bis, J. Fourn. & K.D. Hyde
Notes: The genus Murispora was introduced by Zhang
et al. (2009a) to accommodate Pleospora rubicunda Niessl
which is characterized by immersed, erumpent or nearly
superficial, globose to subglobose, elongate, weakly
123
papillate ascomata that stain the woody substrate purple,
filamentous, narrow, branched, septate, pseudoparaphyses,
8-spored, bitunicate, cylindro-clavate asci, and oval to
ellipsoidal or fusiform, pale or reddish brown, asymmetrical, muriform ascospores, with one side flattened.
Wanasinghe et al. (2015) introduced six species for this
genus which were collected from Italy and the UK. In this
Fungal Diversity (2019) 96:1–242
15
Fig. 5 Amniculicola guttulata (MFLU 18–1326, holotype). a, b Appearance of ascomata on host substrate. c Section through ascoma. d, e
Structure of peridium. f, g Asci. h–l Ascospores. Scale bars: c = 150 lm, f, g = 30 lm, d, e = 20 lm, h–l = 10 lm
paper, we report on Murispora cicognanii as a new record
for China.
Murispora cicognanii Wanasinghe, Camporesi, E.B.G.
Jones & K.D. Hyde, Cryptog. Mycol. 36: 437 (2015)
Facesoffungi number: FoF01108; Fig. 6
Saprobic on decaying wood submerged in freshwater
habitats. Sexual morph Ascomata 200–260 lm high, 160–
250 lm diam., solitary, globose to subglobose, dark brown
to black, semi-immersed to immersed, substrate stained
purple, fused to the host tissue. Peridium comprising 3–4
layers of brown to reddish brown cells of textura angularis,
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with inner 1–2 layers of cells thin-walled and hyaline.
Hamathecium comprising numerous, filamentous, branched, septate, pseudoparaphyses. Asci 120–150 9 20–
25 lm (
x = 135 9 22.5 lm, n = 10), 8-spored, bitunicate,
fissitunicate, cylindric-clavate, short pedicellate, thickwalled at the apex. Ascospores 32–36 9 11–15 lm
(
x = 34 9 13 lm, n = 20), overlapping uniseriate, golden
yellow turning brown when mature, fusiform, asymetrical
with one side flattened, muriform with 5–9 transverse
septa, and 1–2 longitudinal septa, slightly constricted at the
middle septum, conical and narrowly rounded at the ends,
surrounded by a mucilaginous sheath. Asexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in a freshwater stream in
Cangshan Mountain, May 2016, F. Yang, S-757, living
culture (MFLUCC 17–1682).
GenBank numbers: LSU: MK106101, SSU: MK106110,
TEF1-a: MK109804.
Notes: Murispora cicognanii was introduced by
Wanasinghe et al. (2015) based on a collection from Italy
and is only known from the type locality. This species is
characterized by globose to subglobose, immersed ascomata, cylindric-clavate, short pedicellate asci with a minute
ocular chamber and golden yellow, fusiform, asymmetrical, muriform ascospores, turning brown when mature,
with one side flattened and surrounded by a mucilaginous
Fig. 6 Murispora cicognanii (S-757, new record). a Appearance of
ascomata on host substrate. b Section through ascoma. c Structure of
peridium. d Pseudoparaphyses. e Asci. f–i Ascospores. j, k Colonies
on MEA from surface and reverse. Scale bars: b = 100 lm,
e = 50 lm, c, d = 20 lm, f–i = 15 lm
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sheath (Wanasinghe et al. 2015). The morphology of our
new isolate fits well with M. cicognanii. Phylogenetic
analysis of LSU, SSU and TEF1-a sequence data shows
that our isolate clustered with M. cicognanii with good
support (100 ML/1.00 BYPP) (Fig. 7). Based on morphology and phylogeny, we identify our isolate as M.
cicognanii. This is first record for China and the first collection from freshwater.
Amorosiaceae Thambug. & K.D. Hyde
Notes: Thambugala et al. (2015) introduced Amorosiaceae to accommodate the genera Amorosia Mantle & D.
Hawksw and Angustimassarina Thambuga., Kaz. Tanaka
& K.D. Hyde. Amorosiaceae is characterized by immersed
to semiimmersed ascomata with crest-like, papillate ostiole
and hyaline, 1–3-septate ascospores with mucilaginous
sheath and hyphomycetous asexual morph (Thambugala
et al. 2015).
Angustimassarina Thambugala, Kaz. Tanaka & K.D.
Hyde
Notes: Thambugala et al. (2015) introduced Angustimassarina based on the type species Angustimassarina
populi Thambug. & K.D. Hyde. Most Angustimassarina
species are fungicolous and they may be parasitic on other
fungi and appear to grow within other ascomata of other
ascomycetes (Hyde et al. 2017b). Currently, 10 epithets are
listed in Index Fungorum (2019). In this study, we introduce a new Angustimassarina species; A. sylvatica on
Fagus sylvatica from Italy.
Angustimassarina sylvatica N.I. de Silva, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556268; Facesoffungi
number: FoF05992; Fig. 8
Etymology: The specific epithet reflects the host Fagus
sylvatica.
Holotype: MFLU 15-2603
Saprobic on dead twigs of Fagus sylvatica Sexual
morph Ascomata 180–260 lm high, 150–200 lm diam.
(
x = 200 9 180 lm, n = 5), uniloculate, scattered,
immersed to semi-immersed in the plant host tissue, dark
brown to black, globose to subglobose. Ostiole in the
centre without a papilla. Peridium 8–12 lm wide,
unequally thick, comprising brown cells of textura angularis of inner layer and 17–24 lm wide, unequally thick,
comprising irregular dark brown cells of textura prismatica. Hamathecium comprising 1–1.5 lm septate, unbranched, cellular pseudoparaphyses, embedded in gelatinous
matrix, between and above the asci. Asci 95–110 9 8–
12 lm (
x = 98 9 10 lm, n = 15), 8-spored, bitunicate,
fissitunicate, cylindric-clavate, with short pedicel, rounded
at
the
apex.
Ascospores
21–25 9 4–5 lm
(
x = 23 9 4 lm, n = 25), bi-seriate, hyaline, fusiform, 1–
17
septate with 2 pseudosepta, deeply constricted at the septum, widest at the centre and tapering toward the ends,
straight, smooth-walled, guttulate, surrounded by a
mucilaginous sheath. Asexual morph Undetermined.
Material examined: ITALY, Province of Forlı̀-Cesena,
Monte Fumaiolo - Verghereto, on dead land twigs of Fagus
sylvatica (Fagaceae), 5 August 2015, Erio Camporesi,
IT2574 (MFLU 15-2603, holotype), ex-type living culture
(MFLUCC 18-0550).
Culture characteristics: Colonies growing on PDA at
25 C reaching 2 cm in 1 week, irregular, white, fluffy
appearance with undulate edge, reverse white.
GenBank numbers: ITS: MK307843, LSU: MK307844,
SSU: MK314097, TEF1-a: MK360181.
Notes: In our phylogenetic analysis, Angustimassarina
sylvatica groups as a distinct clade with A. alni,
A. premilcurensis and Exosporium stylobatum (Fig. 9).
Angustimassarina sylvatica has longer asci (95–110 lm)
and longer ascospores (21–25 lm) than A. alni (asci
71–89 lm, ascospores 19–22 lm) and A. premilcurensis
(asci 64–93 lm, ascospores 19–23 lm (Tibpromma et al.
2017). Additional details of morphological differences are
mentioned in Table 1. Angustimassarina sylvatica was
isolated from Fagus sylvatica (Sapindaceae) in Italy,
whereas A. alni was recorded from Alnus glutinosa (Betulaceae) in Germany and A. premilcurensis recorded on a
dead branch of Carpinus betulus (Betulaceae) in Italy
(Tibpromma et al. 2017). There are six base pair differences between Angustimassarina sylvatica and A. alni and
four base pair differences between A. sylvatica and
A. premilcurensis for 485 ITS nucleotide sequences. There
is no TEF1-a sequence data available for A. alni and
A. premilcurensis.
Camarosporidiellaceae Wanas., Wijayaw., Crous & K.D.
Hyde
Notes: In an investigation of camarosporium-like taxa,
Wanasinghe et al. (2017a) introduced Camarosporidiellaceae to accommodate the genus Camarosporidiella
which was well-positioned within the suborder Pleosporineae and phylogenetically distinct from other families.
Camarosporidiellaceae species are characterized by
coelomycetous asexual morphs, comprising pycnidial
conidiomata, with a single, papillate ostiole, enteroblastic,
annellidic, integrated to discrete, doliiform, lageniform or
cylindrical, hyaline conidiogenous cells and pale to dark
brown conidia that are phragmosporous to muriform and
mostly ellipsoidal. Their sexual morphs are gregarious to
solitary, globose to subglobose ascomata having a papillate, central ostiole, a peridium containing cell layers of
textura angularis, cylindrical, (2–)4–8-spored asci and
uniseriate, ellipsoidal, brown, muriform ascospores. However, taxonomic classification of Camarosporidiellaceae
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b Fig. 7 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU and TEF1-a sequence data. Seventysix strains are included in the combined analyses which comprised
2640 characters (804 characters for LSU, 995 characters for SSU, 841
characters for TEF1-a) after alignment. Hysterium angustatum (CBS
236.34) and (MFLU 16-1179) (Hysteriaceae, Hysteriales) are used as
the outgroup taxa. Single gene analyses are carried out and the
topology of each tree had clade stability. Tree topology of the
maximum likelihood analysis is similar to the Bayesian analysis. The
best RaxML tree with a final likelihood value of - 19536.000988 is
presented. Estimated base frequencies were as follows:
A = 0.244757, C = 0.246173, G = 0.275709, T = 0.233360; substitution rates AC = 0.971666, AG = 3.423862, AT = 1.366124, CG =
1.101326, CT = 8.907769, GT = 1.000000; gamma distribution
shape parameter a = 0.478984. Bootstrap support values for ML
greater than 75% and Bayesian posterior probabilities greater than
0.95 are given near nodes respectively. The tree is rooted with H.
angustatum (MFLU 16-1179 and CBS 236.34). Ex-type strains are in
bold and black. The newly generated sequences are indicated in blue
species based exclusively on morphological characteristics
is insufficient for designating new species because of their
inadequate phenotypic variability. Mostly taxa are saprobic
and sometimes can be endophytic or pathogenic or potential opportunistic pathogens on leaves and woody materials. The current understanding of ecological and
pathogenic aspects of this group is superficial and extensive sampling should be carried out in different regions and
hosts. Pathogenetic virulence of this family also should be
further investigated with more taxon sampling and DNA
based sequence analyses.
Camarosporidiella Wanas., Wijayaw., K.D. Hyde
Notes: Camarosporidiella was established by Wanasinghe et al. (2017a) with C. caraganicola (Phukhams.,
Bulgakov & K.D. Hyde) Phukhams., Wanas. & K.D. Hyde
as the type species. Wanasinghe et al. (2017a) placed 20
species in this genus based on morphological and multigene phylogenetic anlyses. In a recent study, Hyde et al.
(2018b) added Camarosporidiella populina as a novel
species from Russia. Species of Camarosporidiella are
morphologically less-diverse and distributed throughout a
broad range of environments especially in the Northern
Hemisphere. Camarosporidiella species are saprobes,
endophytes or pathogens of a wide range of hosts
(Wanasinghe et al. 2017a). Species identification based
solely on morphology and plant host association is difficult
since many species have overlapping characters. The genus
is similar to other camarosporium-like genera and distinguishing these genera can be problematic (Wanasinghe
et al. 2017a). LSU is useful for preliminary genus identification. It is recommended to use a combination of LSU,
SSU, ITS and TEF1-a sequence data (Wanasinghe et al.
2017a). In this study, we introduce the 22nd species of this
19
genus, Camarosporidiella mori, which was collected on
Morus alba in Russia.
Camarosporidiella mori Phutthacharoen, Bulgakov,
Wanas. & K.D. Hyde, sp.nov.
Index Fungorum number: IF555709; Facesoffungi
number: FoF0549; Fig. 10
Etymology: The specific epithet ‘‘mori’’ refers to the
plant host genus Morus.
Holotype: MFLU 17–2147
Necrotrophic on branches of Morus alba L. Sexual
morph Undetermined. Asexual morph Conidiomata 370–
520 9 220–430 lm (
x = 428 9 358 lm, n = 10), pycnidial, solitary or gregarious, scattered on wood, immersed to
semi-erumpent, unilocular, dark brown, globose, black in
centre with a papillate ostiole; Pycnidial wall 30–60 lm
(
x = 38.5 lm, n = 10), composed of cells of textura
angularis, multi-layered, with outer layer composed of 4–6
layers of heavily pigmented, light brown cells, inner layer
composed of 3–4 layers, inwardly lighter. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells hyaline, rough, enteroblastic, numerous, clavate, formed from
the inner most layer of the pycnidial wall. Conidia 15.5–
21 9 6.5–9.5 lm (
x = 18 9 8.5 lm, n = 20), hyaline
when immature, brown when mature, oval, with 4-transverse septa and 1-longitudinal septum, smooth and thinwalled, blunt at both ends.
Material examined: RUSSIA, Rostov region, Shakhty
City, trees near buildings, on dying branch of Morus alba
L. (Moraceae), 9 April 2017, Timur S. Bulgakov (MFLU
17–2147, holotype).
GenBank numbers: ITS: MK590359, LSU: MK590358.
Notes: Three species have been found on Morus alba,
viz. Camarosporidiella celtidis, C. moricola and C. mori.
Conidiomata are similar in dimensions and shape. They are
unilocular, globose and black, with papillate ostioles in the
centre. The shape of conidiogenous cells are not significantly differet. The conidia of C. mori are pale brown,
while C. moricola has much darker conidia. Camarosporidiella mori conidia are comparatively larger than
C. moricola (18 9 8.5 lm vs 11 9 5.5 lm). Camarosporidiella moricola and C. mori have conidia with
longitudinal septa, while septa are lacking in C. celtidis. In
the phylogenetic analyses, C. mori groups with C. eufemiaea and C. premilcurensis with high bootstrap support and
is clearly distinct from C. moricola (Fig. 11). Comparison
with the C. eufemiaea and C. premilcurensis asexual morph
characters are currently impossible as they have introduced
only as sexual morphs, while C. mori is known only from
its asexual morph. Therefore, we introduce our novel collection as a new species.
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Fig. 8 Angustimassarina sylvatica (MFLU 15–2603, holotype). a Ascomata on the host material. b, c Sections through ascomata. d Peridium.
e Pseudoparaphyses. f–h Ascospores. i–l Asci. Scale bars: b, c = 80 lm, d, i–l = 50 lm, f–h = 10 lm
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Cucurbitariaceae G. Winter
Notes: The family Cucurbitariaceae, in the order
Pleosporales, was established by Winter (1885) and Cucurbitaria Gray is the type genus (Doilom et al. 2013;
Hyde et al. 2013; Wijayawardene et al. 2014; Li et al.
2016). Species belonging to this family have grouped,
ostiolate ascomata located superficially on their hosts or on
a pseudostromatic structure with cylindrical asci and
phragmosporous or muriform ascospores (Hyde et al.
2013). They are either saprobes or necrotrophs on woody
substrates (Hyde et al. 2013). While Index Fungorum
(2019) lists numerous epithets belonging to Cucurbitariaceae, many of them are not linked to the latter and only a
few species have molecular data in GenBank (Wanasinghe
et al. 2017b; Jaklitsch et al. 2018). The family has been
subjected to several generic additions and exclusions (Barr
1987; de Gruyter et al. 2010, 2013; Hyde et al. 2013;
Doilom et al. 2013). Known asexual morphs of Cucurbitariaceae are pyrenochaeta- or phoma-like (Jaklitsch
et al. 2018). For this reason, Pyrenochaeta De Not. was
accommodated in the family (de Gruyter et al. 2010, 2013;
Chen et al. 2015). Pyrenochaeta quercina Kabát & Bubák,
P. unguis-hominis Punith. & M.P. English, P. cava
(Schulzer) Gruyter, Aveskamp & Verkley, P. hakeae Crous
and P. keratinophila Verkley, C. Ferrer & Gené (Wanasinghe et al. 2017b; Valenzuela-Lopez et al. 2018) were,
however, recently transferred to the novel genus Neocucurbitaria Wanas., E.B.G. Jones & K.D. Hyde introduced
by Wanasinghe et al. (2017b). Valenzuela-Lopez et al.
(2018) introduced another genus, Paracucurbitaria
Valenz.-Lopez, Stchigel, Guarro & Cano, which currently
has two species, Paracucurbitaria corni (Bat. & A.F. Vital)
Valenz.-Lopez, Stchigel, Guarro & Cano., formerly identified as Pyrenochaeta corni (Bat. & A.F. Vital) Boerema,
Loer. & Hamers (Boerema et al. 1996) and a new species,
P. italica Valenz.-Lopez, Crous, Stchigel, Guarro & Cano
to the family. They established the new monotypic genus
Allocucurbitaria Valenz.-Lopez, Stchigel, Guarro & Cano
and the new families Pseudopyrenochaetaceae Valenz.Lopez, Crous, Stchigel, Guarro & Cano and Neopyrenochaetaceae Valenz.-Lopez, Crous, Cano, Guarro &
Stchigel to accommodate taxa previously belonging to the
genus Pyrenochaeta, thereby, eliminating Pyrenochaeta
sensu stricto from Cucurbitariaceae. Additionally, species
of Pyrenochaetopsis Gruyter, Aveskamp & Verkley, also
previously included in Cucurbitariaceae, have been transferred to the new family Pyrenochaetopsidaceae Valenz.Lopez, Crous, Cano, Guarro & Stchigel (Valenzuela-Lopez
et al. 2018). The latest generic additions to the family
Cucurbitariaceae include Astragalicola Jaklitsch & Voglmayr, Cucitella Jaklitsch & Voglmayr, Parafenestella
Jaklitsch & Voglmayr, Protofenestella Jaklitsch &
21
Voglmayr and Seltsamia Jaklitsch & Voglmayr (Jaklitsch
et al. 2018).
Protofenestella Jaklitsch & Voglmayr
Notes: Protofenestella (Cucurbitariaceae, Pleosporales)
was established and defined as the primitive form of the
genus Fenestella Tul. & C. Tul. (Jaklitsch et al. 2018), by
virtue of non-clustered, ill-defined ascomata on host
materials, as well as distant phylogenetic placement. The
genus currently accommodates the single species
Protofenestella ulmi Jaklitsch & Voglmayr, whose sexual
morph was reported on Ulmus sp. from Europe (Jaklitsch
et al. 2018). Its asexual morph was described from culture
(Jaklitsch et al. 2018). In this study, we introduce the
asexual morph of P. ulmi from its host with evidence from
morphology and concatenated phylogenetic analysis based
on ITS, LSU and TEF1-a sequence data.
Protofenestella ulmi Jaklitsch & Voglmayr
Facesoffungi number: FoF05078; Fig. 12
Saprobic on dead twigs and branches of Ulmus pumila
L. Sexual morph See Jaklitsch et al. (2018) for description
and illustrations. Asexual morph Coelomycetous, phomalike. Conidiomata 78–131 lm high, 71–135 lm diam.
(
x = 96 9 96.4 lm, n = 6), globose to depressed subglobose, erumpent through host periderm, solitary or loosely
aggregated in ill-defined groups. Peridium 9–12.8 lm wide
at the top, 5.8–8.9 lm wide at the sides, consisting of 4–6
layers, outer layer comprising olivaceous brown, relatively
thin-walled cells of textura angularis, inner layer becoming
paler and comprising pale brown, thin-walled cells of
textura angularis. Conidiophores reduced to conidiogenous
cells.
Conidiogenous
cells
9–27 9 0.9–2.8 lm
(
x = 14.3 9 1.66 lm, n = 32) hyaline, enteroblastic,
phialidic, lageniform, doliiform to cylindrical, straight to
faintly curved, discrete or integrated. Conidia 2.5–
5.4 9 0.9–2.4 lm (
x = 3.9 9 1.4 lm, n = 76), hyaline,
aseptate, cylindrical or oblong to allantoid, mostly containing 2 small guttules, smooth-walled.
Culture characteristics: Colonies on MEA reaching
1.5 cm diam. after 21 days at 25 C, circular, colony initially greyish olive, eventually turning unevenly brown
after 4 weeks, with limited aerial mycelium, reverse olivaceous dull green to grey, odour indistinct.
Material examined: RUSSIA, Rostov region, Shakhty
City, Cotton fabric microdistrict, artificial grove
(47.72195 N, 40.25177 E), on dead twigs of Ulmus
pumila L. (Ulmaceae), 26 May 2017, Timur S. Bulgakov
(MFLU 17-2047, HKAS 104976), living culture
(MFLUCC 18-0376).
GenBank numbers: ITS: MK418793, LSU: MK418783,
TEF1-a: MK424969.
Notes: The asexual morph of Protofenestella ulmi has so
far only been reported from culture (Jaklitsch et al. 2018).
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Fungal Diversity (2019) 96:1–242
23
b Fig. 9 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, ITS and TEF1-a sequence data.
Related sequences are obtained from GenBank. Melanomma pulvispyrius (CBS 124080) is used as the outgroup taxon. Twenty-nine
strains are included in the combined analyses which comprise 3297
characters (525 characters for ITS, 847 characters for LSU, 984
characters for SSU, 941 characters for TEF1-a) after alignment.
Single gene analyses were carried out and the topology of each tree
had clade stability. Tree topology of the maximum likelihood analysis
is similar to the maximum parsimony analysis. The best RaxML tree
with a final likelihood value of - 12251.741088 is presented. The
matrix had 784 distinct alignment patterns, with 18.70% undetermined characters or gaps. Estimated base frequencies were as follows:
A = 0.244380, C = 0.247658, G = 0.268826, T = 0.239136; substitution rates AC = 1.50180, AG = 2.768806, AT = 1.335846, CG =
1.292688, CT = 6.996209, GT = 1.000000; gamma distribution
shape parameter a = 0.137453. Maximum parsimony analysis of
2585 constant characters and 485 informative characters resulted in
two equally most parsimonious tree of 1370 steps (CI = 0.688,
RI = 0.838, RC = 0.577, HI = 0.312). Bootstrap support values for
ML (first set) and MP equal to or greater than 50% are given above
the nodes. The ex-type strains are in bold and black. The newly
generated sequence is indicated in bold and blue
In the present study, the asexual morph of the species is
illustrated from natural substrate, Ulmus pumila. The strain
obtained from this study clusters with the type strain of P.
ulmi with high booststrap support (100% ML, 100 BYPP)
in Fig. 13. The conidia of P. ulmi observed in our study are
similar in size to those obtained from the ex-holotype
culture
[2.5–5.4 9 0.9–2.4 lm
(this
study)
vs
2.9–5.4 9 1.3–2.2 lm (Jaklitsch et al. 2018)]; slight differences may be accounted for by the fact that the asexual
morph of P. ulmi in this study was observed from natural
substrate while that reported by Jaklitsch et al. (2018) was
obtained from culture.
Dictyosporiaceae Boonmee & K.D. Hyde
Notes: The family Dictyosporiaceae was erected by
Boonmee et al. (2016) with Dictyosporium Corda. as the
type genus with morphological data and multi-gene
analysis. The asexual morphs are cheirosporous hyphomycetes (Boonmee et al. 2016) and the sexual morphs are
characterized by immersed to erumpent or superficial,
globose to subglobose, dark brown to black ascomata,
bitunicate asci and septate, hyaline, sheathed ascospores.
Currently, there are twelve genera in this family (Wijayawardene et al. 2018a).
Dictyocheirospora D’souza et al.
Notes: Dictyocheirospora was introduced by Boonmee
et al. (2016) with D. rotunda D’souza, Bhat & K.D. Hyde
as the type species. Dictyocheirospora species are characterized by dark sporodochial colonies with aeroaquatic
cheiroid dictyospores and all are saprobes (Boonmee et al.
2016). Currently, there are 19 epithets in Index Fungorum
(2019).
Dictyocheirospora aquadulcis Sorvongxay, S. Boonmee &
K.D Hyde, sp. nov.
Index Fungorum number: IF556308; Facesoffungi
number: FoF05963; Fig. 14
Etymology: From the Latin aquadulcis (aqua = water
and dulcis = sweet, in reference to its freshwater habitat.
Holotype: MFLU 18-1088
Saprobic on decaying, submerged wood in freshwater.
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Colonies on natural substrate, superficial,
scattered. Mycelium immersed, branched, septate, pale
brown, smooth-walled. Conidiomata 117–412 lm diam.,
slightly, pulvinate, dark brown. Conidiophores micronematous, undifferentiated from vegetative hyphae. Conidiogenous cells, holoblastic, integrated, terminal, pale
brown.
Conidia
60–80 lm 9 17–29 lm
(
x = 70 9 24 lm, n = 20), solitary, cheiroid, ellipsoid to
cylindrical, rounded at the apex, consisting of 7 rows of
cells, with each row composed of 11–14 cells, with rows
digitate, cylindrical, inwardly curved at apex, arising from
Table 1 Synopsis of Angustimassarina species
Species name
Asci (lm)
Ascospores (lm)
Plant host
Locality
Reference
A. acerina
92–105
21–23
Acer platanoides
Germany
Thambugala et al. (2015)
A. alni
71–89
19–22
Alnus glutinosa
Germany
Tibpromma et al. (2017)
A. arezzoensis
67–95
19–21
Salvia sp.
Italy
Tibpromma et al. (2017)
A. coryli
70–100
20–25
Corylus avellane
Italy
Hyde et al. (2016)
A. italica
78–103
15–22
Ilex aquifolium
Italy
Tibpromma et al. (2017)
A. lonicerae
55–81
19–25
Lonicera sp.
Italy
Tibpromma et al. (2017)
A. populi
80–95
19–22
Populus sp.
Italy
Thambugala et al. (2015)
A. premilcurensis
64–93
19–23
Carpinus betulus
Italy
Tibpromma et al. (2017)
A. quercicola
60–94
17–21
Quercus robur
Germany
Thambugala et al. (2015)
A. rosarum
40–102
16–22
Rosa canina
Italy
Wanasinghe et al. (2018a)
A. sylvatica
95–110
21–25
Fagus sylvatica
Italy
This study
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Fig. 10 Camarosporidiella mori (MFLU 17-2147, holotype). a Herbarium package. b Conidiomata on Morus alba. c Close up of
conidiomata. d Cross section of conidioma. e Section through
conidioma wall. f Conidiogenous cells. g–m Conidia. Scale bars:
c = 1000 lm, d = 100 lm, e = 30 lm, f, g = 10 lm, h–m = 5 lm
a basal cell, euseptate, slightly constricted at septa, brown,
smooth-walled.
Culture characteristics: Conidia germinated on WA
within 24 h. Colonies on MEA reaching 0.2–0.5 cm diam.,
in 5 days at 25 C, with wavy margins, initial white and
later becoming gray.
Material examined: THAILAND, Pha Yao Province, on
decaying wood, in flowing freshwater stream, 11 September 2017, Saranyaphat Boonmee (PK12), (MFLU 18-1088,
holotype), ex-type living culture (MFLUCC 17-2571).
GenBank numbers: ITS: MK634542, LSU: MK634545.
Notes: Dictyocheirospora aquadulcis is similar to D.
heptaspora in having the same conidial shape and size.
However, it differs in having elongate conidia with 11–14
septa, whereas D. heptaspora has ellipsoidal conidia with
10–11 septa. Phylogenetically, D. aquadulcis clustered
with D. heptaspora with moderate bootstrap support
(Fig. 18). We therefore, introduce D. aquadulcis as a new
species in this study based on its morphological
distinctness.
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25
Fig. 11 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU and SSU sequence data for Camarosporidiella species and several closely related genera in
Camarosporidiellaceae. Twenty-seven strains are included in the
combined analyses which comprise 2376 characters (including gaps)
(548 characters for ITS, 850 characters for LSU, 974 characters for
SSU) after alignment. Coniothyrium palmarim (CBS 758.73 and CBS
400.71) (Coniothyriaceae, Pleosporales) are used as the outgroup
taxa. Single gene analyses are carried out and the topology of each
tree had clade stability. Tree topology of the maximum likelihood
analysis was similar to the maximum parsimony analysis and the
Bayesian analysis. The best RaxML tree with a final likelihood value
of - 4091.750594 is presented. The matrix had 130 distinct
alignment patterns, with 5.97% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.252474,
C = 0.213418, G = 0.274008, T = 0.260100; substitution rates AC =
3.818489, AG = 8.876577, AT = 3.746626, CG = 0.477392, CT =
7.229081, GT = 1.000000; gamma distribution shape parameter
a = 0.020000. Maximum parsimony analysis of 2313 constant
characters and 47 informative characters (CI = 0.607, RI = 0.756,
RC = 0.459, HI = 0.393). Bootstrap values for maximum likelihood
and maximum parsimony equal to or greater than 50 and Bayesian
posterior probabilities equal or greater than 0.95 are placed above the
branches respectively. The ex-type strains are in bold and black. The
newly generated sequence is indicated in bold and blue
Dictyocheirospora rotunda M.J. D’souza, Bhat & K.D.
Hyde
Facesoffungi number: FoF05063; Fig. 15
Saprobic on dead wood. Sexual morph Undetermined.
Asexual morph Hyphomycetous. Conidiomata punctiform, sporodochial, velvety, brown to dark brown. Mycelium superficial, with brown, smooth, septate, branched
hyphae. Conidiophores micronematous, pale brown.
Conidiogenous cells 5–8 9 6–9 lm (
x = 6.5 9 7 lm,
n = 20), holoblastic, integrated, terminal, pale brown,
smooth-walled.
Conidia
16–29 9 47–62 lm
(
x = 22 9 55 lm, n = 50), solitary, monoblastic, acrogenous, cheiroid, pale brown to brown, consisting of 6–8
rows of cells, rows digitate, cylindrical, inwardly curved at
the tip, arising from a basal cell, each arm composed of
7–9-septated, constricted at septa.
Culture characteristics: Ascospores germinating on
PDA (potato dextrose agar) within 1 week at 23 C.
Sporulation after 30 days, colony 1.5 cm diam. Initially,
orange from above and reverse, becoming brown, with
concentric zonation, with irregular form of margin, filamentous mycelium, with rough surface and raised
elevation.
Material examined: CHINA, Yunnan Province, Kunming, Songhuaba Lake, on dead wood, 3 September 2017,
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26
Fig. 12 Protofenestella ulmi (MFLU 17-2047, new record). a, b
Appearance of conidiomata on host substrate. c Vertical section
through conidioma. d Peridium. e Conidiogenous cells. f Mature and
immature conidia attached to conidiogenous cells. g–i Conidia. j, k
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Colony on MEA (j from above view, k from below view).
l Conidiomata on colony. m Conidia attached to conidiogenous cell
in culture. Scale bars: a = 1000 lm, b, l = 100 lm, c = 50 lm, d,
e = 20 lm, f = 10 lm, m = 7 lm, g–i = 3 lm
Fungal Diversity (2019) 96:1–242
S.K. Huang (KUN-HKAS 99571), living culture (KUMCC
18-0014).
GenBank numbers: ITS: MK050008, LSU: MK050010,
SSU: MK050009.
Notes: Dictyocheirospora rotunda has been previously
reported from Thailand and northwestern Yunnan Province, China (Boonmee et al. 2016; Wang et al. 2016). In
phylogenetic analysis of combined ITS and LSU sequence
data of Dictyosporiaceae strains, our strain clusters with the
ex-type strain of Dictyocheirospora rotunda (MFLUCC
14-0293) with relatively high-support (Fig. 18). Pairwise
comparison of DNA sequences of ITS and LSU regions
between KUMCC 18-0014 and MFLUCC 14-0293 reveals
they are slightly different. Boonmee et al. (2016) introduced D. rotunda as having 5–7 9 6–7 lm, pale brown
conidiogenous cells and 42–58 9 19–38 lm, cylindrical
conidia. The morphological similarities reveal they are the
same species. Thus, a record of D. rotunda from Kunming,
Yunnan Province is reported.
Dictyocheirospora taiwanense Tennakoon, C.H. Kuo &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556309; Facesoffungi
number: FoF05964; Fig. 16
Etymology: Named after Taiwan, where this fungus was
collected.
Holotype: MFLU 18-0070
Saprobic on decaying wood of Macaranga tanarius (L.)
Müll.Arg. Sexual morph Undetermined. Asexual morph
Hyphomycetous. Conidiomata punctiform, sporodochial,
110–230 lm diam., velvety, dark brown to black. Conidiophores micronematous, pale brown, smooth, thin-walled.
Conidiogenous cells holoblastic, cylindrical. Conidia (72–)
74–84(–86) 9 16–20(–24)
lm
(
x = 77.5 9 18 lm,
n = 30), solitary, brown, ellipsoid to cylindrical, rounded at
the apex, cheiroid, not complanate, consisting of 5 rows of
cells, rows cylindrical, arising from a basal cell, each
composed of 10–13 cells, constricted at septa, without
appendages.
Culture characteristics: Conidia germinating on water
agar within 24 h, germ tubes appearing from the basal cells
of the conidium. Colonies on PDA covering 3 cm diam., in
3 weeks at 27 C, at first white, colony from above: light
orange to yellowish at the margin, yellowish to white at the
centre; reverse, orange to yellowish at the margin, orange
at the centre; mycelium yellowish to white with tufting; not
producing pigments in PDA.
Material examined: TAIWAN, Chiayi, Shihnong Forest
Area, decaying wood of Macaranga tanarius (L.) Müll.Arg. (Euphobiaceae), 25 June 2017, D.S. Tennakoon,
DTW013 (MFLU 18-0070, holotype), ex-type living culture (MFLUCC 17-2654).
GenBank numbers: ITS: MK495821, LSU: MK495820.
27
Notes: Dictyocheirospora taiwanense is typical of Dictyocheirospora in having aeroaquatic cheiroid dictyospores
(Boonmee et al. 2016). Phylogenetically, D. taiwanense
nests independently closely to D. garethjonesii (MFLUCC
16-0909), D. aquatica (KUMCC 15-0305), D. cheirospora
(KUMCC 17-0035) and D. rotunda (MFLUCC 14-0293a,
HKAS 99571) with moderate support (Fig. 18). However,
D. taiwanense differs from D. aquatica and D.
garethjonesii
in
having
larger
conidia
(74–84 9 16–20 lm) with a higher number of cells
(10–13) in each conidial row, whereas the conidia of D.
aquatica (34–42 9 12.5–19.5 lm) and D. garethjonesii
(45.5–54.5 9 15.5–24.5 lm) are comparatively smaller
and comprise 6–8, 7–10 cells in each conidial row (Wang
et al. 2016). In addition, D. pseudomusae and D. vinaya are
also show distinct morphological differences viz. mostly D.
taiwanense conidia has 5 rows (10-13 cells in each row),
whereas D. pseudomusae has 6-7 rows (13–15 cells in each
row) and D. vinaya (6–7 rows, 9–13 cells in each row)
(Tanaka et al. 2015; Boonmee et al. 2016).
Digitodesmium Corda, Weitenweber’s Beitr.
Notes: Digitodesmium was established by Kirk (1981)
with type species Digitodesmium elegans P.M. Kirk. The
asexual morph of the Digitodesmium species is characterized by punctiform, sporodochial conidiomata and
acrogenous, euseptate, cheiroid, digitate conidia, with
apical gelatinous caps. There are six records in Index
Fungorum (2019), however, these are lacking the molecular data with the exception of D. bambusicola.
Digitodesmium chiangmaiense Q.J. Shang & K.D. Hyde
sp. nov.
Indexfungroum number: IF556220; Facesoffungi number: FoF05098; Fig. 17
Etymology: Names reflects the province in Thailand,
from where the holotype was collected.
Holotype KUN-HKAS 102163
Saprobic on decaying wood submerged in stream.
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Colonies punctiform, sporodochial, scattered, dark brown to black, glistening. Conidiophores
micronematous, hyaline to pale brown, unbranched, thinwalled, smooth. Conidiogenous cells monoblastic, integrated, terminal, determinate, hyaline to pale brown,
smooth. Conidia acrogenous, solitary, cheiroid, smoothwalled, complanate, brown to dark brown, consisting of
three arms closely compact with side arms higher than
middle arms, (24–)33–42(–44) 9 (13–)15–18(–21) lm
(
x = 38 9 17 lm, n = 45), 5–7-euseptate in each arm,
without appendages attached at the two outer arms, arms
(20–)27–37(–38) 9 (4–)4–7(–8) lm (
x = 32 9 5 lm,
n = 30), not discrete, unbranched, cylindrical.
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b Fig. 13 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU and TEF1-a sequence data for
Protofenestella species and several closely related genera in Cucurbitariaceae. Related sequences are taken from Wanasinghe et al.
(2017b) and Jaklitsch et al. (2018). Forty-three strains are included in
the combined analyses which comprise 2749 characters (601 characters for ITS, 914 characters for LSU and 1234 characters for TEF1a) after alignment. Pyrenochaetopsis leptospora (CBS 101635)
(Pyrenochaetopsidaceae, Pleosporales) is used as the outgroup taxon.
Single gene analyses are carried out and the topology of each tree has
clade stability. Tree topology of the maximum likelihood analysis is
similar to the Bayesian analysis. The best RaxML tree with a final
likelihood value of - 13415.377769 is presented. The matrix had 741
distinct alignment patterns, with 25.51% undetermined characters or
gaps. Estimated base frequencies were as follows: A = 0.234883,
C = 0.253189, G = 0.270793, T = 0.241136; substitution rates AC =
1.125763, AG = 3.623108, AT = 2.100279, CG = 1.168787, CT =
6.351079, GT = 1.000000; gamma distribution shape parameter
a = 0.142500. Bootstrap values for maximum likelihood equal to or
greater than 60 are placed above and below the branches respectively.
Branches with Bayesian posterior probabilities equal or greater than
0.90 are in bold. The ex-type strains are in bold and black. The newly
generated sequence is indicated in bold and blue
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced from the basal cell.
Colonies on PDA reaching 3–5 mm diam. after 7 days at
room temperature, colonies circular, with fluffy, dense,
white mycelium in the middle and sparse mycelium in the
outer ring on the surface with irregular brown margin; in
reverse, brown in the middle and pale yellow to brown at
the margin.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng District, Mushroom Research Center, on dead
wood, 27 January 2017, Qiuju Shang, M7P4-17 (KUNHKAS 102163, holotype), ex-type living culture
(MFLUCC, ICMP).
GenBank numbers: LSU: MK571766, SSU: MK571775.
Notes: Based on analysis of combined ITS and LSU
sequence data (Fig. 18), Digitodesmium chiangmaiense
(M7P4-17) groups with D. bambusicola (CBS 110279)
with moderate support (57% ML, 52% MP and 0.92
BYPP). Digitodesmium chiangmaiense differs from
D. bambusicola in lacking appendages on the outer of the
arms and having longer conidia (Table 2). In addition,
D. chiangmaiense differs from other Digitodesmium species in the dimension and number of arms (Table 2).
Didymellaceae Gruyter, Aveskamp & Verkley
Notes: The family Didymellaceae was established to
accommodate Didymella, Phoma and phoma-like genera
and is one of the most species rich families in the fungal
kingdom (Chen et al. 2015, 2017), with 31 genera based on
morphology and phylogeny (Chen et al. 2015; Jayasiri
et al. 2017; Thambugala et al. 2017a; Wanasinghe et al.
2018b). Species of this family are distributed in a broad
29
range of environments. Most members are economically
important plant pathogens causing leaf and stem spots and
many are mentioned in quarantine regulations (Thambugala et al. 2017b, 2018; Knight et al. 2018; ValenzuelaLopez et al. 2018). Limited studies have been carried out
on the sexual morphs (Chen et al. 2015; Thambugala et al.
2017a, 2018; Wanasinghe et al. 2018b). The taxonomic
placements of this family were treated based on phylogenetic and morphological analyses (Chen et al. 2015;
Jayasiri et al. 2017; Thambugala et al. 2017a; Wanasinghe
et al. 2018b).
Calophoma Qian Chen & L. Cai
Notes: The genus Calophoma, introduced by Chen et al.
(2015), and is typified by C. clematidina. (Thum.) Q. Chen
& L. Cai and accommodates C. aquilegiicola, C. clematidina, C. clematidis-rectae, C. complanata, C. glaucii and
an insufficiently known species (CBS 186.55) (Chen et al.
2015). The sexual morph has been reported for C. petasitis
(Tibpromma et al. 2017). The asexual morph has been
reported with its unique characteristics such as subglobose,
subcylindrical, ellipsoidal, somewhat obclavate-fusiform
conidia with 0–1 septa and chlamydospores are produced
in one species (Chen et al. 2015, 2017; Tibpromma et al.
2017). Currently there are eight species included in this
genus (Tibpromma et al. 2017). We introduce a novel
species to this genus with support from combined LSU,
ITS, TUB2 and RPB2 sequence data.
Calophoma humuli Thiyagaraja., Bulgakov & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555424; Facesoffungi
number: FoF05077; Fig. 19
Etymology: The specific epithet ‘‘humuli’’ refers to the
host plant genus Humulus.
Holotype: MFLU 17-2205
Saprobic on dead stems of Humulus lupulus L. (Cannabaceae). Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata 190–220 lm high 9 210–
230 lm diam. (
x = 205 9 220 lm, n = 5), pycnidial, subimmersed, solitary, globose, black. Pycnidial wall multilayered, 10–25 lm wide at the base, 10–20 lm wide in
sides, thick, comprising two layers, outer layer heavily
pigmented, thick walled, comprising brown cells of texura
angularis, cells towards the inside lighter, inner layer
composed of hyaline texura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells
phialidic, hyaline, smooth walled, formed from inner layer
of the pycnidium wall. Conidia 3–10 9 3–6 lm
(
x = 6.5 9 3.8 lm, n = 50), oblong, straight, rounded at
the ends, sometimes narrowly rounded at the ends, smooth
and thick-walled, aseptate when immature and becoming
1-septate when mature, initially hyaline, becoming pale
brown at maturity.
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Fig. 14 Dictyocheirospora aquadulcis (MFLU 18–1088, holotype).
a, b Colonies on host surface. c Colony of conidia. d–m Conidium
arms. n Germinating conidium. o, p Culture on MEA at 3 weeks. q, r
Culture on MEA at 7 months. Scale bars: a, b = 200 lm,
c = 100 lm, e–m = 50 lm, d, n = 20 lm
Culture characteristics: Colonies on PDA reaching
40–45 mm diam. after 2 weeks at 20–25 C, dense, circular, slightly raised, surface smooth with even edge, cottony, colony from above: white at the margin, grayish at
the centre; from below: white at the margin, brownish
yellow at the centre; not producing pigmentation on PDA
media.
Material examined: RUSSIA, Rostov region, Krasnosulinsky District, Gornensky protected landscape, edge of
ravine forest (47.86602 N, 40.22959 E), on dead stems of
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31
Fig. 15 Dictyocheirospora rotunda (HKAS 99571, new geographical record). a Material. b Appearance of conidiomata on host. c Conidioma.
d–f Conidia. g Squashed conidium. h–j Arms of conidia. Scale bars: d–g = 20 lm, h–j = 10 lm
Humulus lupulus L. (Cannabaceae), 6 July 2017, Timur S.
Bulgakov (MFLU 17-2205, holotype), ex-type living culture (MFLUCC 18-0101).
GenBank numbers: ITS: MK446317, LSU: MK446318,
RPB2: MK492667, TUB2: MK492666.
Notes: In our multi-gene analyses, Calophoma humuli
forms a monophyletic clade with C. petasitis with strong
support. Calophoma petasitis is reported as the sexual
morph in the genus Calophoma with the characteristics of
subglobose ascomata and smooth-walled, guttulate, hyaline
ascospores. A comparison of LSU, ITS and RPB2 with
these two strains reveals 3/820 (0.36%), 9/441 (2.05%) and
75/797 (9.4%) base pair differences. Even though their
LSU and ITS sequences are quite similar, the RPB2 region
reveals they should be two different species following the
guidelines of Jeewon and Hyde (2016). Therefore, we
introduce our isolate as a new species in Calophoma.
Neodidymelliopsis Qian Chen & L. Cai
Notes: Neodidymelliopsis was introduced by Chen et al.
(2015), with Neodidymelliopsis cannabis (G. Winter) Q.
Chen & L. Cai as the type species (Chen et al. 2015; Hyde
et al. 2016).
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Fig. 16 Dictyocheirospora taiwanense (MFLU 18–0070, holotype).
a Conidiomata on the substrate. b, c Close-up of conidiomata.
d Squash mount of conidia. e, f Squash mount of conidioma with
conidiogenous cells. g–j Conidia. k Germinating conidium. Scale
bars: d = 100 lm, e = 50 lm, f–k = 20 lm
Neodidymelliopsis farokhinejadii S.A. Ahmadpour & M.
Mehrabi-Koushki, Sydowia 69: 175 (2017)
Facesoffungi number: FoF04975; Fig. 20
Saprobic or necrotrophic on dead arial stem of Cirsium
sp. Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata on host 75–105 lm diam.
(
x = 74 lm, n = 10), pycnidial, solitary, scattered, globose
to subglobose, black, semi-immersed to immersed; pycnidial wall pseudoparenchymatous, textura angularis dark
brown to hyline cells. Chlamydospores not observed.
Conidiophores not observed. Conidiogenous cells not
observed. Conidia on host 4.5–7.2 9 2.3–3.6 lm diam.
(
x = 6 9 3.2 lm, n = 40), oblong to ellipsoid, hyaline,
aseptate, smooth-walled; in culture, 4.6–7.5 9 2.4–3.9 lm
diam. (
x = 6 9 3.4 lm, n = 40), ellipsoidal to cylindrical,
sometimes allantoid, hyaline, smooth- and thin-walled,
aseptate, with small polar guttules.
Culture characteristics: Colonies on PDA reach
25–30 mm diam. after 7 days at 25 C, with white aerial
mycelium, surface floccose to wooly, margin regular, pale
greenish olivaceous; reverse dull green.
Material examined: ITALY, Forlı̀-Cesena Province,
near Castrocaro Terme, on dead aerial stem of Cirsium sp.
(Asteraceae), 25 February 2018, Erio Camporesi, living
culture (MFLUCC 18-1569).
Genbank numbers: ITS: MK084580, LSU: MK084581.
Notes: Neodidymelliopsis farokhinejadii was first
reported on dead branches of Eucalyptus sp. and was
subsequently found on Citrus limon, Conocarpus erectus,
Ziziphus sp., Juglans regia and Cupressus sp. (Ahmadpour
et al. 2017) Our new isolate of N. farokhinejadii was collected from Forlı̀-Cesena Province in Italy. This is the first
report of N. farokhinejadii on Cirsium species (Farr and
Rossman 2019). Morphological observations such as spore
characteristics support the inclusion of our taxa into Neodidymelliopsis (Chen et al. 2015). DNA sequence analyses
from the LSU, ITS, TUB2 and RPB2 genes confirms that
our taxon clades together with N. farokhinejadii (Fig. 21).
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Fig. 17 Digitodesmium chiangmaiense (KUN-HKAS 102163, holotype). a, b Colonies on submerged wood. c Squash mount of a
sporodochium. d–g Conidia with conidiophores. h Conidium.
33
i Germinated conidia. j, k Culture characteristics on PDA (j from
above view, k from below view) Scale bars: c = 20 lm, d–i = 10 lm
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Fig. 18 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequences data in Dictyosporiaceae.
Related sequences are taken from Boonmee et al. (2016) and Wang
et al. (2016). Fifty-three strains are included in the combined analyses
which comprise 1405 characters (553 characters for ITS, 852
characters for LSU) after alignment. Letendraea helminthicola (7-3)
(Montagnulaceae, Pleosporales) is used as the outgroup taxon. Single
gene analyses are carried out and the topology of each tree had clade
stability. Tree topology of the maximum likelihood analysis is similar
to the Bayesian analysis and the maximum parsimony analysis. The
best RAxML tree with a final likelihood value of - 8736.227244 is
presented. The matrix had 523 distinct alignment patterns, with
18.97% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.241508, C = 0.237486, G = 0.270256,
T = 0.250751; substitution rates AC = 2.173342, AG = 3.473225,
AT = 3.246966, CG = 0.531302, CT = 36.318285, GT = 1.000000;
gamma distribution shape parameter a = 0.567927. Maximum parsimony analysis of 981 constant characters and 292 informative
characters resulted in two equally most parsimonious trees of 1392
steps (CI = 0.486, RI = 0.744 RC = 0.361, HI = 0.514). Bootstrap
values for maximum likelihood (ML) and maximum parsimony (MP)
equal to or greater than 60 are placed above and below the branches
respectively. Branches with Bayesian posterior probabilities (BYPP)
equal or greater than 0.95 are in bold. The ex-type strains are in bold
and black. The newly generated sequences are indicated in bold and
blue
Neodidymelliopsis sambuci Manawasinghe, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF555378; Facesoffungi
number: FoF04939; Fig. 22
Etymology: Name reflects the host genus.
Holotype: MFLU 18-0177
Saprobic or necrotrophic on dead and dying twigs and
branches of Sambucus nigra. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata on
host 37–82 lm diam. (
x = 58 lm, n = 10), pycnidial,
solitary, scattered, globose, black, semi-immersed to
immersed. Conidiophores not observed. Conidiogenous
cells not observed. Conidia on host 4–7 9 2–3.5 lm diam.
(
x = 6 9 2 lm, n = 40), oblong to ellipsoid, hyaline,
aseptate, smooth-walled; in culture, 5–8 9 1–3.5 lm diam.
(
x = 6 9 2.7 lm, n = 40). Conidial exudates not
observed.Culture characteristics: Colonies on PDA reach
65 mm diam. after 7 days at 25 C, with circular, entire
edge, raised grey olivaceous aerial mycelium, surface
floccose to woolly.
Material examined: ITALY, Province of Ravenna, Santa
Lucia, Faenza, on living branch of Sambucus nigra L.
(Adoxaceae), 5 January 2018, Erio Camporesi (MFLU
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35
Table 2 Conidia of Digitodesmium species discussed in this study
Taxa
D.
bambusicola
D. chiangmaiense
D. elegans
D.
heptasporum
D. intermedium
D.
macrosporum
D. recurvum
Colour
Pale brown
Brown to dark brown
–
Pale brown
Brown to dark
brown
Brown to dark
brown
Pale brown
Dimension (lm)
24–
32.5 9 12.5–
23
(25–)30–45(–
44) 9 (13–
)12–21(–21)
45–
60 9 12–
21
50–
75 9 32.5–
70
39–76 9 25–
35
130–
145 9 19–26
30–
45 9 12.5–
23
Appendages
Yes
No
No
No
No
No
No
Wall
Smooth
Smooth
Smooth
Smooth
Smooth
Smooth
Smooth
Septal pores
Conspicuous
Conspicuous
Inconspicous
Conspicuous
Conspicuous
Conspicuous
Conspicuous
Number of arms
3
3
(2–)3–4(–6)
6–7
3–11
5–8
(2–)4–6(–7)
Number of septa
in each of arm
4–7
5–7
9–12
11–17
7–13
17–19
6–10
Country
Philippines
Thailand
UK
China
Spain
Spain
China
Reference
Cai et al.
(2002)
This study
Kirk. (1981)
Cai et al.
(2003)
Silvera-Simón
et al. (2010)
Silvera-Simón
et al. (2010)
Ho et al.
(1999)
Fig. 19 Calophoma humuli (MFLU 17–2205, holotype). a, b Conidiomata on host. c Section through conidioma. d Section through conidioma
wall. e Conidiogenous cells. f–i Conidia. Scale bars: a, b = 500 lm, c = 100 lm, d = 20 lm, e = 10 lm, f–i = 5 lm
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Fig. 20 Neodidymelliopsis
farokhinejadii (JZB–H380023,
new host record). a, b
Conidiomata on the host tissue.
c Section of the pycnidial wall.
d–h Conidia. i Upper view of
7-day old culture. j Reverse
view of 7-day old culture. Scale
bar: c–h = 10 lm
18-0177, holotype), ex-type living culture (MFLUCC
18-1565).
GenBank numbers: ITS: MH84232, LSU: MH84231,
TUB2: MK049556.
Notes: Morphological characters such as conidial size
and shape (Fig. 22) fit well within the concept of Neodidymelliopsis (Chen et al. 2015). The combined gene
sequence analyses of LSU, ITS, RPB2 and TUB2 genes
confirm that present taxon belongs in genus Neodidymelliopsis in a strongly supported monophyletic lineage
(Fig. 21) within family Didymellaceae. In phylogenetic
analysis Neodidymelliopsis sambuci is in a sister clade with
N. longicolla. Morphologically, N. sambuci can be distinguished from N. longicolla by its smaller conidia
(12–15 9 4–7 lm in N. longicolla). This is the first report
123
of Neodidymelliopsis reported on Sambucus species (Farr
and Rossman 2019).
Nothophoma Qian Chen & L. Cai
Notes: The genus Nothophoma was introduced by Chen
et al. (2015) with the type species Nothophoma infossa
(Ellis & Everh.) Q. Chen & L. Cai. (syn. Phoma infossa)
and five other species transferred from Phoma, namely N.
anigozanthi (syn. P. anigozanthi), N. arachidis-hypogaeae
(syn. P. arachidis-hypogaeae), N. quercina (syn. P.
fungicola), and N. gossypiicola (syn. P. gossypiicola)
(Chen et al. 2015). There are nine species in this genus with
the recently introduced N. variabilis (Valenzuela-Lopez
et al. 2018). Ovoid, oblong to ellipsoidal aseptate conidia
are a characteristic of the genus Nothophoma. This genus
Fungal Diversity (2019) 96:1–242
37
Fig. 21 Phylogram generated
from maximum likelihood
analysis based on combined
LSU, ITS, TUB2 and RPB2
sequence data for
Didymellaceae. Related
sequences are taken from Chen
et al. (2015), Jayasiri et al.
(2017), Thambugala et al.
(2017a) and Wanasinghe et al.
(2018b). Fifty-nine strains are
included in the combined genes
sequence analyses which
comprise total 2683 characters
(960 characters for LSU, 547
characters for ITS, 360
characters for TUB2, 816
characters for RPB2) after
alignment. Leptosphaeria
conidia (CBS 616.75) and L.
doliolum (CBS 505.75)
(Leptosphaeriaceae,
Pleosporales) are used as the
outgroup taxa. Single gene
analyses are carried out and the
topology of each tree had clade
stability. Tree topology of the
maximum likelihood analysis is
similar to the Bayesian analysis.
The best sorting RaxML tree
with a final likelihood value of
- 16885.291744 is presented.
The matrix had 945 distinct
alignment patterns, with 21.62%
undetermined characters or
gaps. Estimated base
frequencies were as follows:
A = 0.241571, C = 0.241856,
G = 0.271818, T = 0.244754;
substitution rates
AC = 1.630628,
AG = 4.058235,
AT = 1.775626,
CG = 0.954941,
CT = 9.716323,
GT = 1.000000; gamma
distribution shape parameter
a = 0.414994. Branches with
Bayesian posterior probabilities
(BYPP) equal or greater than
0.95 are in bold. The ex-type
strains are in bold and black.
The newly generated sequences
are indicated in bold and blue
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Fig. 22 Neodidymelliopsis sambuci (MFLU 18-0177, holotype).
a Appearance of pycnidia on dead branch of Sambucus nigari.
b Appearance of pycnidia under the light microscope. submerged
conidiomata on the host surface. c–e Conidia on host. f Pycnidia on
agar. g pycnidial wall. h Developing hyphae. i–k Conidia on PDA.
l Upper view of 7 days old culture on PDA. m Reverse view of 7 days
old culture on PDA. Scale bars: a = 1000 lm, b, f = 100 lm, c,
k = 20 lm, d, e, i, j = 10 lm
contains ubiquitous fungal species, of which many are
plant pathogens, some of which are of quarantine concern
(Chen et al. 2015).
Pathogenic on leaves of Buxus megistophylla H.Lév.
Sexual morph Undetermined. Asexual morph Conidiomata produced on the PDA surface, 0.19–0.47 mm
(
x = 0.25 mm, n = 10) diam., pycnidial, solitary, scattered,
globose to irregularly shaped, black, ostiolate. Pycnidial
wall multi-layered, composed of pale brown,
Nothophoma quercina (Syd. & P. Syd.) Q. Chen & L. Cai,
Stud. Mycol. 82: 213. 2015.
Facesoffungi number: FoF04974; Fig. 23
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39
Fig. 23 Nothophoma quercina
(JZB380024, new host record).
a, b Conidiomata on the host
tissue. c Section of the pycnidial
wall. d Pycnidia on PDA.
e Conidiogenous cells on
culture. f Conidia on culture.
g Upper view on colonies on
PDA. h Lower view on colonies
on PDA. Scale bar: c = 1 mm,
d–f = 20 lm
pseudoparenchymatous cells. Conidiogenous cells phialidic, hyaline, simple, doliiform to ampulliform, variable in
size. Conidia 4–5.6 9 1.4–3.3 lm (
x = 5.2 9 2.3 lm,
n = 40), variable in size and shape, subglobose to oval or
obtuse, initially hyaline, but brown at maturity, aseptate,
smooth-walled. Conidial exudates not recorded.
Culture characteristics: Colonies on PDA reach 80 mm
diam. after 7 days at 28 C, with regular margin, dull white
aerial mycelium surface floccose to wooly, greenish
olivaceous to olivaceous near the centre and reverse dark
ochreous in the centre and white in the margin.
Material examined: CHINA, Haidian District, Beijing,
on leaves of Buxus megistophylla (Buxaceae), November
2017, D. Harishchandra, living culture (JZB380024,
MFLUCC 18-1568).
GenBank numbers: ITS: MK070136, LSU: MK070139.
Notes: Nothophoma quercina has been reported as a
pathogen on Pistacia vera (Chen et al. 2013), Chaenomeles
sinensis (Yun and Oh 2016), Olea europaea (Moral et al.
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40
2017) and was reported as a saprobe on Quercus sp. in
Ukraine (Chen et al. 2015). This is the first record of N.
quercina on Buxus sp. Phylogenetic analysis with the
combined sequence data of ITS, TEF1-a, TUB2 and RPB2
of our strain (MFLUCC 18-1568) clusters together with the
ex-type strain of Nothophoma quercina (CBS 633.92) with
relatively high bootstrap and Bayesian probabilities (99%
MP/1.00 PP).
Hermatomycetaceae Locq.
Notes: Hashimoto et al. (2017) introduced Hermatomycetaceae for a separate lineage of Hermatomyces species in Pleosporales, with Hermatomyces Speg. as the
generic type. Hermatomycetaceae was introuduced by
Locquin (1984) based on their distinctive characteristics,
recently, the family placement had been formalised with a
robust phylogenetic information (Doilom et al. 2017;
Hashimoto et al. 2017). Several taxa have been introduced
in the family, interestingly, no sexual morph has been
discovered (Koukol et al. 2018; Tibpromma et al.
2016, 2018). In this study, we introduce a novel Hermatomyces species based on the distinctive morphology
and supportive phylogenetic relationships (Fig. 24).
Hermatomyces Speg.
Notes: Hermatomyces was introduced by Spegazzini
(1911) with H. tucumanensis Speg. as a type species. The
genus is commonly present as sporodochial conidiomata
and muriform, lenticular, hyaline or dematiaceous conidia
of one or two types (Chang 1995; Tibpromma et al. 2016;
Hashimoto et al. 2017). Koukol et al. (2018) synonymized
several species of Hermatomyces based on morphological
similarity and possible artefacts in the phylogenetic analysis. Tibpromma et al. (2018) confirm that the species
homogeneity needs sufficient evidence from characters and
TUB2 gene sequence data are still required (Jeewon and
Hyde 2016). Therefore in this study, we follow Tibpromma
et al. (2018) and introducing a novel species of Hermatomyces, H. bauhiniae based on its morphological and phylogenetically distinct from other Hermatomyces species
(Fig. 24).
Hermatomyces bauhiniae Phukhams., D.J. Bhat & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555500; Facesoffungi
number: FoF04827; Fig. 25
Etymology: Name refers to the host plant, Bauhinia
variegata from which this fungus was isolated.
Holotype: MFLU 18-1381
Saprobic on dead branch of Bauhinia variegata L.
Sexual morph Undetermined. Asexual morph Colonies
on natural substrate forming sporodochial conidiomata,
superficial, scattered, circular or oval, blackish brown,
velvety, glistening, consisting of an orbicular, abundantly
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sporulation, conidia readily liberated when agitated, 160–
180 lm wide. Mycelium mostly superficial, composed of a
loose or compact network of repent, branched, septate,
rough-walled, thick-walled, reddish brown to brown
hyphae, 2–4 lm wide; subicular hyphae short, irregularly
geniculate or flexuous, densely packed. Conidiophores 4–
10 9 2–5 lm, micronematous or semimacronematous,
mononematous, cylindrical, erect, smooth or finely verruculose, aseptate, unbranched, often corresponding to
conidiogenous cells, hyaline. Conidiogenous cells 3–
8 9 4–9 lm, holoblastic, monoblastic, integrated, terminal, determinate, cylindrical or slightly subulate, subsphaerical or ampulliform, hyaline. Conidia dimorphic,
solitary, smooth-walled: lenticular conidia: 25–36 9 15–
20 lm (
x = 30 9 18 lm, n = 70), muriform, smooth,
broadly ellipsoidal to oval in front view, brown to dark
brown, slightly constricted at the septa, in side view
composed of one column of 4–6 cells, end cells subhyaline
to pale brown, often carrying remnant of conidiogenous
cell at base; cylindrical conidia: 20–28 9 8–11 lm
(
x = 24 9 9 lm, n = 30), straight or flexuous, septate,
constricted at the septa, consisting of one column, 2–3septate, doliiform, cylindrical or subcylindrical, apical cells
clavate or doliiform, verrucose, apex rounded, basal cells
globose or subglobose, smooth, hyaline.
Culture characteristics: Colonies on MEA at room
temperature (25 8C) reaching 7 cm in 2 weeks, circular
with entire margin, with white–grey mycelium, white at the
margin, smooth at the surface and raised, mycelium
strongly radiating into agar; reverse beige, no sporulation
in cultures.
Material examined: THAILAND, Phrae Province, on
dried branch of Bauhinia variegata (Fabaceae), 25 July
2015, C. Phukhamsakda, S1_02 (MFLU 18-1381, holotype), ex-type living culture (MFLUCC 16-0395, ICMP
21948).
GenBank numbers: ITS: MK443382, LSU: MK443378,
RPB2: MK443386, SSU: MK443380, TEF1-a:
MK443384.
Notes: Hermatomyces bauhiniae is introduced as a new
species based on its distinct morphology and support from
phylogenetic analysis. Hermatomyces bauhiniae has distinct characteristics as compared to the other species. The
lenticular conidia are not round or flat in one plane, and
have well visible conidiogenous cells attached at the bottom of the conidia. In the phylogenetic analysis, H. bauhiniae formed a clade with H. indicus with good support
(MLBS = 82; BYPP = 0.99). Both species were recorded
from angiosperms plants. Nonetheless, H. bauhiniae can be
distinguished from H. indicus by having straight conidia
with only one column, hyaline, and cylindrical conidia,
where as in H. indicus conidia are turbinate, grey-brown at
the apex, and become hyaline towards the base with two
Fungal Diversity (2019) 96:1–242
41
Fig. 24 Phylogram of 50% majority rule consensus tree from the
Bayesian-inference analysis based on combined dataset of LSU, ITS,
TEF1-a, and RPB2 sequence data representing Hermatomycetaceae.
Related sequences are taken from Doilom et al. (2017), Hashimoto
et al. (2017), Koukol et al. (2018) and Tibpromma et al. (2018).
Thirty-nine strains are included in the combined analyses which
comprise 3301 characters (826 characters for LSU, 514 characters for
ITS, 941 characters for TEF1-a, 1020 characters for RPB2) after
alignment. The tree is rooted with Anteaglonium globosum (ANM
925.2) and A. parvulum (MFLUCC 14-0821) in Anteagloniaceae
(Pleosporales). Single gene analyses were also performed to compare
the topology and clade stability with combined gene analyses. Tree
topology of the maximum likelihood analysis is similar to the
Bayesian analysis. Bootstrap support values for maximum likelihood
analysis greater than 70% and clade credibility values greater than
0.90 (the rounding of values to 2 decimal proportions) from Bayesianinference analysis labeled on the nodes. The matrix has 710 distinct
alignment patterns, with 20.65% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.244863,
C = 0.263683, G = 0.262259, T = 0.229195; substitution rates AC =
1.191383, AG = 5.252837, AT = 1.421242, CG = 1.059773, CT =
14.708510, GT = 1.000000; gamma distribution shape parameter
a = 0.766883. The ex-type strains are in bold and black. The newly
generated sequence is indicated in bold and blue
columns. Hermatomyces indicus has larger cylindrical
conidia but smaller lenticular conidia (Prasher and Sushma
2014; Doilom et al. 2017).
Lophiostomataceae Sacc.
Notes: The family was introduced as ‘‘Lophiostomeae’’
by Nitschke (1869) and recorded as ‘‘Lophiostomaceae’’ in
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b Fig. 25 Hermatomyces bauhiniae (MFLU 18-1381, holotype). a–
c Sporodochia on natural substrate. d Vertical section through
sporodochia. e–g Subicular hyphae. h Cylindrical conidia and
lenticular conidia on host substrate. i, j Cylindrical conidia. k–m
Mature lenticular conidia. n Conidia. o Germinated conidium. Scale
bars: a = 1 cm, b = 500 lm, c, d, f = 100 lm, o = 50 lm, h–
m = 20 lm, e, g = 10 lm
Saccardo (1883), but was accepted as Lopiostomataceae in
Mugambi and Huhndorf (2009). Members of this family
are saprobes and are found in terrestrial, aquatic and
marine habitats (Tanaka and Harada 2003; Zhang et al.
2009c; Hirayama and Tanaka 2011; Hyde et al. 2013;
Thambugala et al. 2015; Jones et al. 2015; Tennakoon et al.
2018). A generic re-evaluation of Lophiostomataceae by
Thambugala et al. (2015) based on multi-locus phylogenies
using LSU, SSU, ITS and TEF1-a sequence data, integrated 16 genera into this family, and resolved the phylogenetic
and
morphological
boundaries
of
Lophiostomataceae. Hashimoto et al. (2018) revised the
taxonomical classification of Lophiostoma bipolare (K.D.
Hyde) E.C.Y. Liew, Aptroot & K.D. Hyde, and proposed a
novel genus Lentistoma A. Hashim., K. Hiray. & Kaz.
Tanaka to accommodate Lophiostoma sensu stricto. Four
additional new genera and two new genera combinations
were recognized from the other lineages of the L. bipolare
complex. Therefore, there are currently 23 genera in
Lophiostomataceae.
Vaginatispora K.D. Hyde
Notes: Vaginatispora was introduced by Hyde (1995b)
with V. aquatica K.D. Hyde as the type species. Vaginatispora aquatica was initially placed in Massarinaceae,
but Zhang et al. (2014a) revealed its phylogenetic affinity
with Lophiostomataceae. Subsequently, Thambugala et al.
(2015) considered Vaginatispora as a separate genus within
Lophiostomataceae based on multi-gene phylogeny and
morphology, of which V. aquatica and V. fucklii were
included. Thereafter, V. appendiculata (Wanasinghe et al.
2016), V. armatispora (: Massarina armatispora K.D.
Hyde, Vrijmoed, Chinnarij & E.B.G. Jones) (Hyde et al.
1992; Liew et al. 2002; Wanasinghe et al. 2016), V.
microarmatispora (Devadatha et al. 2017), V. amygdali
and V. scabrispora (Hashimoto et al. 2018) were introduced based on morphology and phylogeny. However, V.
fuckelii was transferred to a new genus Neovaginatispora
(Hashimoto et al. 2018) based on phylogenetic distinction
and its typical characters of erumpent ascomatal and a
thinner peridium. During observations of microfungi from
marine (mangrove) and freshwater habitats, Vaginatispora
palmae sp. nov. and V. armatispora were reported and
illustrated herein. An updated phylogenetic tree is
43
provided, which includes all Vaginatispora taxa and closely related genera in Lophiostomataceae.
Vaginatispora palmae S.N. Zhang, J.K. Liu & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556316; Facesoffungi
number: FoF05089; Fig. 26
Etymology: The epithet reflects the family of host plant.
Holotype: MFLU 18-1586
Saprobic on immersed rachis of Nypa fruticans. Sexual
morph Ascomata in vertical section 250–340 lm high,
215–385 lm diam. (
x = 310 9 326 lm, n = 10), dark
brown to black, scattered, semi-immersed, erumpent, subglobose to elongated, base flatted, coriaceous to carbonaceous. Ostiole crest-like, variable in shape, central
papillate. Peridium 15–38 lm wide, wider at the apex and
thinner at the base, composed of several pale brown to
brown cells of textura angularis, cells towards the inside
lighter, at the outside darker, somewhat flattened, fusing
and with the host tissues. Pseudoparaphyses 1–2.5 lm
wide, hypha-like, numerous, septate, rarely branched and
anastomosed, tapering towards the apex. Asci 89–
115 9 12–20 lm (
x = 100.5 9 16.0 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindric-clavate, with a
short bulbous pedicel, rounded at the apex, with an ocular
chamber.
Ascospores
23–45 9 6–9 lm,
(
x = 35.3 9 7.5 lm, n = 30), hyaline, uniseriate or overlapping to biseriate, 1-septate, occasionally producing
pseudosepta, slightly constricted at the central septum, cell
above central septum swollen, guttulate, smooth-walled,
surrounded by a narrow mucilaginous sheath and drawn out
towards each end to form tapering appendages, 6–8 lm
long. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
PDA within 24 h. Colonies growing on PDA reaching
2 cm diam. after 21 days at 25 C, the off-white hyphae in
first week, then becoming grayish blue and dark bluish,
composed of brown to dark brown, septate, smooth or
verrucose hyphae.
Material examined: THAILAND, Ranong Province, on
immersed rachis of Nypa fruticans Wurmb (Arecaceae), 3
December 2016, S.N. Zhang, SNT92 (MFLU 18-1586,
holotype), ex-type living culture (MFLUCC 18-1526);
ibid. (HKAS 102207, isotype).
GenBank numbers: ITS: MK085055, LSU: MK085059,
SSU: MK085057, TEF1-a: MK087657.
Notes: Most species of Vaginatispora are found in
tropical regions and commonly occur in freshwater and
marine environments, but only one species V. nypae
Jayasiri, E.B.G. Jones & K.D. Hyde (Jayasiri et al. 2019)
has recently been recorded from a Nypa fruticans. Almost
all Vaginatispora species have 1-septate ascospores with
terminal appendages or sheaths (Hashimoto et al. 2018).
123
44
Fig. 26 Vaginatispora palmae (MFLU 18-1586, holotype). a, b
Appearance of stromata on host surface. c Structure of peridium.
d Vertical section through the stromata with ascomata. e Pseudoparaphyses. f–h Ascus. i–k Ascospores. l Ascospore in India Ink, with
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Fungal Diversity (2019) 96:1–242
clear terminal appendages and narrow sheath. Sacle bars:
b = 500 lm, a = 200 lm, d = 100 lm, c, e, f–h = 20 lm, i–
l = 10 lm
Fungal Diversity (2019) 96:1–242
45
Detailed characters of ascospores are useful for distinguishing taxa at the species level, and molecular sequence
data is key for identification of taxa in this group. The
multi-gene analysis includes all species in this genus and
the new isolate Vaginatispora palmae clustered with V.
amygdali (Fig. 28). Vaginatispora palmae differs from V.
amygdali because the latter species has a lateral pad-like
structure within the sheath and an internal chamber at both
ends of the ascospores, while V. palmae lacks those
structures. They also differ in peridium width (15–38 lm
vs. 37.5–62.5 lm), ascus (mean: 100.5 9 16 lm vs.
115 9 18.5 lm)
and
ascospores
size
(mean:
35.3 9 7.5 lm vs. 30.6 9 8.8 lm). In addition, polymorphic nucleotide comparison shows that these two strains
differ in eight positions in the ITS1 and ITS2 regions
including two gaps, and differ in eight positions in the
TEF1-a region. The difference in molecular sequence data
also distinguish V. palmae from V. amygdali, therefore, we
introduce a new species V. palmae.
site1–4–1, (MFLU 18-1229), living culture (MFLUCC
18-0247).
GenBank numbers: ITS: MK085056, LSU: MK085060,
RPB2: MK087669, SSU: MK085058, TEF1-a:
MK087658.
Notes: The strain MFLUCC 18-0247 that was isolated
and described in this study, phylogenetically clustered
together with Vaginatispora armatispora (HKUCC 1562)
(Wanasinghe et al. 2016) with high support (Fig. 28). On
the other hand, these two V. armatispora strains (MFLUCC
18-0247 and HKUCC 1562) have only one base pair difference in the ITS1 region. These two isolates are quite
similar, but isolate MFLUCC 18-0247 has a thicker
peridium than that of HKUCC 1562. Based on such
insignificant differences in morphology and molecular
data, we consider the isolate (MFLUCC 18-0247) and V.
armatispora are conspecific. Vaginatispora armatispora
can be distinguished from other members in Vaginatispora
because it has pale brown ascospores.
Vaginatispora armatispora (K.D. Hyde, Vrijmoed, Chinnaraj & E.B.G. Jones) Wanas., E.B.G. Jones & K.D. Hyde
Facesoffungi number: FoF05060; Fig. 27
Saprobic on decaying wood, submerged wood in
freshwater and marine habatits. Sexual morph Ascomata
325–370 lm
diam.
300–340 lm
high,
(
x = 350 9 317 lm, n = 10), scattered, usally immersed
or semi-immersed, globose to subglobose, black. Ostiole
black, crest-like, central, filled with brown cells. Peridium
20–32 lm wide, 2–3 layers, composed of dark brown to
hyaline, thick-walled cells of textura porrecta at side and
textura angularis at the base. Hamathecium comprising 2–
3 lm wide, septate, hyaline, pseudoparaphyses composing
situated between and above the asci, embedded in a
gelatinous
matrix.
Asci
94–118 9 15–17 lm
(
x = 106 9 16 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical-clavate, short pedicellate, apically
rounded with an ocular chamber, hyaline to subhyaline.
Ascospores 29–34 9 7–9 lm (
x = 32 9 8 lm, n = 30),
fusiform, hyaline to pale brown, smooth-walled, 1-septate,
constricted at the septum, tapering to pointed apices, distinct large guttules in each cells, with distinct hyaline
appendages, surrounded by a narrow mucilaginous sheath
or not. Asexual morph undetermined.
Culture characteristics: Ascospores germinating on
PDA within 24 h. Colonies on MEA at room temperature
reaching 3 cm diam. in three weeks, mycelium pale brown
at first, grayish brown after 3 weeks, composed pale brown
to brown, septate, smooth hyphae.
Material examined: THAILAND, Prachuap Khiri Khan
Province, Bang Saphan District, Sai Khu Waterfall, on
submerged decaying wood, August 2017, V. Kumar,
Massariaceae Nitschke
Notes: The family Massariaceae was introduced by
Nitschke (1869) to accommodate the genus Massaria De
Not. with M. inquinans (Tode) De Not. as the type species.
The family is characterized by immersed, globose, subglobose to pyriform, ascomata, with a thin-walled peridium
comprising cells of textura angularis, oblong to cylindrical
asci with a wide ocular chamber and refractive ring, and
large, oblong to ellipsoidal ascospores surrounded by a
gelatinous sheath (Voglmayr and Jaklitsch 2011; Hyde
et al. 2013, 2016). Based on combined LSU, SSU and
TEF1-a sequence data, Hyde et al. (2016) introduced the
genus Neomassaria Mapook, Camporesi & K.D. Hyde in
Massariaceae but Ariyawansa et al. (2018) introduced
Neomassariaceae Ariyawansa, Jaklitsch & Voglmayr to
accommodate this genus. Based on morphology and LSU
phylogeny, Huanraluek et al. (2018) described the genus
Massarioramusculicola Huanraluek., Thambugala & K.D.
Hyde. At present, the family comprises 31 Massaria species and a single Massarioramusculicola species. In this
study, we introduced a new genus Paramassaria typified
with Paramassaria samaneae.
Paramassaria Samarak., & K.D. Hyde, gen. nov.
Index Fungorum number: IF555521; Facesoffungi
number: FoF05213
Etymology: Named because of its morphological similarity to Massaria.
Saprobic on branches. Sexual morph Ascomata
immersed or semi-immersed in the yellow stained host,
solitary, scattered, coriaceous, globose to subglobose,
brown to dark brown. Ostiole central, raised, flattened
around the dome-shaped ostiole, surrounded by blackish
stromatic zone on the host, minutely papillate. Peridium
123
46
Fig. 27 Vaginatispora armatispora (MFLU 18-1229, new record).
a Ascomata on submerged wood. b Section of ascoma. c–e Section of
peridium. f Pseudoparaphyses. g–j Asci. k–m Ascospores.
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Fungal Diversity (2019) 96:1–242
n Germinating ascospore. o, p Culture on PDA. Scale bars:
b = 100 lm, c–e, g–n = 20 lm, f = 10 lm
Fungal Diversity (2019) 96:1–242
47
Fig. 28 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU, SSU, TEF1-a and RPB2 sequence data
for Vaginatispora species and several closely related genera in
Lophiostomataceae. Related sequences are taken from Thambugala
et al. (2015), Wanasinghe et al. (2016), Devadatha et al. (2017) and
Hashimoto et al. (2018). Thirty-four strains are included in the
combined analyses which comprise 4243 characters (605 characters
for ITS, 832 characters for LSU, 895 characters for SSU, 894
characters for TEF1-a, 1017 characters for RPB2) after alignment.
Angustimassarina populi (MFLUCC 13–0034) and A. acerina
(MFLUCC 14-0505) (Amorosiaceae, Pleosporales) are used as the
outgroup taxa. Single gene analyses are carried out and the topology
of each tree had clade stability. Tree topology of the maximum
likelihood analysis is similar to the maximum parsimony analysis and
the Bayesian analysis. The best RaxML tree with a final likelihood
value of –17911.101212 is presented. The matrix had 1059 distinct
alignment patterns, with 20.20% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.248793,
C = 0.250421, G = 0.268979, T = 0.231807; substitution rates AC =
1.458334, AG = 3.325243, AT = 1.282598, CG = 1.395939, CT =
8.065988, GT = 1.000000; gamma distribution shape parameter
a = 0.592418. Maximum parsimony analysis of 3144 constant
characters and 854 informative characters resulted in two equally
most parsimonious tree with a tree length as 2508 (CI = 0.622,
RI = 0.822, RC = 0.511, HI = 0.378). Bootstrap values for maximum
likelihood (ML) and maximum parsimony (MP) equal to or greater
than 75 are placed above and below the branches respectively.
Branches with Bayesian posterior probabilities (BYPP) equal or
greater than 0.95 are in bold. The ex-type strains are in bold and
black. The newly generated sequences are indicated in bold and blue
wide around the ostiole, outer layer; thick-walled, reddish
to dark brown cells of textura angularis, inner layer; thinwalled hyaline cells of textura angularis. Hamathecium
comprising wide, hyaline, cylindrical to filiform, septate,
branched pseudoparaphyses. Asci 8-spored, bitunicate,
oblong to cylindrical, absence of prominent ocular chamber, shortly pedicellate. Ascospores overlapping 1–2-seriate, ellipsoid to broadly fusiform, initially hyaline,
1-septate becoming brown and 3-septate, constricted at the
septum, surrounded by a hyaline gelatinous sheath at
maturity. Asexual morph Undetermined.
Notes: Paramassaria samaneae was collected on a dead
branch of Samanea saman from Thailand. This taxon is
similar to Massaria in its ascomata, asci and ascospore
characters. The wide ostiole and absence of an ocular
chamber in the asci distinguishes our taxon from Massaria.
However, the combined phylogenetic analyses (Fig. 29)
separate our taxon as a basal clade to Massaria. The closest
BLAST search with the LSU and TEF1-a sequences of
MFLU 17-1551 were GQ221892 (with 93% identity)
Rhytidhysteron opuntiae and FJ161103 (with 93% identity)
Gloniopsis praelonga belonging to Hysteriaceae. Only ITS,
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Fungal Diversity (2019) 96:1–242
b Fig. 29 Phylogram generated from Bayesian analysis based on
combined LSU, SSU and TEF1-a sequence data for Massariaceae
and several closely related families in Pleosporales. Related
sequences are taken from Voglmayr and Jaklitsch (2011), Hyde
et al. (2016) and Ariyawansa et al. (2018). One-hundred and one
strains are included in the combined analyses which comprise 2519
characters (806 characters from LSU, 840 characters for SSU, 873
characters for TEF1-a) after alignment. Bayesian posterior probabilities (BYPP) equal to or greater than 0.9 are on branches. The newly
generated sequences are indicated in bold and blue
LSU and SSU sequences are available for Massarioramusculicola and resulted in inconsistent topologies with
the absence sequence data. In addition, we observed that
our strains have probabilities to cluster in Aigialacaeae as a
basal clade. However, Massarioramusculicola differs from
Massaria and Paramassaria by having a peridium with two
strata with dark brown to hyaline cells of textura angularis,
rarely branched, septate and trabeculate pseudoparaphyses
and 3-septate hyaline ascospores. Including parallel morphology to Massaria, we prefer to place our strains into a
new genus in Massariaceae. Voglmayr and Jaklitsch (2011)
suggested that the ascospore and pseudothecia characters
are important for the identification of phylogenetically
distinct Massaria species. The strong host-specificity
among Massaria species makes it complex for accurate
identification (Voglmayr and Jaklitsch 2011). It might also
not be a surprise if future discoveries of more species
within Massariaceae converge the clade and there is a need
to aggregate them into one genus, but it is unwise to do this
at this stage.
Type species: Paramassaria samaneae Samarak., &
K.D. Hyde
Paramassaria samaneae Samarak & K.D. Hyde, sp. nov.
Index Fungorum number: IF555522; Facesoffungi
number: FoF05214; Fig. 30
Etymology: Name based on the host Samanea, from
which it was collected.
Holotype: MFLU 17-1551
Saprobic on a dead branch of Samanea saman. Sexual
morph Ascomata 520–580 lm high (
x = 540 lm, n = 15),
410–560 lm diam. (
x = 490 lm, n = 15), immersed or
semi-immersed in the yellow stained host, solitary, scattered, coriaceous, globose to subglobose, brown to dark
brown. Ostiole central, flattened on the top, surrounded by
blackish stromatic zone on the host, ostiolar canal with
hyaline
periphyses.
Peridium
20–43 lm
wide
(
x = 33.5 lm, n = 25), inner cell layer hyaline, thin and
outer cell layer comprising dark brown textura angularis
cells. Hamathecium comprising 4.2–10 lm wide
(
x = 6.6 lm, n = 25), hyaline, cylindrical to filiform, septate, branched pseudoparaphyses. Asci 190–215 9 45.5–
56 lm
(
x = 212.5 9 50.5 lm,
n = 20),
8-spored,
49
bitunicate, oblong to cylindrical, short pedicellate. Ascospores 52.5–71.5 9 13.5–18.5 lm (
x = 61.5 9 16 lm,
n = 50), l/w 3.9, overlapping 1–2-seriate, brown, ellipsoid
to broadly fusiform, 3-septate, constricted at the septa, with
4 guttlues, surrounded by hyaline gelatinous sheath
observed clearly when mounted in Indian ink. Asexual
morph Undetermined.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Mae Fah Luang University, on a dead
branch of Samanea saman (Fabaceae), 2 August 2017, MC.
Samarakoon, SAMC002 (MFLU 17-1551, holotype;
HKAS 102338, isotype).
GenBank numbers: LSU: MK108190, MK108191, SSU:
MK108187, MK108188, TEF1-a: MK105747, MK105747.
Phaeosphaeriaceae M.E. Barr
Notes: Phaeosphaeriaceae is the largest family in
Pleosporales and was introduced by Barr (1979). It is
typified with Phaeosphaeria with the type species Phaeosphaeria oryzae (Phookamsak et al. 2014). Phaeosphaeriaceae comprises 52 genera (Wijayawardene et al. 2018a).
Phaeosphaeriaceae is a heterogenous family of saprobes,
pathogens and endophytes on plants (Tennakoon et al.
2016a; Phookamsak et al. 2017; Tibpromma et al. 2018).
The family is characterized by bitunicate asci with hyaline,
yellow or brown spores that are aseptate or septate and
holoblastic or enteroblastic conidiogenesis. Seven new
species belonging to Muriphaeosphaeria (M. angustifoliae), Neosetophoma (N. miscanthi, N. salicis), Phaeopoacea (P. asparagicola), Phaeosphaeria (P. penniseti)
and Nodulosphaeria (N. thalictri, N. aquilegiae) are
introduced.
Muriphaeosphaeria Phukhamsakda et al.
Notes: The genus Muriphaeosphaeria was introduced by
Phukhamsakda et al. (2015) based on the type species
Muriphaeosphaeria galatellae found on dead and drying
stems of Galatella villosa (L.) Rchb.f. (Asteraceae) in
Russia. The morphological characters of the genus comprise superficial ascomata with a peridium containing thinwalled brown cells of textura angularis, cellular pseudoparaphyses and muriform ascospores. The asexual
morph is coelomycetous and is characterized by conidiomata with a thick, hyaline, inner wall layer and cylindrical to subclavate, 1–3-transversely septate, brown
conidia (Phukhamsakda et al. 2015). Muriphaeosphaeria
contains three species (Index Fungorum 2019). Cultures
and sequences are available.
Muriphaeosphaeria angustifoliae D. Pem, Gafforov &
K.D. Hyde, sp. nov.
Index Fungorum number: IF555432; Facesoffungi
number: FoF05109; Fig. 31
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b Fig. 30 Paramassaria samaneae (MFLU 17–1551, holotype). a, b
Ascomata on the substrate. c Horizontal section of ascoma. d Vertical
section of ascoma. e Ostiole section. f Peridium (in water).
g Pseudoparaphyses. h–j Asci. k–s Ascospores (s-in Indian ink).
Scale bars: a = 1000 lm, b–d = 200 lm, e = 100 lm, h–j = 50 lm,
f, g, k–s = 20 lm
Etymology: Name reflects the host from which the fungus was isolated.
Holotype: TASM 6138
Saprobic on dead branches of Perovskia angustifolia.
Sexual morph Ascomata 200–600 lm high 9 200–
500 lm diam. (
x = 228.2 9 226.5 lm, n = 10), scattered
or sometimes clustered, or solitary, semi-immersed to
superficial, globose to subglobose, brown to dark brown,
with papillate central ostiole. Ostiole 17–18 lm high 9
28–33 lm wide, slightly raised, centrally located, lacking
periphyses. Peridium 70–90 lm thick, smooth-walled
comprising 2–3 layers of brown to dark brown, pseudoparenchymatous cells of textura angularis. Hamathecium
comprising numerous, dense, 1–1.5 lm wide, filiform,
broadly cellular pseudoparaphyses, branched and anastomosing, embedded in gelatinous matrix. Asci 125–
175 9 13–15 lm (
x = 148 9 14.5 lm, n = 10), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate,
short pedicellate or subsessile, apically rounded with an
ocular
chamber.
Ascospores
21–28 9 9–11 lm
(
x = 25.9 9 10.1 lm, n = 10), overlapping uniseriate,
initially hyaline, becoming yellowish brown at maturity,
broadly fusiform, multiseptate, constricted at the middle
septa, narrowly rounded at both ends, smooth-walled,
swollen, lacking a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 48 h. Colonies growing on MEA, reaching
3 cm diam. in 1 week at 16 C. Mycelium superficial,
surface smooth, circular, hairy, margin well defined, shiny,
from above white, reverse pale yellow and grey in the
middle.
Material examined: UZBEKISTAN, Surxondaryo,
Boysun, Omonxona Village, South-Western Hissar
Mountains, on dead branches of Perovskia angustifolia
(Lamiaceae), 6 April 2016, Y. Gafforov, YG-S29-2
(TASM 6138, holotype; MFLU 17-0070, isotype), ex-type
living culture (MFLUCC 18-1124).
GenBank numbers: ITS: MK101002, LSU: MK100999,
SSU: MK100997.
Notes: Muriphaeosphaeria angustifoliae constitutes a
strongly supported independent lineage closely nested with
M. ambrosiae (MFLU 15-1971). Muriphaeosphaeria
ambrosiae is only known from its asexual morph and
cannot
be
compared
to
M.
angustifoliae.
51
Muriphaeosphaeria angustifoliae is morphologically more
similar to M. galatellae (Phukhamsakda et al. 2015) in
having superficial ascomata and broadly fusiform multiseptate ascospores lacking a mucilaginous sheath. However,
M.
angustifoliae
has
larger
ascomata
(200–600 lm 9 200–500 lm
v/s
114–180 9 167–263 lm), a wider peridium (70–90 lm
v/s 12–26 lm), narrower hamathecium (1–1.5 lm v/s
2–3 lm) and longer asci (125–175 9 13–15 lm v/s
53–86 9 9–17 lm). We therefore, introduce M. angustifoliae as a new species in Phaeosphaeriaceae.
Neosetophoma Gruyter et al.
Notes: Neosetophoma was introduced by de Gruyter
et al. (2010) and is typified by N. samararum (Desm.)
Gruyter, Aveskamp. & Verkley. Neosetophoma comprises
15 species (Wijayawardene et al. 2017a) and contains
important plant pathogens, saprobes and soil fungi
(Phookamsak et al. 2014; Karunarathna et al. 2017). The
genus is characterized by globose to irregular conidiomata
with papillate ostioles and yellowish conidia attenuated at
one end (Wijayawardene et al. 2017a). In this study
Neosetophoma miscanthi and Neosetophoma salicis are
introduced.
Neosetophoma miscanthi Karun., C.H. Kuo & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556266; Facesoffungi
number: FoF05848; Fig. 32
Etymology: Name reflects the host genus, Miscanthus
from which the fungus was isolated.
Holotype: MFLU 18-2675
Saprobic on dead stems of Miscanthus gigantius (Poaceae), as raised, black, shiny globular structures on the host
surface. Sexual morph Ascomata 90–130 lm high, 110–
120 lm diam. (
x = 109.5 9 115 lm, n = 5), solitary,
gregarious, immersed under epidermis, becoming erumpent
through host surface, subglobose, or polygonal at sides,
uni-loculate, glabrous, ostiolate, with minute papilla;
papilla 30–50 lm long (n = 5), carbonaceous, easily broken. Peridium 19–20 lm wide, thin-walled, of equal
thickness, composed of few layers of dark brown to brown,
pseudoparenchymatous cells of textura angularis. Hamathecium of dense, filamentous, septate, cellular pseudoparaphyses, embedded in a hyaline gelatinous matrix.
Asci 45–50 9 10–12 lm (
x = 48 9 10.5 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindric-clavate to
obclavate, slightly broad at the base with short rounded
pedicel, apically rounded. Ascospores 18–21 9 4.7–
5.3 lm (
x = 20 9 5 lm, n = 40), overlapping 1–2-seriate,
hyaline, fusiform, with acute ends, occasionally 3-septate,
cell near the septum slightly larger, smooth-walled, guttulate. Asexual morph Undetermined.
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Fig. 31 Muriphaeosphaeria angustifoliae (TASM 6138, holotype).
a, b Appearance of ascomata on host surface. c Section through an
ascoma. d Peridium. e Hamathecium. f. Ostiole. g-i Asci. j-l
Ascospores. m Ascospore germination on MEA after 48 h. n, o
Culture characters on MEA (n from above view, o from below view).
Scale bars: a = 2000 lm, b = 500 lm, c = 100 lm, g–i = 50 lm, d,
f, m = 20 lm, e, j–l = 10 lm
Culture characteristics: Ascospores germinating on
PDA within 12 h reaching 20 mm diam. after 7 d at 25 C,
circular, floccose, flat with even margin, initially pale
yellow becoming greenish grey; reverse yellowish grey.
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53
Fig. 32 Neosetophoma miscanthi (MFU 18-2675, holotype). a, b
Appearance of ascomata on Miscanthus gigantius. c. Section through
ascoma. d. Section through ostiole. e. Section through peridium. f.
Pseudoparaphyses. g–j. Different developing stages of the asci. l–n.
Ascospores. o. Germinating ascospore. Scale bars: c–e = 50 lm,
f = 20 lm, g–o = 10 lm
Material examined: TAIWAN, Chia Yi Province,
Kwang Hwa, Miscanthus gigantius (Poaceae), 18 March
2018, A. Karunarathna AKTW 31 (MFLU 18-2675,
holotype), ex-type living culture (FU31023).
GenBank numbers: ITS: MK503820, LSU: MK503826,
SSU: MK503832.
Notes: Neosetophoma miscanthi is morphologically
similar to N. guiyangensis but is distinct in its asci and
ascospore characters. In N. miscanthi asci are cylindricclavate to obclavate, broad at the base whereas the asci of
N. guiyangensis are cylindrical to cylindric-clavate. Furthermore, N. miscanthi generally has smaller ascomata,
peridia and asci parameters than N. guiyangensis (Hyde
et al. 2018a). Neosetophoma miscanthi prominently consists with 3-septate ascospores, while N. guiyangensis
prominently consisting with 1–3(–5)-septate ascospores.
Neosetophoma miscanthi was found on Poaceae while N.
guiyangensis was found on tree branches. In phylogenetic
analysis the N. miscanthi and N. guiyangensis separates
with high support (90% ML/0.99 BYPP)
Associated with twigs and branches. Sexual morph
Undetermined. Asexual morph Conidiomata 8–10 9 3–
4.5 lm, immersed in host tissue, scattered, erumpent, discoid, circular, with locules, ostiolate. Ostioles at the same
level, with flattened top. Peridium brown to pale brown,
comprising a few layers of cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells lining the inner cavity, hyaline, smooth,
phialidic with prominent periclinal thickening and thick
channel, globose to doliiform, 8–10 9 3–4.5 lm, formed
from the inner most layer of pycnidial wall, hyaline,
smooth-walled. Conidia (6.9–)8–10 9 2–2.8(–3.1) lm
(
x = 8.7 9 2.8 lm, n = 30), unicellular, allantoid, hyaline,
smooth-walled with thick layer.
Material examined: UZBEKISTAN: Surxondaryo Province, Boysun District, Qizilnaur Village, South-Western
Hissar Mountains, twigs and branches of Salix sp. (Salicaceae), 17 June 2016, Y. Gafforov, YG-S109-2 (TASM
6147, holotype; MFLU 17-0118, isotype), ex-type living
culture (MFLUCC).
GenBank numbers: ITS: MK608025, LSU: MK608026.
Notes: Neosetophoma salicis is introduced as a new
species based on phylogenetic analysis, as it forms a separate branched as a sister taxon to N. rosae Jayasiri et al.,
which was isolated from twigs and branches of Rosa
canina L. in Italy (isolate MFLUCC 17-0844, Wanasinghe
Neosetophoma salicis Norphanphoun, Gafforov, T.C. Wen
& K.D. Hyde, sp. nov.
Index Fungorum number: IF555517; Facesoffungi
number: FoF06000; Fig. 33
Etymology: The specific epithet reflects the host genus.
Holotype: TASM 6147
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Fig. 33 Neosetophoma salicis (TASM 6147, holotype). a Stromatal
habit in wood. b Fruiting bodies on host surface. c Surface of fruiting
bodies showing the black ostioles. d Cross section of conidioma.
e Peridium. f Ostiolar. g–h Conidiogenous cell containing conidia.
i Conidia. Scale bars: d = 100 lm, e, f = 50 lm, g, h = 10 lm,
i = 5 lm
et al. 2018a) with strong support (Fig. 38). However, they
are different in morphological characteristics, the conidia
of N. salicis are shorter than N. rosae (8.7 9 2.8 versus
11.9 9 2.5 lm) and conidia lack septa.
Shoemaker (1976) re-circumscribed Nodulosphaeria and
transferred various species to Ophiobolus. Later, many
species were added to the genus based on morphology
(Shoemaker 1984; Shoemaker and Babcock 1987; Zhang
et al. 2012). There are 71 epithets listed under Nodulosphaeria (Index Fungorum 2019). Mapook et al. (2016)
revised the genus and confirmed its phylogenetic placement in Phaeosphaeriaceae.
Nodulosphaeria Rabenh., Klotzschii Herb.
Notes: The genus Nodulosphaeria was introduced by
Rabenhorst (1858) with N. hirta Rabenh. as type species. It
is characterized by brown setae at the ascomata apex, and
three- to multi-septate ascospores, with a swollen cell and
some with terminal appendages (Holm 1961; Shoemaker
1984). The genus was re-examined by Zhang et al. (2012)
and Phookamsak et al. (2014) based on type material of N.
hirta. Nodulosphaeria was previously treated as a synonym
of Leptosphaeria, but later reinstated by Holm (1957).
Nodulosphaeria was accommodated in Phaeosphaeriaceae
by Barr (1979) based on morphology and similarity to
Ophiobolus as the ascospores form enlarged cell(s).
123
Nodulosphaeria aquilegiae Chaiwan, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555508; Facesoffungi
number: FoF05200; Fig. 34
Fig. 34 Nodulosphaeria aquilegiae (MFLU 17-1273, holotype). a, c, c
d Ascomata on a dead stem of Aquilegia viscosa. b, i Peridium. e, f
Section of ascomata. g Pseudoparaphyses. h-n Asci. o-r Ascospores.
Scale bars: c, d = 200 lm, b, e, f–i = 50 lm, j–n = 20 lm, o–
r = 10 lm
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55
123
56
Etymology: Name reflects the host from which the
fungus was isolated.
Holotype: MFLU 17-1273
Saprobic on dead stem of Aquilegia viscosa. Sexual
morph Ascomata 150–195 lm high 9 95–195 lm diam.
(
x = 167.8 9 142.2 lm, n = 5), superficial, solitary, scattered, uniloculate, globose to subglobose, dark brown to
black, conspicuous on the host. Ostioles 50–60 9 45–
60 lm (
x = 53.5 9 52.5 lm, n = 5), protruding from the
centre of the ascomata, without setae. Peridium 40–80 lm
wide, comprising several layers of brown to dark brown
cells of textura angularis, outer layer darkened cells and
inner layer pale brown to hyaline cells. Hamathecium
comprising numerous, filiform, septate, hyaline pseudoparaphyses. Asci 70–100 9 7–11 lm (
x = 83 9 9 lm,
n = 20), 8-spored, bitunicate, short pedicellate with clubshaped pedicel, or sessile, apically rounded. Ascospores
20–30 9 4.5–5.5 lm (
x = 25 9 5 lm, n = 20), overlapping 1–2-seriate, initially hyaline, becoming yellowish
brown at maturity, fusiform, ellipsoidal to subcylindrical,
slightly curved, upper part shorter and wider than the lower
part, 4-transversely septate, constricted at the middle septum, ends remaining lighter and cone-shaped, with narrowly rounded ends, wall firm, smooth-walled, sometimes
guttulate, with appendages (3–5 lm long, 3–4 lm wide) at
both ends. Asexual morph: undetermined.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Passo la Calla - Santa Sofia, on a dead aerial stem of
Aquilegia viscosa Gouan (Ranunculaceae), 14 July 2017,
Erio Camporesi, IT3398 (MFLU 17-1273, holotype).
GenBank numbers: ITS: MK579853, LSU: MK579854,
SSU: MK579855, TEF1-a: MK579856.
Notes: Nodulosphaeria aquilegiae was found on dead,
aerial stems of Aquilegia viscosa in Italy. Ascomatal
characteristics resemble N. multiseptata and N. sambuci in
having superficial, solitary, uniloculate, globose to subglobose ascomata (Tibpromma et al. 2017). This is the first
record of a Nodulosphaeria species on Aquilegia viscosa.
Nodulosphaeria thalictri D. Pem, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555433; Facesoffungi
number: FoF05110; Fig. 35
Etymology: Name reflects the host from which the fungus was isolated.
Holotype: MFLU 17-1241
Saprobic on dead stems of Thalictrum sp. Sexual
morph Ascomata 217–305 lm diam. 9 185–304 lm high
(
x = 263.7 9 217.9 lm, n = 10), immersed or semi-immersed, solitary or scattered, subglobose to obpyriform,
dark brown to black, papillate. Ostiole 70–75 lm diam.
(
x = 71.2 lm, n = 10), papillate, protruding from substratum with numerous internal brown to dark brown cells of
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textura globulosa. Peridium 12–16 lm wide, comprising
4–5 layers of dark brown cells of textura angularis, flattened at the inside. Hamathecium comprising 1.5–2.5 lm
wide, filiform, septate, branching pseudoparaphyses. Asci
77–100 9 7–12 lm (
x = 84.1 9 10.1 lm, n = 10), 8spored, bitunicate, cylindric-clavate, slightly curved, short
pedicellate, apically rounded, with an ocular chamber.
Ascospores
29–35 9 4–5 lm
(
x = 32.3 9 4.5 lm,
n = 20), overlapping 3–4 seriate, hyaline when immature,
becoming pale yellowish to greenish at maturity, long
fusiform, 7–8 septate, straight or slightly curved, thick and
smooth-walled, with terminal appendages. Asexual morph
Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 24 h. at 16 C and germ tubes produced from
middle septa. Colonies growing on MEA, reaching 1 cm
diam. in 1 week. Mycelium superficial, medium sparse,
irregular, flat, smooth, effuse, slightly irregular margin,
from above whitish, reverse yellowish to pale brown.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Lago Pontini – Bagno di Romagna, on dead aerial
stems of Thalictrum sp. (Ranunculaceae), 5 July 2017, E.
Camporesi (MFLU 17-1241, holotype; HKAS 97500,
isotype), ex-type living culture (MFLUCC 18-1138).
GenBank numbers: ITS: MK 101000, LSU: MK
100998, SSU: MK 101001, TEF1-a: MK128971.
Notes: Nodulosphaeria thalictri is similar to N. hirta
(type species of Nodulosphaeria) in having cylindric-clavate, slightly curved, short pedicellate asci and a peridium
with dark brown cells of textura angularis, however N.
thalictri has narrower pseudoparaphyses (1.5–2.5 lm v/s
2.5–3.5 lm),
shorter
and
wider
ascospores
(
x = 32.3 9 4.5 lm v/s x = 55 9 5 lm) and the presence
of terminal appendages (Phookamsak et al. 2014). Our
phylogenetic analyses place N. thalictri close to N. aconiti
(MFLUCC 13–0728). Nodulosphaeria thalictri is similar to
N. aconiti in having a peridium composed of cells of textura angularis and branching pseudoparaphyses (Mapook
et al. 2016). However, N. thalictri has longer asci
(84 9 10 lm v/s 69 9 9 lm), more septa in the ascospores septation (7–8 v/s 4–septate), has long fusiform
ascospores rather than cylindric-fusiform and possesses
terminal appendages, which are lacking in N. aconiti. Nodulosphaeria thalictri is unique in having an ostiole with
numerous internal brown to dark brown cells of textura
globulosa, protruding externally, a feature not recorded in
other Nodulosphaeria species.
Phaeopoacea Thambug., Dissan. & K.D. Hyde
Notes: Phaeopoacea was introduced by Thambugala
et al. (2017b) to accommodate the Phaeopoacea species
occurring as a saprobe on Poaceae (Hyde et al. 2017b;
Thambugala et al. 2017b). Currently, the genus comprises
Fungal Diversity (2019) 96:1–242
57
Fig. 35 Nodulosphaeria thalictri (MFLU 17-1241, holotype). a, b
Appearance of ascomata on host surface. c Section through an
ascoma. d Ostiole. e Peridium. f Hamathecium. g-i Asci. j-m
Ascospores, arrow showing terminal appendages. n Germinated
ascospore on MEA after 24 h. o, p Culture characters (o from above
view, p from below view). Scale bars: a, b = 500 lm, c = 50 lm,
d = 30 lm, g–i = 25 lm, e, n = 20 lm, f, j–m = 10 lm
three species: P. festucae (type species), P. phragmiticola
and P. muriformis. Sexual morphs of Phaepoacea are
characterized by subepidermal ascomata with a central
papillate ostiole, bitunicate asci and ascospores tapering
towards the ends (Shoemaker and Babcock 1989; Hyde
et al. 2017b). The asexual morphs from this genus have
pycnidial conidiomata with brown conidia (Thambugala
et al. 2017b). In this study, we introduce a novel species,
Phaeopoacea asparagicola from Asparagaceae (Fig. 39).
globose to compressed globose, coriaceous, dark brown to
black, rough-walled, with short hyphae projecting from
peridium, ostiolate. Ostiole centrally located, papillate,
without periphysoids. Peridium 11–29 lm wide, composed
of 4–7 layers of reddish brown to dark brown cells of
textura angularis, inner layer composed of hyaline gelatinous cells. Hamathecium composed of numerous, dense,
long, 2–7 lm wide (
x = 5 lm, n = 50), filiform, transversely septate, branched, anastomosing, cellular pseudoparaphyses.
Asci
67–129 9 12–16 lm
(
x = 98 9 14 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate to broad cylindrical, with furcate
pedicel, with ocular chamber visible when immature. Ascospores 22–33 9 6–10 lm (
x = 27 9 8 lm, n = 50), biseriate or overlapping, initially hyaline, becoming yellowish to brown at maturity, broad fusiform, narrow
towards the apex, 3–7-transversely euseptate, constricted at
the septa, with cell above central septum wider, smoothwalled, indentations present, surrounded by a 7–9 lm
wide, mucilaginous sheath. Asexual morph Undetermined.
Phaeopoacea asparagicola Phukhams., Akulov & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555501; Facesoffungi
number: FoF04826; Fig. 36
Etymology: Name refers to the host plant, Asparagus sp.
Holotype: MFLU 18-1380
Saprobic on dead stems of Asparagus sp. Sexual morph
Ascomata 200–365 9 110–314 lm (
x = 283 9 220 lm,
n = 10), semi-immersed to immersed under epidermal tissue, solitary or sometimes gregarious, confluent, scattered,
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b Fig. 36 Phaeopoacea asparagicola (MFLU 18-1380, holotype).
a Appearance of ascomata on host surface. b Close up of ascomata.
c, d Vertical sections of ascomata. e Section of partial peridium layer.
f Cellular pseudoparaphyses. g–i Development state of asci. j–o
Development state of ascospores. p Ascospores stained with india ink,
sheath surrounding the entire ascospore. Scale bar: b = 500 lm, c,
d = 100 lm, e–i = 50 lm, j–p = 10 lm
Culture characteristics: Colonies on MEA, reaching
50 mm diam. after 4 weeks at 18 C. Culture olive-green
radiating yellowish toward the edge, with dense mycelia,
circular, flat, dull, fimbriate, radially furrowed, and slightly
covered with white aerial mycelium; reverse black with
radiating cream mycelium.
Material examined: UKRAINE, Odessa region, Lyman
district, Regional landscape park Tiligulskyt, on the overwintered stems of Asparagus sp. (Asparagaceae), 1 May 2014,
A. Akulov, EX CWU (MYC) AS 5825 (MFLU 18-1380,
holotype), ex-type living culture (MFLUCC 16-0379).
GenBank numbers: ITS: MK443383, LSU: MK443379,
RPB2: MK443387, SSU: MK443381, TEF1-a: MK443385.
Notes: Phaeopoacea asparagicola is introduced as a
new species based on morphological and phylogenetic
evidence. The species formed a moderately-supported
clade with P. festucae and P. phragmiticola (MLBS = 66;
BYPP = 0.99). The morphology of P. asparagicola is
similar to P. phragmiticola in having sub-epidermal, globose, papillate ascomata, cylindrical with short stalk asci,
and conical at the apex with pale brown ascospores
(Shoemaker and Babcock 1989; Hyde et al. 2017b).
Besides these characters, P. asparagicola can be distinguished from P. phragmiticola and P. muriformis by having larger ascomata and asci and ascospores only have
longitudinal septa.
Phaeosphaeria I. Miyake
Notes: Phaeosphaeria was introduced by Miyake (1909)
with P. oryzae I. Miyake as the type species (Phookamsak
et al. 2014). Previously, Phaeosphaeria was considered as a
synonym of Leptosphaeria. However, based on both morphology and molecular analyses, it was considered that
Phaeosphaeria differs from Leptosphaeria in having pseudoparenchymatous peridium, stagonospora-like asexual
morphs and mostly monocotyledonous hosts while Leptosphaeria has scleroplectenchymatous peridium, phoma-like
asexual morphs and mostly dicotyledonous hosts (Schoch
et al. 2009; Zhang et al. 2012). Phaeosphaeria species are
probably cosmopolitan in distribution, since they have been
recorded from both temperate and tropical countries (i.e.
Canada, China, Germany, Japan, Thailand, and Switzerland)
(Shoemaker and Babcock 1989; Phookamsak et al.
2014, 2019). There are 211 epithets for Phaeosphaeria in
59
Index Fungorum (2019). We follow the latest treatment and
updated account of Phaeosphaeria in Phookamsak et al.
(2019). Phaeosphaeria penniseti is introduced as a novel
species based on morphological and phylogenetic evidence
(Figs. 37, 38).
Phaeosphaeria penniseti Karun., C.H. Kuo & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556267; Facesoffungi
number: FoF05849; Fig. 37
Etymology: Name reflects the host genus, Pennisetum.
Holotype: MFLU 18-2674
Saprobic on dead stems of Pennisetum purpureum (Poaceae). Sexual morph Ascomata 133–184 lm high 150–
243 lm diam. (
x = 158.5 9 196.5 lm, n = 5), black,
immersed under host epidermis, visible as raised, to semiimmersed, solitary, subglobose to ampulliform, uni-loculate,
ostiolate, with minute papilla. Peridium 18–20 lm wide at the
sides, comprising 3–5 layers, of brown to dark brown pseudoparenchymatous cells, of textura angularis, hyaline to pale
brown, flattened towards the inner layers. Hamathecium
comprising numerous, filamentous, branched, septate pseudoparaphyses. Asci 55–70 9 10–15 lm (
x = 62.5 9 12.5
lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindricclavate, pedicellate, rounded at the apex and with a shallow
ocular chamber. Ascospores 26–29 9 4–5 lm (
x = 28 9 5
lm, n = 40), overlapping, 2-seriate, hyaline, becoming brown
at maturity, narrowly fusiform, 5-septate, slightly wider at the
second cell from above, guttulate at the beginning, rounded at
the ends. Asexual morph: Undetermined.
Material examined: TAIWAN, Chia Yi Province,
Kwang Hwa, Pennisetum purpureum (Poaceae), 18 March
2018, A. Karunarathna AKTW 24 (MFLU 18-2674,
holotype), ex-type living culture (FU31020).
GenBank numbers: ITS: MK503819, LSU: MK503825,
SSU: MK503831.
Notes: In the NCBI BLAST search, Phaeosphaeria
penniseti shows higher percentage identity to P. oryzae
(99.80%). Phylogenetically, Phaeosphaeria penniseti
forms a sister lineage with P. oryzae, the type species of the
Phaeosphaeria with moderate support in maximum likelihood analysis (89% ML, Fig. 38). It differs from P. oryzae
in having 5-septate ascospores, whereas, P. oryzae has
3-septate ascospores (Phookamsak et al. 2014).
Pseudoberkleasmiaceae Phukhams & K.D. Hyde, fam.
nov.
Index Fungorum number: IF555489, Facesoffungi
number: FoF05311
Saprobic on decaying wood. Sexual morph Undetermined. Asexual morph Hyphomycetous, dictyosporous.
Colonies on natural substratum sporodochia, superficial,
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Fig. 37 Phaeosphaeria penniseti (MFLU 18-2674, holotype). a, b
Appearance of ascomata on the host (Pennisetum purpureum,
Poaceae). c Section through ascoma. d Section through ostiole.
e Section through peridium. f Pseudoparaphyses (stained in cotton
blue). g–i Different developing stages of the asci. j–m Ascospores.
Scale bars: c, d = 50 lm, e = 25 lm, f–m = 10 lm
compact, scattered, irregular, dark-brown to black, glistening. Mycelium immersed in the substrate, septate, branched.
Conidiophores micronematous, mononematous, reduced,
hyaline. Conidiogenous cells holoblastic, monoblastic, integrated, terminal, determinate. Conidia acrogenous, solitary,
broadly ellipsoidal to obovoid, muriform, guttulate, smoothwalled, brown, olivaceous green, with or without guttules,
usually with conidiogenous cell attached.
Type genus: Pseudoberkleasmium pandanicola Tibpromma & K.D. Hyde, Fungal Diversity: 52 (2018)
Notes: Pseudoberkleasmiaceae is introduced to accommodate a berkleasmium-like hyphomycete that is phylogenetically placed in the order Pleosporales. The family
comprises Pseudoberkleasmium chiangmaiense and P.
pandanicola, the latter as the generic type. Berkleasmium is
polyphyletic, with species having similar characteristics,
but which are phylogenetically distinct (Pinnoi et al. 2007;
Hu et al. 2010; Lu et al. 2018b). A study by Tanney and
Miller (2017) was able to obtained several fresh collections
of Berkleasmium and compared those with the reference
specimens of Berkleasmium concinnum. Re-examination of
the type species placed the generic type of Berkleasmium in
Tubeufiales. In this study, Pseudoberkleasmiaceae is
Fig. 38 Phylogram generated from maximum likelihood analysis c
based on combined ITS, LSU, SSU and TEF1-a sequence data for
genera in Phaeosphaeriaceae. Related sequences are taken from
Wanasinghe et al. (2018a) and Phookamsak et al. (2019). Onehundred and sixty-one strains are included in the combined analyses
which comprise 3307 characters (419 characters for ITS, 824
characters for LSU, 990 characters for SSU, 1074 characters for
TEF1-a) after alignment. Staurosphaeria rhamnicola (MFLUCC
17-0813, MFLUCC 17-0814) (Montagnulaceae, Pleosporales) are
used as the outgroup taxa. Single gene analyses are carried out and the
topology of each tree had clade stability. Tree topology of the
maximum likelihood analysis is similar to the maximum parsimony
analysis and the Bayesian analysis. The best RaxML tree with a final
likelihood value of - 29921.960589 is presented. The matrix had
1190 distinct alignment patterns, with 23.56% undetermined characters or gaps. Estimated base frequencies were as follows:
A = 0.243950, C = 0.233656, G = 0.265190, T = 0.25720; substitution rates AC = 1.334366, AG = 3.046037, AT = 2.327147, CG =
0.800430, CT = 6.652827, GT = 1.000000; gamma distribution
shape parameter a = 0.527656. Maximum parsimony analysis
resulted 3276 constant characters and 629 informative characters.
Bootstrap values for maximum likelihood (ML) and maximum
parsimony (MP) equal to or greater than 65% are placed above and
below the branches respectively. Branches with Bayesian posterior
probabilities (BYPP) equal or greater than 0.90 are in bold. The extype strains are in bold and black. The newly generated sequences are
indicated in bold and blue
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61
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62
related to the family Hermatomytaceae with relatively
strong statistical support.
Pseudoberkleasmium Tibpromma & K.D. Hyde
Notes: The monotypic genus Pseudoberkleasmium was
introduced by Tibpromma et al. (2018) with P. pandanicola Tibpromma & K.D. Hyde as the type species. Pseudoberkleasmium is characterized by hyaline subglobose
conidiogenous cells and acrogenous, broadly ellipsoidal to
obovoid, muriform, brown or olivaceous green, and guttulate conidia. In this paper, we introduce a new Pseudoberkleasmium species, P. chiangmaiense, based on
phylogenetic and morphological evidence.
Pseudoberkleasmium chiangmaiense Y.Z. Lu & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555595; Facesoffungi
number: FoF05310; Fig. 39
Etymology: chiangmaiense, referring to collecting site.
Holotype: MFLU 17-1118
Saprobic on decaying wood. Sexual morph Undetermined Asexual morph Hyphomycetous, dictyosporous.
Colonies on natural substratum sporodochia, superficial,
compact, scattered, irregular, black, glistening. Mycelium
immersed in the substrate, composed of septate, branched,
smooth, thin-walled, hyaline to pale brown, 2 lm wide
hyphae. Conidiophores micronematous, mononematous,
reduced, hyaline, smooth-walled. Conidiogenous cells 12–
18 9 12–18 lm (
x = 15 9 14 lm, n = 20), holoblastic,
monoblastic, integrated, terminal, determinate, subglobose,
with guttulate, hyaline. Conidia 30–35 9 15–20 lm
Fig. 38 continued
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(
x = 33 9 18 lm, n = 20), acrogenous, solitary, broadly
ellipsoidal to obovoid, flattened, muriform, guttulate,
smooth-walled, brown, usually with conidiogenous cell
attached.
Culture characteristics: Conidia germinating on water
agar and germ tubes produced from conidia within 12 h.
Colonies growing on PDA, circular, with flat surface, edge
entire, reaching 28 mm in 3 weeks at 28 C, white to pale
brown in PDA medium. Mycelium superficial and partially
immersed, branched, septate, hyaline to pale brown,
smooth.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng District, Mushroom Research Center, on
decaying wood, 25 January 2017, Yong-Zhong Lu, MRC
23 (MFLU 17-1118, holotype), ex-type living culture
(MFLUCC 17-1809).
GenBank numbers: ITS: MK131259, LSU: MK131260,
TEF1-a: MK131261.
Notes: Pseudoberkleasmium chiangmaiense is similar to
P. pandanicola in having hyaline subglobose conidiogenous cells and acrogenous, broadly ellipsoidal to obovoid, muriform, brown and guttulate conidia, but differs by
its larger conidiogenous cells (12–18 9 12–18 lm vs
5–11 9 9–12 lm). Phylogenetically, Pseudoberkleasmium
chiangmaiense shares a sister relationship to P. pandanicola with good bootstrap support (100% ML/1.00 BYPP)
(Fig. 40); the phylogeny also indicates that they are distinct
species.
Fungal Diversity (2019) 96:1–242
63
Fig. 39 Pseudoberkleasmium chiangmaiense (MFLU 17-1118, holotype). a Colonies on substrate. b–j Conidia. k Germinating conidium. l, m
Colony on PDA (l from above view, m from below view). Scale bars: a = 200 lm, b–k = 20 lm, l, m = 20 mm
Pyrenochaetopsidaceae Valenzuela-Lopez, Crous, Cano,
Guarro & Stchigel
Notes: ‘Cucurbitariaceae’ was revealed to represent five
distinct
families,
i.e.
the
newly
erected
Neopyrenochaetaceae, Parapyrenochaetaceae, Pseudopyrenochaetaceae and Pyrenochaetopsidaceae, and Cucurbitariaceae sensu stricto with a narrowed concept
(Valenzuela-Lopez et al. 2018). Pyrenochaetopsis is the
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Fungal Diversity (2019) 96:1–242
b Fig. 40 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, ITS, TEF1-a, and RPB2 sequence
data representing Pseudoberkleasmiaceae and the closely related
families in Pleosporales order. Related sequences are taken from
Tibpromma et al. (2018). Sixty-four strains are included in the
combined analyses which comprise 4340 characters (842 characters
for LSU, 1030 characters for SSU, 512 characters for ITS, 926
characters for TEF1-a, 1030 characters for RPB2) after alignment.
Lophiostoma crenatum (CBS 629.86) and L. arundinis (CBS 621.86)
in Lophiostomataceae (Pleosporales) are used as the outgroup taxa.
Single gene analyses were also performed to compare the topology
and clade stability with combined gene analyses. Tree topology of the
maximum likelihood analysis is similar to the Bayesian analysis. The
best RaxML tree with a final likelihood values of - 38637.764110 is
presented. The matrix had 1968 distinct alignment patterns, with
33.91% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.245895, C = 0.251442, G = 0.271420,
T = 0.231244; substitution rates AC = 1.592217, AG = 3.496391,
AT = 1.418709, CG = 1.209916, CT = 8.405355, GT = 1.000000;
gamma distribution shape parameter a = 0.569582. Bootstrap values
for maximum likelihood (ML) equal to or greater than 70% and clade
credibility values greater than 0.90 (the rounding of values to 2
decimal proportions) from Bayesian-inference analysis labelled on the
nodes. The ex-type strains are in bold and black. The newly generated
sequence is indicated in bold and blue
type genus of Pyrenochaetopsidaceae. The general characters of the family Pyrenochaetopsidaceae included
glabrous or setose pycnidia with apapillate or papillate
ostiolate necks, phialidic conidiogenous cells and aseptate,
cylindrical to allantoid conidia. Members of
Pyrenochaetopsidaceae are saprobes from various niches,
such as water, plants, soil and air samples (de Gruyter et al.
2010, 2013; Valenzuela-Lopez et al. 2018). Some species
e.g. Pyrenochaetopsis poae, have been repeatedly isolated
as endophytes of graminaceous plants (Crous et al. 2014).
Other species such as P. globosa, P. paucisetosa and
Neopyrenochaetopsis hominis, have been isolated from
sinusitis, toe nail and skin tissue of humans as weak
pathogens (Valenzuela-Lopez et al. 2018).
Pyrenochaetopsis Gruyter, Aveskamp & Verkley
Notes: Pyrenochaetopsis was erected by de Gruyter
et al. (2010) with P. leptospora (Sacc. & Briard) Gruyter,
Aveskamp & Verkley as the type species (de Gruyter et al.
2010). The genus is characterized by setose pycnidia with
apapillate or papillate ostioles, and simple, aseptate,
cylindrical to allantoid conidia.
Pyrenochaetopsis sinensis G.S. Li, J.M. Liang & L. Cai,
sp. nov.
Index Fungorum number: IF556011; Facesoffungi
number: FoF05965; Fig. 41
Etymology: Named after the country where the type was
collected.
Holotype: HMAS 248045
65
Saprobic on rhizosphere soil of turfgrasses. Sexual
morph Undetermined. Asexual morph Hyphae pale white,
smooth- and thin-walled, septate, 2–5 lm wide. Conidiomata pycnidial, pale brown, solitary or confluent,
immersed (OA), subglobose or globose, 100–180 9 100–
155 lm, with apapillate ostiolar neck. Pycnosclerotia
forming after 25 days culturing on OA, brown to dark
brown, globose or subglobose, 55–60 lm in diam. Setae
absent. Pycnidial wall of textura angularis, 3–5 layered,
15–25 lm thick, composed of brown, flattened polygonal
cells of 3–6 lm diam. Conidiogenous cells phialidic,
hyaline,
smooth-walled,
subglobose,
3–6 9 2–5
(
x = 4 9 3, n = 30) lm. Conidia aseptate, hyaline,
smooth- and thin-walled, cylindrical to allantoid, 3–
4.5 9 1–2 (
x = 4 9 2, n = 40) lm, usually with two
guttules.
Culture characteristics: Colonies on OA reaching
23 mm diam. after 7 d at 25 ± 1 C, flattened, greenish
grey (26B2); reverse olive grey (26E3). Colonies on MEA
reaching 19 mm diam. after 7 d at 25 ± 1 C, floccose,
pale grey (26B1) to olive (26C2); reverse olive (26F3).
NaOH spot test negative. Crystals absent.
Material examined: CHINA, Beijing, in rhizosphere soil
of Poa pratensis (Poaceae), 26 August 2017, J.M. Liang
(HMAS 248045, holotype), ex-type living culture
CGMCC 3.19296 = LC12199; Paratypes: China. Beijing,
in rhizosphere soil of Digitaria sanguinalis (Poaceae), 21
August 2017, J.M. Liang LC12200); China, Beijing, in
rhizosphere soil of Poa pratensis (Poaceae), 21 August
2017, J.M. Liang LC12197; ibid. LC12198.
GenBank numbers: ITS: MK348586, MK348587,
MK348584, MK348584, LSU: MK348581, MK348580,
MK348582, MK348583, RPB2: MK355077, MK355078,
TUB2: MK348221, MK348220, MK348222, MK348223.
Notes: Pyrenochaetopsis sinensis is phylogenetically
distinct from other species in this genus, clustering sister to
P. microspore (Fig. 42). Pyrenochaetopsis sinensis is
easily distinguished from P. microspora by the absence of
setae covering on the surface of pycidium (Gruyter and
Boerema 2002). In addition, P. sinensis is the only species
of Pyrenochaetopsis producing pycnosclerotia.
Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray.
Notes: The family Tetraplosphaeriaceae accommodates
seven genera viz. Ernakulamia Subram. (Subramanian
1994), Polyplosphaeria Kaz. Tanaka & K. Hiray. (Tanaka
et al. 2009), Pseudotetraploa Kaz. Tanaka & K. Hiray.
(Tanaka et al. 2009), Quadricrura Kaz. Tanaka, K. Hiray.
& Sat. Hatak. (Tanaka et al. 2009), Shrungabeeja V.G. Rao
& K.A. Reddy (Rao and Reddy 1981), Tetraploa Berk. &
Broome, and Triplosphaeria Kaz. Tanaka & K. Hiray.
(Tanaka et al. 2009). Ernakulamia is placed in this family
based on morphological and phylogenetic analyses
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Fig. 41 Pyrenochaetopsis sinensis (HMAS 248045, holotype). a–
c Colony on OA, MEA and SNA (front and reverse). d Pycnidia
forming on OA. e Pre-pycnidial structures. f Pycnidia. g, h Pycnidia
wall. i1–i5 Conidiogenous cell. j Conidia. k Pycnosclerotium on OA.
l–m Pycnosclerotium. Scale bars: e = 50 lm, g = 20 lm (apply to l),
h = 10 lm (apply to i, j, m, i1–i5)
(Delgado et al. 2017). Although Pseudotetraploa and
Shrungabeeja were not included in this family by
Wijayawardene et al. (2018a), these genera should be
retained. Pseudotetraploa is distinct from Tetraploa by its
obpyriform to long obpyriform, pseudoseptate conidia
composed of 4 to 8 columns with 4 (rarely 6 to 8) setose
appendages (Hyde et al. 2013). Pseudotetraploa also
formed an independent clade in the molecular analysis and
distinguished from Tetraploa (Tanaka et al. 2009).
Shrungabeeja was identified as a distinctive genus within
Tetraplosphaeriaceae in Ariyawansa et al. (2015). Tetraploa, Polyplosphaeria and Triplosphaeria are characterized with almost hyaline 1(–3)-septate ascospores as
Massarina-like sexual morphs and several setose
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67
Fig. 42 Phylogram generated from maximum likelihood (ML) analysis inferred from a four-locus concatenated alignment (ITS, LSU,
RPB2 and TUB2) for Pyrenochaetopsis species and closely related
species. Related sequences are taken from Valenzuela-Lopez et al.
(2018) and Crous et al. (2014). Twenty-two strains are included in the
multi-locus analyses which comprise a total of 2454 characters (443
characters for ITS, 817 characters for LSU, 297 characters for TUB2
and 897 characters for RPB2) after alignment. Neopyrenochaetopsis
hominis (CBS 143033) (Pyrenochaetopsidaceae, Pleosporales) is used
as the outgroup taxon. Tree topology of the ML analysis is similar to
that of the Bayesian analysis. The best RAxML tree with a final
likelihood value of - 8929.890374 is presented. The matrix had 573
variable sites. Estimated base frequencies were as follows:
A = 0.2382, C = 0.2531, G = 0.2663, T = 0.2423; substitution rates
AC = 1.4907, AG = 4.6367, AT = 1.4907, CG = 1.0000, CT =
10.5414, GT = 1.0000; gamma distribution shape parameter
a = 0.5740. Bootstrap values (1000 replicates) over 75% for ML
and Bayesian posterior probability (PP) over 0.95 are marked at the
left of every node (ML/BYPP). Type strains are in bold and the newly
generated sequences are indicated in blue
appendages as tetraploa-like asexual morphs (Tanaka et al.
2009; Hyde et al. 2013). The sexual morphs of the genera
Ernakulamia,
Pseudotetraploa, Shrungabeeja and
Quadricrura are unknown. Most taxa of Tetraplosphaeriaceae were found on bamboo and grasses.
are 20 epithets listed under the genus in Index Fungorum
(2019). However, only five species of Tetraploa have
molecular data (NCBI: www.ncbi.nlm.nih.gov; accessed
1st May 2019). Tetraploa has relatively small globose
ascomata, narrowly fusiform ascospores having appendage-like sheath, and conidia with four setose appendages
and is mainly known from monocotyledons especially
grasses and bamboo (Tanaka et al. 2009; Goonasekara
et al. 2018; Farr and Rossman 2019). A new record of
Tetraploa nagasakiensis from China is reported in this
paper.
Tetraploa Berk. & Broome
Notes: Hyde et al. (2013) and Wijayawardene et al.
(2014) recommended the use of Tetraploa over Tetraplosphaeria based on the close phylogenetic relationships between the type species of these two genera and
there are more species accommodated in Tetraploa, having
priority of the most named uses. Thus, Rossman et al.
(2015b) adopted Tetraploa over Tetraplosphaeria. There
Tetraploa nagasakiensis (Kaz. Tanaka & K. Hirayama)
Kaz. Tanaka & K. Hirayama
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Facesoffungi number: FoF05080; Fig. 43
Basionym: Tetraplosphaeria nagasakiensis Kaz. Tanaka
& K. Hiray., in Tanaka et al., Stud. Mycol. 64: 180 (2009)
Saprobic on dead bamboo branches. Sexual morph
Ascomata 170–200 lm high, 190–260 lm diam.,
immersed under host epidermis, solitary, scattered, visible
as small minute black spots or papilla on host issue, dark
Fig. 43 Tetraploa nagasakiensis (HKAS 101756, new geographical
record). a–c Host and ascomata. d Vertical section of ascoma.
e Peridium of ascoma. f Pseudoparaphyses. g–k Asci. l Culture from
above and below views. m, n Ascospores. o Germinating ascospore.
Scale bars: i–k, o = 20 lm, e, g, h = 15 lm, f, m, n = 10 lm
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brown to black, uni-loculate, globose to subglobose or
ampulliform, coriaceous, with ostiole. Peridium 17–24 lm
wide, comprising several layers, with outers layers composed of dark brown to black cells of textura angularis,
inner layers composed of light brown to hyaline pseudoparenchymatous cells arranged in textura angularis. Hamathecium of 1.7–2.4 lm, septate, branched, broad
pseudoparaphyses, anastomosing near the apex. Asci 98–
130 9 14–18 lm (
x = 114 9 16 lm, n = 20), 8-spored,
bitunicate, fissitunicate, clavate, pedicellate with furcate
ends, apically rounded with well-developed ocular chamber. Ascospores 30–34.5 9 4–5 lm (
x = 32.3 9 4.6 lm,
n = 20), overlapping 2–3-seriate, hyaline, 1-septate, narrowly fusiform, smooth-walled, multi-guttulate. Asexual
morph Refer to Tanaka et al. (2009).
Culture characteristics: Ascospores germinating on
PDA within 24 h. Colonies on PDA reaching 30–35 mm
diam. after 4 weeks at 20–25 C, circular, slightly raised to
umbonate, pale black at the middle, white at the margin
from the above and below, no pigment in agar.
Material examined: CHINA, Yunnan Province, Honghe
Prefecture, Pingbian County, nearby Tuanpo Reservoir, on
dead bamboo branches, 21 September 2017, H.B. Jiang,
Pb001 (HKAS 101756), living culture (KUMCC 18-0109).
GenBank numbers: ITS: MK079890, LSU: MK079891,
SSU: MK079888.
Notes: Although the new collection (HKAS 101756) has
eight different ITS base pairs as compared to the type strain
of Tetraploa nagasakiensis, they are still close to each
other in multi-phylogenetic analysis based on ITS, LSU
and SSU sequenced data (Fig. 44). They are also morphologically very similar. Tetraploa nagasakiensis was
reported only from Japan (Tanaka et al. 2009), but our new
strain was collected in China.
Torulaceae Corda
Notes: The family Torulaceae was introduced by Corda
(Sturm 1829). Two genera, Dendryphion Wallr. and Torula
Pers., are accepted within this family. Su et al. (2016) and
Li et al. (2016) introduced the genera Neotorula Ariyaw.,
Z.L. Luo & K.D. Hyde and Sporidesmioides Jin F. Li,
Phook. & K.D. Hyde within Torulaceae, respectively.
Torula Pers.
Notes: The genus Torula was established by Persoon
(1794), typified by T. herbarum (Pers.) Link as the type
species. Torula is characterized by 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; these conidiogenous cells are consequently termed ‘corona cells’ (Crane and Miller 2016). This genus is
problematic as many species await redescription (Seifert
et al. 2011). More sequence data are required in the future
69
study of the genus. A key to Torula and similar genera was
provided by Crane and Miller (2016).
Torula breviconidiophora C.G. Lin & K.D. Hyde, sp. nov.
Index Fungorum number: IF555428; Facesoffungi
number: FoF05069; Fig. 45
Etymology: In reference to the short conidiophores.
Holotype: MFLU 18-1693
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies effuse on the
natural substrate, scattered, hairy, dark brown. Mycelium
immersed to superficial, composed of pale brown, septate,
branched hyphae. Conidiophores macronematous,
mononematous, subcylindrical, erect, septate, smooth,
straight or flexuous, subhyaline to dark brown, with
ampulliform cells, 3.5–28 lm long, 3–8 lm wide at the
widest part. Conidiogenous cells mono- to polyblastic,
integrated, terminal, on conidiophores, terminal or intercalary in conidial chains, doliiform to ellipsoid, pale
brown, collapsing in above half when old, 4–7 lm long,
3–7 lm wide. Conidia phragmosporous, in branched
chains, acrogenous, dry, cylindrical, medium to dark
brown, with apical cell subhyaline or pale brown, 1–4septate, deeply constricted at septa, verrucose, rounded at
both ends, easily separating, 8–21 lm long (x
¯ = 15.2 lm,
n = 60), 3.5–7 lm wide (x
¯ = 5.2 lm, n = 60).
Culture characteristics: Conidia germinating on PDA
within 12 h. Colonies on PDA effuse, greyish brown to
dark brown, reaching a diam. of 4-6 cm in 10 days at
25 C.
Material examined: THAILAND, Chang Rai Province,
Muang District, Mae Fah Luang University, near S7
building, on decaying wood, 20 July 2016, Chuangen Lin,
ML 1-4 (MFLU 18-1693, holotype; HKAS 102196, isotype), ex-type living culture (KUMCC 18-0130).
GenBank numbers: ITS: MK071670, LSU: MK071672,
SSU: MK071697, TEF1-a: MK077673.
Notes: This species is most similar to T. mackenziei
which is characterized by greyish brown conidia composed
of moniliform cells and 2–3 septa (Li et al. 2017). However, the conidia of T. breviconidiophora are medium to
dark brown, with 1–4-septate, subhyaline or pale brown
apical cells. From the phylogenetic trees of Bayesian
inference, maximum parsimony and maximum likelihood
analyses based on combined ITS, LSU, SSU and TEF1-a,
this species is closest to T. chromolaenae and T. mackenziei (Fig. 47). Torula chromolaenae is distinct from other
Torula species in having smaller and doliiform conidiogenous cells and 2–3-septate conidia (Li et al. 2017).
Torula breviconidiophora differs from T. chromolaenae by
its longer conidiophores (3.5–28 lm vs 5–6.3 lm).
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70
Fig. 44 Phylogram generated from maximum likelihood analysis
based on a combined ITS, LSU, and SSU sequence dataset for
Tetraploa species and several closely related genera in
Tetraplosphaeriaceae. Related sequences are taken from Tanaka
et al. (2009) and Ariyawansa et al. (2015). Twenty-nine strains are
included in the combined genes sequence analyses which comprise
total 3225 characters including gaps (584 characters for ITS, 1300
characters for LSU, 1341 characters for SSU) after alignment.
Lophiotrema neoarundinaria KT 856 (Lophiotremataceae, Pleosporales) is used as the outgroup taxon. Single gene analyses are carried
out and the topology of each tree had clade stability. Tree topology of
the maximum likelihood analysis is similar to the Bayesian analysis.
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Fungal Diversity (2019) 96:1–242
The best sorting RaxML tree with a final likelihood value of
- 8716.750658 is presented. The matrix had 430 distinct alignment
patterns, with 15.62% undetermined characters or gaps. Estimated
base frequencies were as follows: A = 0.251593, C = 0.234741,
G = 0.277315, T = 0.236350; substitution rates AC = 4.295630,
AG = 4.189995, AT = 2.538258, CG = 1.455605, CT = 14.145092,
GT = 1.000000; gamma distribution shape parameter a = 0.501945.
Bootstrap values for maximum likelihood (ML) equal to or greater
than 70% are shown on the left. Branches with Bayesian posterior
probabilities (BYPP) equal or greater than 0.90 are shown on the
right. The ex-type strains are in bold and black. The newly generated
sequence is indicated in blue
Fungal Diversity (2019) 96:1–242
71
Fig. 45 Torula breviconidiophora (MFLU 18-1693, holotype). a, b Host material. b Conidiophores on leaf surface. c, d Conidiophores,
conidiogenous cells and conidia. e–g Conidia. Scale bars: c–g = 10 lm
Torula polyseptata C.G. Lin & K.D. Hyde, sp. nov.
Index Fungorum number: IF555429, Facesoffungi
number: FoF05070; Fig. 46
Etymology: In reference to the 2–8-septate conidia.
Holotype: MFLU 18-1694
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies effuse on the
natural substrate, scattered, hairy, dark brown. Mycelium
immersed to superficial, composed of pale brown, septate,
branched hyphae. Conidiophores macronematous,
mononematous, subcylindrical, erect, septate, smooth,
straight or slightly flexuous, dark brown to black, 10–
40 lm long, 3.5–8 lm wide at the widest part. Conidiogenous cells holoblastic, mono- to polyblastic, integrated,
terminal or intercalary in conidial chains, doliiform, pale
brown, collapsing in above half when old, 4.5–8.5 lm
long, 4.5–8 lm wide. Conidia phragmosporous, in branched chains, acrogenous, dry, cylindrical, medium to dark
brown, 2–8-septate, deeply constricted at septa, verrucose,
rounded at both ends, easily separating, 10–40 lm long
(
x = 19.3 lm, n = 35), 3.5–7.5 lm wide (
x = 5.5 lm,
n = 35).
Culture characteristics: Conidia germinating on PDA
within 12 h. Colonies on PDA effuse, greyish brown to
dark brown, reaching a diam. of 4–6 cm in 10 days at
25 C.
Material examined: THAILAND, Chang Rai Province,
Muang District, Mae Fah Luang University, on decaying
wood, 20 July 2016, Chuangen Lin, ML 21-2 (MFLU
18-1694, holotype; HKAS 102197, isotype), ex-type living
culture (KUMCC 18-0131).
GenBank numbers: ITS: MK071671, LSU: MK071673,
SSU: MK071698, TEF1-a: MK077674.
Notes: This species is most similar to Torula pluriseptata in their conidial morphology, however, the conidiophores of T. polyseptata are longer than T. pluriseptata
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Fig. 46 Torula polyseptata (MFLU 18-1694, holotype). a, b Host material. b Conidiophores on leaf surface. c–e Conidiophores, conidiogenous
cells and conidia. f–j Conidia. Scale bars: c = 20 lm, d–j = 10 lm
(10–40 lm vs 2.8–4.3 lm). Torula polyseptata forms a
clade together with strains of T. chiangmaiensis with 98%
MP bootstrap support, 97% ML bootstrap support and
0.97% Bayesian posterior probabilities within the genus
Torula (Fig. 47). Torula polyseptata is distinct from T.
chiangmaiensis by the size and number of septa in the
conidia.
The ex-type strains are in bold and black. The newly
generated sequences are indicated in bold and blue. The
tree is rooted with Dendryphion nanum (HKAS 84010).
Trematosphaeriaceae K.D. Hyde, Y. Zhang ter, Suetrong
& E.B.G. Jones
Notes: Trematosphaeriaceae was introduced by Suetrong et al. (2011b) to accommodate the genera
123
Falciformispora K.D. Hyde, Halomassarina Suetrong et al.
and Trematosphaeria Fuckel. These genera mainly comprise fungi inhabiting mangrove wood in marine environments and wood in freshwater (Hyde 1989; Suetrong et al.
2011b). The main distinguishing characters of this family
are medium-sized rounded ascomata with a papillate ostiole, a relatively wide, coriaceous peridium, cellular pseudoparaphyses and cylindro-clavate asci. The ascospores are
two-celled or many celled, hyaline or brown. We introduce
Falciformispora aquatica as a new species based on evidence from morphology and phylogenetic analyses of
combined LSU, SSU, RPB2 and TEF1-a sequence data
(Fig. 49).
Fungal Diversity (2019) 96:1–242
73
Fig. 47 Phylogenetic tree generated from ML analysis based on
combined ITS, LSU, SSU and TEF1-a sequence data for the genus
Torula. Related sequences are taken from Crane and Miller (2016),
Su et al. (2016) and Li et al. (2017). Twenty-eight strains are included
in the combined analyses which comprise 3139 characters (552
characters for ITS, 844 characters for LSU, 887 characters for SSU,
856 characters for TEF1-a) after alignment. The best RaxML tree
with a final likelihood value of - 7637.920840 is presented. The
matrix had 454 distinct alignment patterns, with 31.43% undetermined characters or gaps. Estimated base frequencies were as follows:
A = 0.242476, C = 0.262534, G = 0.274745, T = 0.220245; substitution rates AC = 3.217101, AG = 3.132488, AT = 2.857294, CG =
1.885404, CT = 12.537654, GT = 1.000000; gamma distribution
shape parameter a = 0.02. Maximum parsimony analysis of 2817
constant characters and 204 informative characters resulted in 126
equally most parsimonious tree of 567 steps (CI = 0.704, RI = 0.785,
RC = 0.552, HI = 0.296). For the Bayesian analysis, two parallel runs
with six chains were run for 1000,000 generations and trees were
sampled every 100th generation, resulted in 20002 trees from two
runs of which 15002 trees were used to calculate the posterior
probabilities (each run resulted in 10001 trees of which 7501 trees
were sampled). Bootstrap support values for maximum parsimony
(MP) and maximum likelihood (ML) greater than 50% and Bayesian
posterior probabilities greater than 0.8 are indicated above or below
the nodes as MLBS/MPBS/BYPP
Falciformispora K.D. Hyde
Notes: Falciformispora was established by Hyde (1992)
as a monotypic genus and was assigned to the Pleosporaceae when compared with Setosphaeria, which has an
Exserohilum asexual morph and is exclusively parasitic on
Gramineae. Subsequently, Raja et al. (2008) collected
Falciformispora species in freshwater in Florida and found
that it is closely related to Chaetomastia rather than Setosphaeria, but is distinct in having hyaline ascospores.
Suetrong et al. (2009) recorded it from the terrestrial oil
palm (Elaeis guineensis) in Thailand. Suetrong et al.
(2011b) showed that Falciformispora forms a well-supported clade with Trematosphaeria pertusa and Halomassarina thalassiae in Trematosphaeriaceae based on
phylogenetic analysis. Therefore, this genus was placed in
Trematosphaeriaceae. Currently, F. lignatilis, F. senegalensis and F. tompkinsii are listed in Index Fungorum
(2019).
Falciformispora aquatica D.F. Bao, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF555415; Facesoffungi
number: FoF05059; Fig. 48
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Fig. 48 Falciformispora aquatica (MFLU 18-1228, holotype). a Ascomata on submerged wood. b Section of ascoma. c, d Section of
peridium. e Pseudoparaphyses. f–k Asci. l-n Ascospores.
o Germinating ascospore. p, q Culture on PDA. Scale bars:
b = 100 lm, f–k = 30 lm, c–e, l–o = 20 lm
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18-1228
Saprobic on decaying wood, submerged wood in
freshwater. Asexual morph Undetermined. Sexual morph
Ascomata 260–330 lm diam. 230–300 lm high
(
x = 286 9 260 lm, n = 10) solitary to gregarious,
immersed in the host tissue, globose to subglobose,
coriaceous, papillate, ostiolate, dark grey to black. Peridium up to 25–30 lm wide, composed of outer thick-walled
angular or rounded greyish brown cells, and inner hyaline
cells of textura prismatica. Hamathecium comprising 2–
3.5 lm wide, numerous, septate, cellular pseudoparaphyses, hyaline, branched, surrounded by a gelatinous matrix.
Asci 115–167 9 28–47 lm (
x = 147 9 36 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindro-clavate, short
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pedicellate.
Ascospores
48–59 9 10–16 lm
(
x = 55 9 13 lm, n = 30), fusiform to clavate, hyaline,
straight or slightly curved, 5–6-septate, mostly 6-septate,
slightly constricted at all septa, tapering to narrow both of
ends, surrounded by a mucilaginous sheath and a single
scythe-like appendage at the apex.
Culture characteristics: Ascospores germinating on
PDA within 24 h. Colonies on MEA at room temperature
reaching 2.5 cm diam. in three weeks, mycelium pale
brown to grayish brown after 3 weeks, composed brown to
dark brown, septate, smooth or verrucose hyphae.
Material examined: Thailand, Prachuap Khiri Khan
Province, Sai Khu Waterfall, on submerged decaying
wood, August 2017, Vinit Kumar., site1–2–1, (MFLU
18–1228, holotype), ex-type living culture (MFLUCC
18–0212).
GenBank numbers: LSU: MK063643, RPB2:
MK099810, TEF1-a: MK099811.
Notes: The phylogenetic analyses showed that Falciformispora aquatica grouped with members of Falciformispora, but separated in an independent lineage with
relatively high bootstrap support (100% ML/1.00 BYPP,
Fig. 49). Falciformispora aquatica is similar to F. lignatilis, as they have similar asci (bitunicate, fissitunicate,
with a short pedicel) and ascospores (fusiform to clavate,
hyaline). However, F. aquatica has larger asci
(115–167 9 28–47 vs. 110–136 9 20–32 lm), and larger
ascospores (48–59 9 10–16 vs. 42–50 9 7.5–10 lm).
Moreover, F. aquatica ascospores are mostly 6-septate,
whereas those of F. lignatilis over 7-septate.
Minutisphaerales Raja, Oberlies, Shearer & A.N. Mill.
Notes: The order Minutisphaerales was introduced by
Raja et al. (2015) based on a LSU, ITS, SSU and MCM7
sequence dataset. It is typified by the sexual genus Minutisphaera with the chemical analysis carried out for M.
aspera Raja, Oberlies, Shearer & A.N. Mill. and M.
parafimbriatispora Raja, Oberlies, Shearer & A.N. Mill.
Jayasiri et al. (2018) included a new family
Acrogenosporaceae in Minutisphaerales.
Acrogenosporaceae Jayasiri & K.D. Hyde
Notes: Acrogenosporaceae is a monotypic family with
Acrogenospora as the type genus. Rossman et al. (2015b)
suggested to protect Acrogenospora over Farlowiella (Art.
59.1). Ellis (1972) reported an Acrogenospora asexual
morph accompanies F. australis Dennis on the host substrate, while Mason (1941) showed the connection between
A. megalospora (Berk. & Broome) Goh, K.D. Hyde &
C.K.M. Tsui and F. armichaeliana (Berk.) Sacc. based on a
cultural study. With seven strains related to this group,
Jayasiri et al. (2018) carried out the phylogenetic analyses
which supported the connection between Acrogenospora
and Farlowiella.
75
Acrogenospora M.B. Ellis
Notes: The holomorph genus Acrogenospora is characterized by laterally compressed hysterothecia with a
prominent sunken slit, aseptate, hyaline pedicellate ascospores, macronematous brown conidiophores, monoblastic,
terminal or intercalary conidiogenous cells and globose,
elliposoid or obovoid, olivaceous to brown conidia. Among
the available sequenced strains of the genus, many do not
have any associated morphological descriptions. We
therefore designate a reference specimen for the type species A. sphaerocephala and introduce a new taxon,
A. thailandica, based on phylogenetic analyses and morphological characters.
Acrogenospora sphaerocephala (Berk. & Broome) M.B.
Ellis, Dematiaceous Hyphomycetes: 114 (1971)
: Monotospora sphaerocephala Berk. & Broome, Ann.
Mag. nat. Hist., Ser. 3 3: 361 (1859)
Index Fungorum number: IF308236; Facesoffungi
number: FoF04687; Fig. 50
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies sparse, scattered,
black, glistening, hairy. Mycelium mostly immersed, consisting of septate, thin-walled, smooth, hyaline to pale
brown hyphae. Conidiophores macronematous, mononematous, (155–)215–320(–360) 9 (3.5–)4.8–7.5(–9.5) lm
(
x = 275 9 6 lm, n = 15), solitary, erect, mostly flexuous,
septate, slightly tapering towards the apex, mid to dark
brown, paler and rounded at the apex, smooth, thick-walled
at the base. Conidiogenous cells monoblastic, integrated,
intercalary, mid brown, cylindrical, with percurrent proliferations, sometimes flexuous at the proliferation. Conidia acrogenous, 18–30 lm (
x = 24 lm, n = 40) diam.,
holoblastic, olive-green to brown, spherical or subspherical, unicellular, smooth and thick-walled, guttulate, truncate at the base.
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced from both ends. Colonies on PDA, reaching 5–10 mm diameter after two weeks
at 25 C in natural light, with dense grayish green aerial
mycelium on the surface, black in reverse with entire
margin.
Material examined: THAILAND, Prachuap Khiri Khan
Province, near 1230.1950 N, 9931.3500 E, on decaying
wood submerged in a freshwater stream, 25 December
2014, Jaap van Strien, Site 5-14-2 (MFLU 18-1130, reference specimen designated here), living culture
(MFLUCC 16-0179, GZCC 15-0071).
GenBank numbers: ITS: MH606233, LSU: MH606222,
RPB2: MH626448.
Notes: The genus Acrogenospora includes 11 species
and two of them have been linked to sexual morphs in
Farlowiella. The synonymy of Acrogenospora with
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Fig. 49 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, TEF1-a and RPB2 sequence data for
Trematosphaeriaceae and several closely related families in Dothideomycetes. Related sequences are taken from Suetrong et al. (2009)
and Wanasinghe et al. (2018a), forty-three strains are included in the
combined analyses which comprise total 3628 characters (1119
characters for LSU, 983 characters for SSU, 821 characters for TEF1a, 705 characters for RPB2) after alignment. Pleospora herbarum
(CBS 191.86) (Pleosporaceae, Pleosporales) is used as the outgroup
taxon. Single gene analyses are carried out and the topology of each
tree had clade stability. Tree topology of the maximum likelihood
analysis is similar to the maximum parsimony analysis and the
Bayesian analysis. The best RaxML tree with a final likelihood value
of –24083.791079 is presented. The matrix had 1258 distinct
alignment patterns, with 18.62% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.250098,
C = 0.240190, G = 0.272034, T = 0.237678; substitution rates AC =
1.254339, AG = 3.593578, AT = 1.238765, CG = 1.071163, CT =
7.108157, GT = 1.000000; gamma distribution shape parameter
a = 0.194764. Bootstrap values for maximum likelihood (ML) and
equal to or greater than 75% and Bayesian posterior probabilities
(BYPP) equal or greater than 0.95 are placed above the branches. The
newly generated sequence is indicated in bold and blue
Farlowiella has long been accepted based on the distinctive
morphology of the asexual morphs (Ellis 1971, 1976; Goh
et al. 1998a; Rossman et al. 2015b). The generic name
Acrogenospora has been protected over its sexual morph
Farlowiella (Rossman et al. 2015b). In this study, molecular data has been generated to support its asexual-sexual
connection. Acrogenospora sphaerocephala was designated as the type species in the genus (Ellis 1971) and it is
a frequently observed species with a worldwide distribution
(Goh et al. 1998a).
Phylogenetic analyses indicate that our collection A.
sphaerocephala (MFLUCC 16-0179) formed a distinctive
branch in the genus, close to A. carmichaeliana (FMR
11021, CBS 179.73, CBS 164.76 and CBS 206.36).
However, the identification of these A. carmichaeliana
(Berk.) Rossman & Crous collections is questionable as
only molecular sequence data was provided and sequence
data is unavailable for the ex-type strain. Acrogenospora
sphaerocephala was thought to be the asexual morph of F.
carmichaeliana due to the information in GenBank. It must
be wrongly identified as A. megalospora was introduced as
the asexual morph of Farlowiella carmichaeliana (Ellis
1971; Goh et al. 1998a) and the combination of the holomorph species was suggested as Acrogenospora carmichaeliana (Rossman et al. 2015b). Our collection was
identified as A. sphaerocephala with the morphological
characters well-matched with the original diagnosis.
Acrogenospora sphaerocephala is morphologically similar
to A. megalospora except A. sphaerocephala has globose
conidia, while A. megalospora produces broadly ellipsoidal
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Fig. 50 Acrogenospora sphaerocephala (MFLU 18-1130, reference
specimen). a Colony on substrate. b, c Conidiophores with conidia. d,
e Conidiophores. f, g Conidiogenous cells with conidia. h–j Conidia.
77
k Germinated conidium on PDA. l, m Culture (l from above view,
m from below view). Scale bars: a = 200 lm, b–d = 100 lm, e–
g = 50 lm, h = 30 lm, k = 20 lm, i, j = 10 lm
123
78
or obovoid conidia (Ellis 1971; Goh et al. 1998a; Cai et al.
2006). Additionally, our collection has similar size of
conidiophores (155–360 9 3.5–9.5 lm) with given by
Ellis (1971) (up to 380 lm long, 5–8 lm wide at the apex
and 9–11 lm wide at the base) and similar conidial size
(18–30 lm, x = 24 lm) with given by Ellis (1971) (15–
33 9 14–33 lm, x = 28 9 27 lm). Thus, a reference
specimen for A. sphaerocephala is designated here.
Acrogenospora thailandica J. Yang & K.D. Hyde, sp. nov.
Index Fungorum number: IF555499; Facesoffungi
number: FoF04676; Fig. 51
Etymology: Referring to the distribution in Thailand.
Holotype: MFLU 18-1129
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies sparse, scattered,
dark brown, glistening, hairy. Mycelium mostly immersed,
consisting of septate, hyaline to pale brown hyphae.
Conidiophores 850–950 9 3.5–8 lm, macronematous,
mononematous, solitary, erect, straight or slightly flexuous,
septate, long cylindrical, slightly tapering towards the apex,
pale to dark brown, paler towards the apex, smooth, thickwalled at the base. Conidiogenous cells monoblastic,
integrated, intercalary, brown, cylindrical. Conidia
acrogenous, 15.5–24.5 lm (
x = 19 lm, n = 35) diam.,
holoblastic, olive-green to dark brown, spherical or subspherical, unicellular, smooth and thick-walled, guttulate,
truncate at the base.
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced from both ends. Colonies on PDA slow growing, reaching 7–10 mm diameter
after two weeks at 25 C in natural light, circular, with dark
grayish green aerial mycelium on the surface, black in
reverse with entire margin.
Material examined: THAILAND, Trat Province,
Amphoe Ko Chang, 12080 N, 102380 E, on decaying wood
submerged in a freshwater stream, 27 April 2017, YongZhong Lu, YJT-30-2 (MFLU 18-1129, holotype; HKAS
102140, isotype), ex-type living culture (MFLUCC
17-2396).
GenBank numbers: ITS: MH606234, LSU: MH606223,
RPB2: MH626449, SSU: MH606221.
Notes: Phylogenetic analyses indicate that Acrogenospora thailandica forms a basal branch in the genus together
with its sister taxon Acrogenospora sp. (JX-43) with strong
support. Yang et al. (2016c) reported Acrogenospora sp.
(JX-43) with LSU, SSU and ITS sequence data and only
mentioned that Acrogenospora sp. (JX-43) is morphologically similar to A. sphaerocephala. We therefore identify
Acrogenospora sp. (JX-43) as A. thailandica based on the
phylogenetic result (Fig. 52). Acrogenospora thailandica
resembles A. sphaerocephala in having relatively long
conidiophores, monoblastic, intercalary conidiogenous
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cells and globose conidia of similar size, but A. thailandica
has much longer conidiophores (850–950 lm long) than
those of A. sphaerocephala (155–360 lm long).
Dothideomycetes orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Notes: The order Asterinales is poorly understood due to
minimal sequence data and its taxonomic placement is
mainly based on morphology. There are different interpretations of this order. Hongsanan et al. (2014) included
only a single family Asteraceae in Asterinales, while
Guatimosim et al. (2015) added Parmulariaceae to the
order. The phylogenetic placement of Asterinales are also
different in the above studies. Until now, there is no convincing evidence that can support the placement of this
family and it is considered as a polyphyletic group. The
enrichment of sequence data of taxa of Asterinales will
make contributions towards further understanding the black
mildews.
Asterinaceae Hansf.
Notes: The family Asterinaceae was established as a
member of Myriangiales by Hansford (1946). Species of
Asterinaceae are characterized by dark brown hyphae with
hyphopodia, dark brown thyriothecia with stellate dehiscence and dark brown, 1-septate ascospores. Phylogenetic
studies have provided several different interpretations of
this family (Hongsanan et al. 2014; Guatimosim et al.
2015). There is presently no convincing evidence that can
support the placement of this family and it is considered as
a polyphyletic group. In this study, we provide two new
collections with sequences data that cluster in Asterinales
sensu lato.
Lembosia Lév.
Notes: Lembosia was established by Leveillé (1845)
with descriptions of L. dendrochili Lév., L. drimydis Lév.,
L. macula Lév. and L. tenella Lév. The genus was placed in
Lembosiaceae by Hosagoudar et al. (2001). The genus is
characterized by oval, elongate thyriothecia with X-,
Y-shaped, or longitudinal dehiscence and with lateral
appressoria on the hyphae. Hongsanan et al. (2014) referred
the genus in Asterinaceae and treated Lembosiaceae as a
synonym.
Lembosia xyliae X.Y. Zeng, T.C. Wen & K.D. Hyde
Facesoffungi number: FoF00933; Figs. 53, 54
Colonies epiphyllous, circular, dense, single to confluent. Hyphae superficial, straight to substraight, dark brown,
branching alternate to opposite at acute to wide angles,
reticulate. Hyphopodia subglobose, 1-celled, alternate,
lateral, antrorse. Sexual morph Thyriothecia dense, elongate, with stellate dehiscence, 350–500 9 200–300 lm
(
x = 400 9 250 lm, n = 20). Pseudoparaphyses filiform,
Fungal Diversity (2019) 96:1–242
Fig. 51 Acrogenospora thailandica (MFLU 18-1129, holotype).
a Colony on substrate. b, c Conidiophores with conidia. d, e
Conidiogenous cells with conidia. f, g Conidia. h Germinated
conidium on PDA. i Reproduced conidiophores and conidia from
79
culture. j, k Culture (j from above view, k from below view). Scale
bars: a = 500 lm, b, c = 200 lm, i = 100 lm, d = 30 lm, e–
h = 20 lm
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Fig. 52 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, TEF1-a and RPB2 sequence data for
Acrogenosporaceae. Nine strains are included in the combined genes
sequence analyses which comprise total 4150 characters. Hysterographium fraxini (CBS 242.32 and CBS 109.43) is selected as the
outgroup taxa. Single gene analyses are carried out and the topology
of each tree with clade stability. Tree topology of the maximum
likelihood analysis is similar to the Bayesian analysis. The best
sorting RaxML tree with a final likelihood valued of - 8661.40414 is
presented. The matrix had 269 distinct alignment patterns, with
44.96% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.259633, C = 0.231472, G = 0.273835,
T = 0.235059;
substitution
rates
AC = 27.402898,
AG =
169.064131, AT = 43.635801, CG = 74.684163, CT = 684.643950,
GT = 1.000000; gamma distribution shape parameter a = 0.194764.
Bootstrap values for maximum likelihood (ML) and equal to or
greater than 70% and Bayesian posterior probabilities (BYPP) equal
or greater than 0.95 are placed above the branches. The newly
generated sequences are indicated in bold and blue
dense, numerous, septate, hyaline. Asci 8-spored, bitunicate, ellipsoid to subglobose, sessile, 40–60 9 30–50 lm
(
x = 50 9 40 lm, n = 30). Ascospores obovoid to ellipsoid, 2-celled, slightly constricted at the septum, lower cell
slightly longer and narrower, hyaline with two oil drops in
each cell when immature, becoming brown at maturity, 25–
32 9 12–17 lm (
x = 29 9 14 lm, n = 30). Asexual
morph Undetermined
Material examined: THAILAND, Chiang Rai Province,
Doi Mae Salong, on leaves of Shorea roxburghii (Dipterocarpaceae), 22 June 2015, XY Zeng (MFLU 16-0068);
THAILAND, Amphoe Ko Chang, Yuttha Navi Ko Chang
Memorial, on leaves of Shorea roxburghii (Dipterocarpaceae), 27 April 2017, XY Zeng (MFLU 17-1052).
GenBank numbers: ITS-LSU: MK660012 (MFLU
17-1052); LSU: MK660011 (MFLU 16-0068).
Notes: This species was introduced by Ariyawansa et al.
(2015) with only a single LSU sequence data. The holotype
was found on leaves of Xylia sp., but it seems incorrectly
identified. In this study, we sequenced both the host and the
fungal species by following Zeng et al. (2018). Results
indicate that the species is associated with the host Shorea
roxburghii, which is very similar to Xylia species, and two
new gene sequences (ITS and SSU) are generated. Lembosia shoreae (R.W. Ryan) B. Song & Hosag., which
synonymized from Morenoella shoreae, is the only species
reported from Shorea, but the original description is
unavailable. Therefore, we would like to report a new host
record and update sequence data for this species.
123
Morenoina Theiss.
Notes: A recent description of Morenoina was mentioned in Tibpromma et al. (2017) where the genus was
placed in Asterinales genera incertae sedis.
Morenoina palmicola J. Fröhl., K.D. Hyde & Joanne E.
Taylor
Facesoffungi number: FoF04833; Fig. 55
Isotype: MFLU 15-0030
Saprobic on petiole of Salacca sp. Sexual morph
Thyriothecia 155–895 (up to 2400) lm long, 80–190 lm
wide (
x = 455 9 135 lm, n = 30), occurring on host surfaces, solitary, aggregated, or gregarious, easily removed
from the host surface, superficial, ellipsoid, oblong, curved,
X- or Y-shaped, flat, with longitudinal, slit-like opening,
Fungal Diversity (2019) 96:1–242
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Fig. 53 Lembosia xyliae (MFLU 16-0068, new host record). a Host
leaves. b, c Thyriothecia on leaf surface. d Squash mount of
thyriothecium. e Cross section of thyriothecium. f Hyphae with
hyphopodia. g–i Young asci. j–m Young ascospores. Scale bars: b–
d = 200 lm, e, f = 50 lm, g–m = 20 lm
linear fissure, which are branched at the margin, from the
centre to the outer rim, free hyphae and appressoria at the
margin. Upper wall comprises linear cells, with irregular,
filiform hyphae, radiating from the centre to the outer rim.
Asci 19–26 9 10–12 lm (
x = 24 9 11 lm, n = 10), 8spored, bitunicate, globose to subglobose or clavate, or
saccate to globose, apedicellate, with a distinct, thickened
apical
region.
Ascospores
9–14 9 4–7 lm
(
x = 12 9 6 lm, n = 20), oblong or fusiform, wider at the
apex, with slightly acute ends, 1-septate, with two large
guttules in each cell, hyaline, smooth-walled. Asexual
morph Undetermined.
Culture characteristics: Ascospore germinating on
MEA within 24 h and germ tube produced from both end
cells. Colonies on MEA reaching 3–4 cm diam., after two
weeks, grey to olivaceous, dense, with a fairly fluffy surface, hyphae, septate, branched, and smooth-walled.
Material examined: THAILAND, Krabi Province, on
dead petiole of Salacca sp. (Arecaceae), 8 December 2014,
Sirinapa Konta, KBR05 (MFLU 15-0030, isotype), ex-type
living culture (MFLUCC 15-0284).
GenBank numbers: ITS: MK120273, LSU: MK120272,
SSU: MK120299.
Notes: Morenoina has long taxonomic confusion concerning its familial placement. Until recently, there was no
sequence data to confirm the relationships of this genus.
Theissen (1913) introduced Morenoina with M. antarctica
as the type species. Doidge (1942) synonymised Morenoina with Lembosia and Von Arx and Müller (1975)
placed Morenoina in Leptopeltidaceae. Fröhlich and Hyde
(2000) described and illustrated M. palmicola in
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Fig. 54 Lembosia xyliae (MFLU 17-1052, new host record). a Host
leaves. b Thyriothecia on leaf surface. c Squash mount of thyriothecium. d Hyphae with hyphopodia. e Young ascus. f–i Ascospores
from young to mature state. Scale bars: b, c = 100 lm, d = 50 lm,
e = 20 lm, f–i = 10 lm
Asterinaceae. Lumbsch and Huhndorf (2010) also suggested that its placement should be in Asterinaceae. Hongsanan et al. (2014) transferred Morenoina to
Aulographaceae based on morphological characters. Tibpromma et al. (2017) introduced a new species M.
calamicola and showed its unstable phylogenetic placement. There are 26 epithets of Morenoina listed in Index
Fungorum (2019). We collected a fresh specimen which is
similar to M. palmicola J. Fröhl. et al. but from a dead
petiole of Salacca (Arecaceae) collected in Krabi,
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83
Fig. 55 Morenoina palmicola (MFLU 15-0013, new host record).
a Appearance of thyrothecia on host substrate. b, c Close up of
thyrothecia. d Cell walls of thyrothecium with radial arrangement.
e Asci. f, g Ascospores. h, i Germinated ascospores. j, k Culture on
MEA. Scale bars: a = 500 lm, b, c = 200 lm, d, e = 10 lm, f–
i = 5 lm
Thailand. The holotype was found on Calamus (Arecaceae)
in Australia (Fröhlich and Hyde 2000). Since they have
similar features it is wise to introduce our novel isolate as a
new host record of Morenoina palmicola and this is the
second species which has DNA based sequence data for
phylogenetic analyses. In our analyses of combined LSU,
and SSU sequence data, Morenoina palmicola and M.
calamicola did not group in Asterinaceae sensu stricto.
Both species have a close phylogenetic affiliation to
Melaspileellaceae and Stictographaceae without statistical
support (Fig. 56). Limited taxon sampling in the phylogenetic analyses may have resulted in inadequate resolution
of this genus. Wider taxon sampling and accurate taxonomic information based on morphological examination of
specimens, coupled with phylogenetic data are needed, to
better integrate Morenoina into an appropriate taxonomic
system.
Botryosphaeriales C.L. Schoch
Notes: Botryosphaeriales are considered as an important
group in Dothidiomycetes due to the ecological and economic significance (Mehl et al. 2014). Many are classified
as plant pathogens, known to cause many diseases on wide
range of important plants. Some cause opportunistic
infections in humans (de Hoog et al. 2000), and most are
endophytes (Slippers & Wingfield 2007) and saprobes in
plants. Based on morphological and molecular phylogenetic sequence data, Botryophaeriales comprise
six families, Aplosporellaceae, Botryosphaeriaceae,
Melanopsaceae, Phyllostictaceae, Planistromellaceae and
Saccharataceae (Dissanayake et al. 2016, 2017a; Phillips
et al. 2019).
Botryosphaeriaceae Theiss. & P. Syd.
Notes: Botryosphaeriaceae was introduced as a family in
Botryosphaeriales by Schoch et al. (2006) and it represents
the predominant family in the order in comprising 23
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b Fig. 56 Phylogram generated from maximum likelihood analysis
based on combined LSU, and SSU sequence data. Related sequences
are taken from from Dai et al. (2018). Eighty-five strains are included
in the combined analyses which comprise a total of 3041 characters
(1973 characters for LSU and 1068 characters for SSU) after
alignment. Caliciopsis pinea (AFTOL-ID 1869) is used as the
outgroup taxon. Single gene analyses were carried out and the
topology of each tree had clade stability. Tree topology of the
maximum likelihood analysis is similar to the Bayesian analysis. The
best RaxML tree with a final likelihood value of - 27484.335626 is
presented. The matrix had 1767 distinct alignment patterns, with
52.88% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.252599, C = 0.225029, G = 0.296253,
T = 0.226118; substitution rates AC = 0.842370, AG = 2.253531,
AT = 0.847156, CG = 1.108746, CT = 5.466539, GT = 1.000000;
gamma distribution shape parameter a = 0.400506. Bootstrap values
for maximum likelihood (ML) equal to or greater than 50 and
Bayesian posterior probabilities (BYPP) equal to or greater than 0.9
are placed above the branches respectively. The newly generated
sequences are indicated in bold and blue
genera and 187 species (Dissanayake et al. 2016). Species
of Botryosphaeriaceae have a cosmopolitan distribution on
a wide range of plant hosts, as endophytes, saprobes and
plant pathogens.
Dothiorella Sacc., Michelia 2(6): 5 (1880)
Notes: Dothiorella species are characterized by conidia
that become pigmented and 1-septate, while they are still
attached to the conidiogenous cells (Phillips et al. 2013).
Presently, 30 species are accepted in the genus (Dissanayake et al. 2016). Wide host ranges and morphological
plasticity within the genus have made it impossible to
identify species based only on morphology. Phillips et al.
(2008) have introduced the genus Spencermartinsia to
accommodate the Dothiorella-like species but Yang et al.
(2016b) have synonymized Spencermartinsia species into
genus Dothiorella because they found that apiculate
ascospores were not reliable to use for separating these two
genera. Therefore we also have treated them as a single
genus, Dothiorella.
Dothiorella plurivora (Abdollahz., Javadi & A.J.L. Phillips) Tao Yang & Crous, in Yang, Groenewald, Jami,
Cheewangkoon, Abdollahzadeh, Lombard & Crous 2016
Facesoffungi number: FoF05785; Fig. 57
Saprobic on dead wood. Sexual morph Undetermined.
Asexual morph Coelomycetous. Conidiomata 155–
260 lm diam., stromatic, superficial, unilocular, glabrous,
brown to black, globose to subglobose, papillate. Ostiole
central, short, lined with periphyses. Wall of conidiomata
20–45 lm membranaceous, composed of dark brown, or
brown to hyaline cells of textura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells 5–
9 9 1.5–5 lm (
x = 7.5 9 3.5 lm, n = 30), enteroblastic,
phialidic, cylindrical to subcylindrical, hyaline, smooth-
85
walled, arising from the inner layers of conidioma. Conidia
14–22 9 5–10 lm (
x = 18 9 8 lm, n = 50), ellipsoidal to
oval, hyaline when young, becoming to brown when
mature, smooth-walled, guttulate, 1-septate.
Culture characteristics: Ascospores germinating on
PDA (potato dextrose agar) within 2 days at 23 C.
Sporulation after 1 week, colony 2 cm diam. gray from
above and reverse, with white mycelia radiating outwards,
with irregular form of margin, filamentous mycelium, with
rough surface.
Material examined: CHINA, Yunnan Province, Kunming, Songhuaba Lake; on dead wood, 3 September 2017;
S.K. Huang (KUN HKAS 99572), living culture (KUMCC
18-0013).
GenBank numbers: ITS: MK459467, LSU: MK459468.
Notes: Spencermartinsia plurivora has been isolated
from soil in Iran (Abdollahzadeh et al. 2014). Multi-gene
analysis showed that Spencermartinsia to have close
affinities with Dothiorella (Phillips et al. 2013; Slippers
et al. 2013). Therefore, Yang et al. (2016b) placed
Spencermartinsia as a synonym of Dothiorella. In phylogenetic analysis of Dothiorella, our strain clusters with the
ex-type strain of Dothiorella plurivora (IRAN 1557C).
Dothiorella plurivora is characterized by 7–10 9 3–5 lm
conidiogenous cells and ellipsoid to ovoid, brown, 1-septate conidia (20–25 9 10–13 lm) (Abdollahzadeh et al.
2014). Morphological similarities reveal it is similar to our
taxon.
Dothiorella rhamni Wanas., Bulgakov, E.B.G. Jones &
K.D. Hyde, in Li et al., Fungal Diversity 78: 253 (2016)
Facesoffungi number: FoF01668; Fig. 58
Saprobic or weak pathogen on dead twigs of Cercis
canadensis L. (Fabaceae). Sexual morph Undetermined.
Asexual morph Conidiomata 310–370 lm high 9 360–
500 lm diam. (
x = 339 9 425 lm, n = 10), pycnidial,
stromatic, mostly solitary, semi-immersed to immersed in
the host, dark brown to black, apapillate. Peridium multilayered 50–57 lm wide at the base, 68–83 lm wide in
sides, comprising 8–10 layers, heavily pigmented, thickwalled, comprising blackish to dark brown, textura angularis cells. Conidiogenous cells 9–15 lm high 9 2–4 lm
wide, holoblastic, cylindrical to subcylindrical, hyaline, the
first conidium produced holoblastically and subsequent
conidia enteroblastically, swollen at the base, discrete,
producing a single conidium at the apex. Conidia 15–
24 9 7–11 lm (
x = 20.9 9 8.9 lm, n = 30), initially
hyaline, unicellular, becoming cinnamon to sepia and
1-septate, while still attached to conidiogenous cells;
detached conidia, hyaline, sepia or dark brown, unicellular
or 1-septate moderately thick-walled, wall externally
smooth, roughened on the inner surface, oval to ovoid,
widest in the centre, apex obtuse, base truncate or rounded.
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Fig. 57 Dothiorella plurivora (HKAS 99572, new record). a Appearance of conidiomata on host. b, c Conidiomata in vertical
section. d Wall of conidioma. e–f Conidiogenous cells with
developing conidia. g–i Conidia. j Germinated conidia. Scale bars:
b, c = 100 lm, d = 50 lm, j = 20 lm, e–i = 10 lm
Culture characteristics: Colonies on PDA, circular,
fimbriate, crenate edged, initially both surfaces white and
turning to grey towards centre after 4 days, fast growing,
reach 70 mm diam. in 6 days at 28 C.
Material examined: RUSSIA, Rostov region, Shakhty
City, Alexandrovsky (Central) Park (47.7058991 N,
40.2053368E), on dead and dying twigs of Cercis
canadensis L., 1 March 2016, Timur S. Bulgakov (MFLU
16–1545), living culture (MFLUCC 19-0001).
GenBank numbers: ITS: MK064561, TEF1-a:
MK078544.
Notes: Dothiorella rhamni was introduced by Li et al.
(2016). There are several records of this species from
Rhamnus cathartica in Russia, Tamarix gallica and
Rhamnus alaternus in Italy (Dissanayake et al. 2016; Li
et al. 2016). Based on our phylogenetic analysis of combined ITS and TEF1-a sequence data of Dothiorella species (Fig. 61), our strain (MFLUCC 19-0001) clustered
with the ex-type strain of D. rhamni (MFLUCC 14-0902)
and the two specimens share similar morphological
characters.
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87
Fig. 58 Dothiorella rhamni (MFLU 16-1545, new record). a Appearance of conidiomata on host substrate. b Vertical section of a
conidioma. c Peridium of conidioma. d, e Immature and mature
conidia attached to conidiogenous cells. f, g Mature conidia. h Culture
characters on PDA. Scale bars: b, c = 50 lm, d = 20 lm, f = 20 lm,
e, g = 10 lm
Dothiorella styphnolobii Brahmanage, Bulgakov & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555482; Facesoffungi
number: FoF05095; Fig. 59
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Fig. 59 Dothiorella styphnolobii (MFLU 17–2256, holotype). a, b
Appearance of conidiomata on Styphnolobium japonicum. c Vertical
section of conidioma. d Peridium. e Developing stages of conidia on
conidiogenous cells. f Immature conidium attach to conidiogenous
cells. g–j Conidia. Scale bars: b = 1000 lm, c = 100 lm, d,
e = 20 lm, f–j = 10 lm
Etymology: The specific epithet ‘‘styphnolobii’’ refers to
the host plant genus Styphnolobium.
Holotype: MFLU 17-2256
Saprobic on dead twigs and branches of Styphnolobium
japonicum (L.) Schott (Fabaceae). Sexual morph Undetermined. Asexual morph Conidiomata 425–500 9 160–
180 lm (
x = 460 9 170 lm, n = 30), pycnidial, stromatic,
mostly solitary, semi-immersed to immersed, dark brown
to black, ostiolate. Ostiole apapillate. Peridium 30–65 lm
wide, comprising 6–8 layers of heavily pigmented, thickwalled, blackish to dark brown, angular cells. Conidiogenous cells 7–9 9 2–4 lm, holoblastic, conical, hyaline,
swollen at the base, discrete, producing a single conidium
at the apex, proliferating at the same level giving rise to
periclinal thickenings. Conidia 18–22 9 8–10 lm
(
x = 20 9 8 lm, n = 30), initially hyaline, unicellular,
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guttulate, becoming pale brown and 1-septate, while still
attached to conidiogenous cells; detached conidia, hyaline,
dark brown, unicellular or 1-septate, slightly concentric,
oval to ovoid, apex obtuse, base truncate or rounded.
Culture characteristics: Colonies on PDA at 25 C
reaching 5 cm in 7 days, ash at first, becoming blackish
when mature and reverse ash becoming blackish when
mature.
Material examined: RUSSIA, Republic of Crimea,
Feodosia Municipality, Karadag Nature Reserve, the park
of Karadag Biological Station, on dead and dying twigs and
branches of Styphnolobium japonicum (L.) Schott, 21 June
2016, Timur S. Bulgakov (MFLU 17-2256, holotype), extype living culture (JZB3150013).
GenBank numbers: ITS: MH880849, TEF1-a:
MK069594.
Notes: Our new collection shares similar morphological
characters with Dothiorella, such as 1-septate conidia that
are dark at an early stage of development (Phillips et al.
2005, 2013). In our phylogenetic analyses, D. styphnolobii
showed close phylogenetic affinities to D. juglandis
(Fig. 61). Dothiorella styphnolobii resembles D. italica in
the characteristics of conidiomata and conidia, but are
distinct in the shape of conidiogenous cells that are subcylindrical in D. italica and conical in D. styphnolobii
(Dissanayake et al. 2017a). Dothiorella styphnolobii can
also be distinguished from D. italica by the conidiomatal
width (160–180 lm vs 680 lm), conidiogenous cells
(7–9 9 2–4 lm vs 8–15 9 3–6 lm) and conidia
(18–22 9 8–10 lm vs 28.7–43.2 9 13.2–17.5 lm) (Dissanayake et al. 2017a). However, they formed two distinct
lineages in our phylogram with the maximum likelihood
support of 80% ML (Fig. 61).
Dothiorella symphoricarposicola W.J. Li, J.K. Liu & K.D.
Hyde, Cyptogamie Mycologie 35(3): 265 (2015)
Facesoffungi number: FoF04935; Fig. 60
Saprobic on dead, aerial branch of Laburnum anagyroides Medik. Sexual morph Undetermined. Asexual
morph Conidiomata 0.17–0.53 mm (
x = 0.39 mm,
n = 10) diam., solitary, scattered, immersed to semi-immersed, partially erumpent at maturity, pyriform, unilocular, globose, black, ostiolate. Ostiole 30–60 lm diam.,
single, central, with a well-developed neck, thick-walled,
sometimes papillate. Peridium multi-layered, 45 lm wide
at the base, 20–38 lm wide in sides, outer layer composed
of 4–5 layers of thick, brown cells, inner 3–4 layers of
hyaline cells of textura angularis, cells towards inner layer
become paler. Conidiophores reduced to conidiogenous
cells.
Conidiogenous
cells
9–14 9 3–5.3 lm
(
x = 11.5 9 4.2 lm, n = 10), phialidic, cylindrical, sometimes slightly curved, hyaline, smooth, thick-walled,
formed from the inner most layer of pycnidial wall.
89
Conidia 14–21 9 7–11.5 lm (
x = 18.4 9 9 lm, n = 30),
ovoid, straight or slightly curved, guttulate, initially aseptate, hyaline, becoming 1-septate, pale to dark brown at
maturity, smooth-walled, rounded at both ends.
Culture characteristics: Colonies on PDA, circular,
fimbriate, crenate edged, with both surfaces white at first,
becoming grey towards centre and finally becoming black,
fast growing, and reach 6 cm diam. after 4 days at 28 C.
Material examined: ITALY, Forlı̀-Cesena Province,
Corniolo – Santa Sofia, on dead, aerial branch of Laburnum
anagyroides Medik. (Fabaceae), 15 May 2016, Erio Camporesi, IT-2971 (MFLU 16-1355), living culture
(MFLUCC 18-0092, KUMCC 18-0414).
GenBank numbers: ITS: MH819800, TEF1-a:
MH853685.
Notes: Dothiorella symphoricarposicola was introduced
by Li et al. (2014). There are several records of this species
from Symphoricarpos sp., Sambucus nigra, Laurus nobilis
and Laburnum alpinum in Italy (Li et al. 2014; Dissanayake et al. 2016). Here we introduce D. symphoricarposicola as a new host record. Based on our
phylogenetic analysis of combined ITS and TEF1-a
sequence data of Botryosphaeriaceae species (Fig. 61), our
strain (MFLUCC 18-0092) clusters with the ex-type strain
of D. symphoricarposicola (MFLUCC 13-0497). The two
strains share similar morphological characters. However,
our strain has slightly larger conidiomata (0.17–0.53 mm)
and conidiogenous cells (9–14 9 3–5.3 lm) compared to
the conidiomata (0.2–0.3 mm) and conidiogenous cells
(4–12 9 1.5–6 lm) of the ex-type strain (Li et al. 2014).
Lasiodiplodia Ellis & Everh.
Notes: This genus comprises 53 species (Dissanayake
et al. 2017a). Both sexual and asexual morphs have been
reported within the genus (Alves et al. 2008; Tennakoon
et al. 2016b). For the species differentiation morphology is
not a reliable character and species can be resolved using
combined ITS and TEF1-a sequence data (Phillips et al.
2013; Slippers et al. 2014).
Lasiodiplodia iraniensis Abdollahz., Zare & A.J.L. Phillips [as ‘iranensis’], Persoonia 25: 8 (2010)
Facesoffungi number: FoF04922; Fig. 62
Saprobic on dried pods of Cassia sp. Sexual morph
Undetermined. Asexual morph Appearing as raised spots
on the host. Coelomycetous. Conidiomata 210–405 lm
high 9 90–336 lm diam. (
x = 283 9 194 lm, n = 10),
stromatic, pycnidial, immersed, aggregated, unilocular,
globose to depressed globose. Peridium 20–55 lm wide,
composed of 3–6-layers of pale brown to brown, smoothwalled cells of textura angularis. Pseudoparaphyses 2.1–
3.9 lm wide, aseptate, hyaline. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 10–22 lm 9 3–
6.3 lm (
x = 13 9 4.4 lm, n = 20), lining the pycnidial
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Fig. 60 Dothiorella symphoricarposicola (MFLUCC 18-0092, new
host record). a Appearance of conidiomata on dead branch of
Laburnum anagyroides. b Longitudinal section of a conidioma.
c Longitudinal section of a conidioma wall showing cell organization.
d–f Conidiogenous cells with developing conidia. g Conidia. h, i
Colony on PDA (h from below view, i from above view). Scale bars:
a = 0.5 mm, b = 100 lm, c–g = 20 lm
cavity, holoblastic, subcylindrical, hyaline, smooth-walled.
Conidia 14–28 9 8–14.5 lm (
x = 22.5 9 12, n = 25)
ovoid, rarely subovoid to ellipsoid-ovoid, with a broadly
rounded apex and truncate base, hyaline, thick-walled,
smooth-walled.
Culture characteristics: Colonies on PDA reaching
55 mm diam. after 7 days at 25 C, circular, smooth
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margin, white at first, ash to greenish black after 2 weeks,
flat on the surface, with aerial mycelium, reverse black.
Material examined: THAILAND, Chiang Rai Province,
on dried pods of Cassia sp. (Fabaceae), 22 December 2016,
R.H. Perera, RHP 37 (MFLU 17-2847), living culture
(MFLUCC 18-0586).
GenBank numbers: ITS: MH107831, LSU: MH107832,
TEF1-a: MH107833, TUB2: MH107834.
Notes: The new collection is a typical Lasiodiplodia
species, and it shares the similar morphology with Lasiodiplodia iraniensis which the dimensions of their conidiophores and conidia are almost identical (Abdollahzadeh
et al. 2010). Conidia of L. iraniensis become 1-septate and
dark brown with longitudinal striations with the time
(Abdollahzadeh et al. 2010). However, we did not observe
released mature conidia in our collection. The multi-gene
phylogenetic analysis showed that our newly obtained
strain (MFLUCC 18-0586) clustered together with Lasiodiplodia iraniensis (Fig. 63). Therefore, we identify our
collection as L. iraniensis and it is reported here as a new
record to Thailand and Cassia sp.
Jahnulales K.L. Pang, Abdel-Wahab, El-Shar., E.B.G.
Jones & Sivichai
Notes: Jahnulales was erected by Pang et al. (2002)
which comprises freshwater lignicolous ascomycetes
(Huang et al. 2018). This order is phylogenetically related
to the Dothideales, Patellariales, and Pleosporales (Campbell et al. 2007). Jahnulales species often occur on rotting
or soft submerged corticated or decorticated wood (Inderbitzin et al. 2001; Suetrong et al. 2011a; Tanaka et al. 2015;
Huang et al. 2018). They produce ascomata with multilayered peridial walls, composed of large cells, stalked and/
or sessile bitunicate asci, and one-septate ascospores with
appendages or gelatinous sheaths (Pang et al. 2002; Suetrong et al. 2011a; Jones et al. 2015). Two families, Aliquandostipitaceae and Manglicolaceae are currently
accepted within this order (Jones et al. 2015; Wijayawardene et al. 2018a).
Aliquandostipitaceae Inderb.
Notes: Aliquandostipitaceae, typified by Aliquandostipite Inderb. is characterized by globose to subglobose
ascomata, and one-septate ascospores (Inderbitzin et al.
2001). The sexual morph genera, Aliquandostipite, Jahnula
and Megalohypha (Kirschstein 1936; Inderbitzin et al.
2001; Ferrer et al. 2007), and asexual morph genera, Brachiosphaera, Speiropsis and Xylomyces are currently
accommodated within this family (Tubaki 1958; Descals
et al. 1976; Goos et al. 1977; Campbell et al. 2007; Suetrong et al. 2011a; Wijayawardene et al. 2018a).
91
Jahnula Kirschst.
Notes: Jahnula was introduced by Kirschstein (1936)
and typified by Jahnula aquatica (Kirschst.) Kirschst.
Jahnula species have been reported from wood or decorticated wood in freshwater habitats (Hawksworth 1984;
Hyde 1993; Hyde and Goh 1998; Ho et al. 2002; Suetrong
et al. 2011a; Fournier et al. 2015; Huang et al. 2018). The
genus is polyphyletic, however, Jahnula sensu stricto
accommodates J. aquatica, J. granulosa, and J. rostrata
(Suetrong et al. 2011a). We introduce an additional taxon
to this genus from decaying wood submerged in a river in
Australia.
Jahnula queenslandica Dayarathne, Fryar & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555384; Facesoffungi
number: FoF04945; Fig. 64
Etymology: The name queenslandica refers to the geographic location where the specimen was collected.
Holotype: AQ522780
Saprobic on dead wood. Sexual morph Ascomata 280–
330 9 200–300 lm (
x = 350 9 250 lm, n = 5), perithecial, solitary, superficial to semi-immersed, unilocular,
obpyriform to subglobose, dark brown to black, papillate,
ostiolate. Ostiole central, composed of hyaline periphyses.
Peridium 28–40 lm thick, membranous, composed of
brown cells of textura angularis. Hamathecium comprising
1.5–2 lm wide, septate, branched, filiform pseudoparaphyses, embedded in a gelatinous matrix. Asci 140–
200 9 12–22 lm (
x = 180 9 16 lm, n = 20), 8-spored,
bitunicate, fissitunicate, cylindrical, pedicellate, rounded at
apex, with a distinct ocular chamber. Ascospores 20–
34 9 8–16 lm (
x = 25 9 12 lm, n = 30), uniseriate, initially light brown, becoming dark brown at maturity, oval
to broadly ellipsoid, aseptate, with a mammiform apex,
slightly curved, smooth-walled, often with two large guttules, apiosporous, rounded at lower end, with hyaline cell
at lower end, up to 2–3 lm, lacking a sheath. Asexual
morph undetermined.
Material examined: AUSTRALIA, Queensland, Daintree National Park, fast running stream with clear water and
a rocky base, S 16.170779, E 145.405268, on decaying
wood submerged in a river, 14 April 2015, Sally Fryar and
Toby Cawson, CT110 (AQ522780, holotype; MFLU
18-1692, isotype).
GenBank numbers: LSU: MH878780, ITS: MH878782.
Notes: Jahnula queenslandica has obpyriform to subglobose ascomata, septate, branched, filiform pseudoparaphyses, cylindrical asci and oval to broadly ellipsoid
ascospores that are characteristic of Jahnula (Raja et al.
2008; Sivichai and Boonyuen 2010; Suetrong et al. 2011a;
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b Fig. 61 Phylogram generated from maximum parsimony analysis
based on combined ITS and TEF1-a sequence data for Dothiorella
species in family Botryosphaeriaceae. Related sequences are taken
from Wanasinghe et al. (2018a) and GenBank. Sixty-six strains are
included in the combined analyses which comprise 828 characters
after alignment. Lasiodiplodia theobromae (CBS 164.96)
(Botryosphaeriaceae and Botryosphaeriales) is used as the outgroup
taxon. Values for maximum likelihood equal to or greater than 60 and
Bayesian posterior probabilities equal or greater than 0.90 are placed.
Maximum parsimony analysis of 85 parsimony informative characters
resulted in a most parsimonious tree (CI = 0.545, RI = 0.824,
RC = 0.449, HI = 0.455). The ex-type strains are in bold and black.
The newly generated sequences are in bold and blue
Fournier et al. 2015; Huang et al. 2018). However, J.
queenslandica is unique from other Jahnula species by
having apiospores. According to our phylogram (Fig. 65)
J. queenslandica (MFLU 18-1692) is closely related to J.
sunyatsenii (UBC F13876) (Inderb.) K.L. Pang, E.B.G.
Jones & Sivichai with high statistical support (91% ML/
0.99 BYPP). Jahnula queenslandica can be distinguish
from J. sunyatsenii by its aseptate, dark brown apiospores,
while ascospores of J. sunyatsenii are 1-septate, light
brown with two bipolar, helmet-shaped appendages, tending to unite over the respective ends (Raja and Shearer
2006).
Tubeufiales Boonmee & K.D. Hyde
Notes: The order Tubeufiales was introduced by Boonmee et al. (2014) for a single family, Tubeufiaceae. Later,
Suetrong et al. (2014) introduced the second family
Wiesneriomycetaceae into Tubeufiales. Recently, based on
divergence time estimates, Liu et al. (2017) synonymized
the orders Bezerromycetales and Wiesneriomycetales
under Tubeufiales.
Tubeufiaceae M.E. Barr
Notes: Barr (1979) introduced the family Tubeufiaceae
typified by Tubeufia. According to recent study by Lu et al.
(2018b) this family comprises 38 genera. The sexual
morphs of Tubeufiaceae are characterized by having
superficial ascomata, pseudoparaphysate hamathecium,
bitunicate asci, and multi-septate, hyaline to pale brown
cylindrical ascospores (Barr 1980; Lu et al. 2017; Liu et al.
2018; Brahmanage et al. 2017; Phookamsak et al. 2018).
Hyphomycetous asexual morphs of Tubeufiaceae are often
helicosporous, while some are chlamydosporous and
phragmosporous (Boonmee et al. 2011, 2014; Brahmanage
et al. 2017; Doilom et al. 2017; Luo et al. 2017; Liu et al.
2018; Lu 2018).
Kamalomyces R.K. Verma, N. Sharma & Soni
Notes: The genus Kamalomyces introduced by Verma
et al. (2008), is typified by K. indicus R.K. Verma, N.
Sharma & Soni. Kamalomyces has a unique set of
93
characters, such as ascomata on a subiculum of black
hyphae, with solitary, gregarious, subglobose to lemoniform ascomata with short stalks and lacking ostioles,
bitunicate, broadly cylindrical to clavate asci and hyaline,
vermiform, crowded, septate ascospores (Verma et al.
2008; Dubey and Neelima 2013; Boonmee et al.
2011, 2014; Phookamsak et al. 2018; Lu et al. 2018a, b).
Kamalomyces currently comprises four species viz. K.
bambusicola Phook., Y.Z. Lu & K.D. Hyde, K. indicus, K.
mahabaleshwarensis Rashmi Dubey & Moonamb. and K.
thailandicus Phook., Y.Z. Lu & K.D. Hyde (Verma et al.
2008; Dubey and Neelima 2013; Phookamsak et al. 2018).
The phylogenetic relationships of Kamalomyces were
reported by Phookamsak et al. (2018). We introduce a
novel marine species to this genus based on combined
LSU, ITS and TEF1-a sequence data.
Kamalomyces mangrovei Dayarathne & K.D. Hyde, sp.
nov.
Index Fungorum number: IF555385; Facesoffungi
number: FoF04946; Fig. 66
Etymology: Epithet derived from the mangrove habitat
that species found.
Holotype: MFLU 18-1691
Saprobic on decaying, submerged wood in a mangrove
stand. Sexual morph Ascomata 280–315 9 250–300 lm,
superficial, solitary to gregarious, embedded in a subiculum
of crowded, black, septate, thick-walled hyphae, superficial,
solitary, gregarious, globose to subglobose, glabrous, shortstalked, apapillate, with indistinct ostiolate. Peridium 30–
45 lm wide, comprising light brown cells of textura angularis, and inwardly small, subhyaline cells of textura prismatica. Hamathecium comprising numerous, 1.5–2 lm
wide, filiform, septate, branched, hyaline pseudoparaphyses.
Asci 148–180 9 16–20 lm (
x = 165 9 18 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindrical to clavate, short
pedicellate, apically rounded, with an ocular chamber. Ascospores 52–67 9 6.5–8.5 lm (
x = 62 9 7.5 lm, n = 30),
2–3-seriate, hyaline becoming light brown when mature,
elongate cylindrical to fusiform-clavate, tapering towards
the lower cells, enlarged at the 4th and 5th cell, straight or
slightly curved, 8–9 septa, distoseptate.
Culture characteristics: Colonies on PDA reaching
3 cm diam. after 30 days at 25 C, circular, smooth margin
white at first, dark gray to black after 6 weeks, flat on the
surface, without aerial mycelium, reverse brownish black.
Material examined: THAILAND, Ranong Province,
Maung District, Mu 4 Tombol Ngao, Ranong Mangrove
Research Center (GPS: 9430 to 9570 N; 98290 to 98390 E)
on decaying, submerged wood of mangrove, 7 December
2016, Monika C. Dayarathne, MCD 053 (MFLU 18-1691,
holotype), ex-type living culture (MFLUCC 17-0407,
TBRC).
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Fig. 62 Lasiodiplodia iraniensis (MFLU 17-2847, new host/geographical record). a Herbarium material. b Conidiomata on host
substrate. c, d Sections through conidiomata. e Peridium. f Paraphyses
in Congo red. g, h Conidiophores with conidia. i–k Conidia.
l Germinating conidium. m, n Colony on PDA. Scale bars: c,
d = 200 lm, e = 50 lm, f–i = 20 lm
GenBank numbers: ITS: MH878781, LSU: MH878779,
SSU: MH878796, TEF1-a: MH886508.
Notes: Kamalomyces mangrovei is characterized by
solitary to gregarious, globose to subglobose, short-stalked
ascomata, lacking ostioles embedded in a subiculum of
black hyphae, bitunicate, cylindrical to clavate asci and
hyaline, elongate cylindrical to fusiform-clavate, septate
ascospores. Kamalomyces mangrovei is morphologically
best fit Kamalomyces (Verma et al. 2008; Dubey and
Neelima 2013; Boonmee et al. 2011, 2014, Phookamsak
et al. 2018; Lu et al. 2018b). Kamalomyces mangrovei
closely resembles K. bambusicola Y.Z. Lu & K.D. Hyde
and K. thailandicus Phook., Y.Z. Lu & K.D. Hyde due to
its ascomatal characters, asci and ascospore shape. However, K. mangrovei differs significantly from K. bambusicola and K. thailandicus by having fewer ascospore septa
(8–9 vs 27–30 and 33–36 septa) and being swollen at the
4th and 5th cells. The novel species occurred on a mangrove species while K. bambusicola and K. thailandicus
have been reported from bamboo. This is the first record of
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Tubeufiaceae from a marine habitat (Jones et al. 2015,
2019). According to our phylogenetic analyses with concatenated LSU, ITS and TEF1-a sequence data, K. mangrovei clustered within the genus Kamalomyces with high
statistical support (84% ML/75% MP/0.99 BYPP, Fig. 67).
Eurotiomycetes O.E. Erikss. & Winka
Chaetothyriomycetidae Doweld
Chaetothyriales M.E. Barr
Notes: Barr (1987) established the order Chaetothyriales
based on the presence of periphysoids in the ascomata.
However, Loculoascomycetes was suggested to be paraphyletic and Chaetothyriales had closer affinity with
Eurotiales based on SSU sequence data (Haase et al. 1995;
Spatafora et al. 1995; Berbee 1996; Silva-Hanlin and
Hanlin 1999). Doweld (2001) introduced the subclass
Chaetothyriomycetidae for Chaetothyriales. Chaetothyriales comprises ecologically diverse species including
opportunistic pathogens on humans (Réblová et al. 2013;
de Hoog 2014; Hyde et al. 2018a, b). Five families are
currently recognized in Chaetothyriales, viz. Chaetothyriaceae, Cyphellophoraceae, Epibryaceae, Herpotrichiellaceae, and Trichomeriaceae (Teixeira et al. 2017).
Herpotrichiellaceae Munk
Notes: The family Herpotrichiellaceae was introduced
by Munk (1953), based on the type genus Herpotrichiella
Petr., along with Berlesiella Sacc., Capronia Sacc., Dictyotrichiella Munk and Didymotrichiella Munk. Sexual
morphs in this family are characterized by ascomata with
setae and ostioles, bitunicate, saccate to ovoid asci with a
thickened apex and pale grey to brown ascospores (Munk
1953), while asexual morphs are diverse dematiaceous
hyphomycetes (Gueidan et al. 2014; Klaubauf et al. 2014;
Liu et al. 2015; Tian et al. 2016; Dong et al. 2018).
However, the classification of some hyphomycetes in this
family has been controversial. Some previously morphologically-based species were excluded from Herpotrichiellaceae with molecular evidence. For example,
Veronaea simplex Papendorf was transferred to Venturiaceae (Arzanlou et al. 2007). Some genera, e.g. Cladophialophora and Exophiala require further study to confirm
their natural classification (Liu et al. 2015; Hyde et al.
2016; Tian et al. 2016; Dong et al. 2018). Currently,
Herpotrichiellaceae comprises 15 genera and resides in the
order Chaetothyriales (Liu et al. 2015; Hyde et al. 2016;
Wijayawardene et al. 2018a).
Thysanorea Arzanlou
Notes: Arzanlou et al. (2007) studied ramichloridiumlike species and revealed that the genus Periconiella was
polyphyletic. Thysanorea was therefore introduced to
accommodate Periconiella papuana Aptroot as T. papuana
(Aptroot) Arzanlou, W. Gams & Crous (Arzanlou et al.
95
2007). Thysanorea differs from Periconiella in branching
pattern of conidiophores and prominent denticle-like
conidiogenous loci (Arzanlou et al. 2007). Dong et al.
(2018) introduced the second species, T. aquatica W.
Dong, H. Zhang & K.D. Hyde from an aquatic habitat.
Only two species were accepted in Thysanorea (Arzanlou
et al. 2007; Dong et al. 2018).
Thysanorea uniseptata N.G. Liu & K.D. Hyde, sp. nov.
Index Fungorum number: IF555387; Facesoffungi
number: FoF04949; Fig. 68
Etymology: Named after the fact that conidia are
1-septate.
Holotype: MFLU 18-1723
Saprobic on submerged wood. Sexual morph Undetermined. Asexual morph Colonies on natural substrate
effuse, scattered, velvety, dark brown to black, with greyish white, glistening heads of conidia at apex. Mycelium
partly immersed, partly superficial, composed of pale
brown to brown, branched, septate, smooth hyphae. Conidiophores up to 310 lm long, 5.5–8 lm wide at base, 3–
5 lm wide in upper half, macronematous, mononematous,
branched at apex, erect, straight or broadly curved, septate,
constricted at septa near apex, not constricted at septa
below, thick-walled, smooth-walled, brown below, paler
towards apex. Conidiogenous cells 6–11.5 9 2–4 lm
(
x = 8.9 9 3.2 lm, n = 15), polyblastic, terminal or
intercalary at main stem and fertile branches, hyaline or
subhyaline, urniform, conidiogenous loci obvious. Conidia
6.5–9 lm long, 2–3 lm wide at the septum
(
x = 7.6 9 2.5 lm, n = 30), acropleurogenous, solitary,
pyriform, hyaline and aseptate when young, subhyaline,
(0–)1-septate when mature, not constricted at the septa,
broadly round at the apex, attenuate and narrowly truncate
at the base, with darkened hilum, smooth-walled.
Culture characteristics: Conidia germinated on water
agar media within 24 h. Germ tubes produced from one or
both ends of conidium. Colonies growing on PDA circular,
edge entire, with umbonate surface, greyish brown to
brown from above, dark brown from reverse, mycelium
dense, fluffy.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Ban Nang Lae Nai, on decaying wood
submerged in a freshwater stream, 6 March 2018, N.G. Liu,
CR067 (MFLU 18-1723, holotype), ex-type living culture
(MFLUCC 18-0701).
GenBank numbers: ITS: MH883033, LSU: MH883031,
SSU: MH883049.
Notes: Thysanorea uniseptata resembles T. aquatica and
T. papuana in having obvious denticle-like conidiogenous
loci and pyriform, (0–)1-septate conidia. However, conidia
of T. aquatica have prominent guttules and are constricted
at septa, features that distinguish it from T. uniseptata.
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96
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Fungal Diversity (2019) 96:1–242
b Fig. 63 Phylogram generated from maximum parsimony analysis
based on combined ITS and TEF1-a sequence data for Lasiodiplodia
species in family Botryosphaeriaceae. Related sequences are taken
from Wanasinghe et al. (2018a). Sixty-three strains are included in the
combined analyses which comprise 1300 characters after alignment.
Neofusicoccum luteum (CBS 110497) (Botryosphaeriaceae and
Botryosphaeriales) is used as the outgroup taxon. Tree topology of
the maximum likelihood analysis is similar to the maximum
parsimony analysis and the Bayesian analysis. Maximum parsimony
analysis of 575 constant characters and 79 informative characters
resulted in most parsimonious tree (CI = 0.551, RI = 0.769, RC =
0.423, HI = 0.449). The best RaxML tree with a final likelihood
value of - 4380.160038 is presented. The matrix had 317 distinct
alignment patterns, with 36.03% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.215525,
C = 0.287133, G = 0.258862, T = 0.238480; substitution rates AC =
0.872639, AG = 3.555191, AT = 1.330285, CG = 1.135935, CT =
5.468515, GT = 1.000000; gamma distribution shape parameter
a = 0.196200. Bootstrap values for maximum parsimony (MP) and
maximum likelihood (ML) equal to or greater than 65 and Bayesian
posterior probabilities (BYPP) equal or greater than 0.90 are placed
below the branches respectively. The ex-type strains are in bold and
black. The newly generated sequence is indicated in bold and blue
Thysanorea uniseptata differs from T. papuana in having
hyaline or subhyaline conidiogenous cells, while the latter
are golden to dark brown. Moreover, T. uniseptata has
smaller conidiogenous cells and larger conidia than those
of T. papuana (6–11.5 vs 5–20 lm long and 6.5–9 vs
5–6 lm long). Phylogenetic analysis showed that
T. uniseptata is phylogenetically distinct from T. aquatica
and T. papuana (Fig. 69).
Eurotiomycetidae Geiser & Lutzoni
Eurotiales G.W. Martin ex Benny& Kimbr.
Notes: Currently, the order Eurotiales comprise three
families, Aspergillaceae Link, Thermoascaceae Apinis and
Trichocomaceae E. Fisch (Houbraken and Samson 2011).
Some species in the Eurotiales are able to grow in extreme
environments, such as low water activity, low acidity, low
levels of oxygen, or at extreme high temperatures (Houbraken and Samson 2011; Houbraken et al. 2014).
Aspergillaceae Link
Notes: The family Aspergillaceae was erected by Link
(1826) and belongs to Eurotiales. The most well-known
species of this family belongs to genera Aspergillus P.
Micheli ex Haller and Penicillium Link (Houbraken and
Samson 2011; Houbraken et al. 2014). Several species
belong to Aspergillaceae have the ability to produce a
diversity of secondary metabolites (or extrolites), organic
acids and diverse enzymes that degrade a wide variety of
complex biomolecules (Geiser et al. 2006; Pitt and Hocking 2009; Samson et al. 2010; Houbraken and Samson
2011), while others can cause disease in humans.
97
Penicillium Link
Notes: The genus is subdivided in two subgenera
(Aspergilloides and Penicillium) and 26 sections (Houbraken and Samson 2011; Houbraken et al. 2016). Species
of Penicillium are ubiquitous and known to cause rot in
fruits and bulbs of plants, including P. expansum (apples
and pears), P. digitatum, P. italicum and P. ulaiense (citrus
fruits), P. tulipae (tulip bulbs) and P. allii (garlic) (Balgrie
2003; Samson and Frisvad 2004; Valdez et al. 2006).
Currently, the genus contains more than 350 accepted
species (Visagie et al. 2014).
Section Fasciculata contains species that are found on
stored or manufactured foods, and have rough-walled
conidiophore stipes and subglobose conidia (Houbraken
et al. 2016). Species in this section are able to grow at low
water activities, and low temperature (with the exception of
species in series Verrucosa) (Houbraken et al. 2016).
Section Fasciculata is subdivided in four series: Camemberti, Corymbifera, Viridicata, Verrucosa. Six species are
currently accepted in series Corymbifera (P. albocoremium, P. allii, P. hirsutum, P. hordei, P. radicicola, P.
tulipae) and most of those species (except P. hordei) have a
strong association with flower bulbs and other plant roots
(Houbraken et al. 2016).
During an investigation of the fungal diversity on roots
of deodeok (Codonopsis lanceolata), rotten apple (Malus
pumila), freshwater, and soil in Korea, five new species of
Penicillium were isolated and these are described here
based on phylogenetic analyses (Figs. 70, 71, 72, 73) and
morphological characteristics (Figs. 74, 75, 76, 77, 78).
Penicillium circulare Hyang B. Lee, P.M. Kirk & T.T.T.
Nguyen sp. nov.
Index Fungorum number: IF555413; Facesoffungi
number: FoF05793; Fig. 74
Etymology: circulare, referring to the formation of two
concentric rings on YES media.
Holotype: CNUFC-GEU220-1
Macromorphology: CYA, 25 C: Sporulation on CYA
moderate; green in sporulating area with white mycelium
in centre, yellow at the margins; radial wrinkles; exudate
absent; soluble pigment absent; reverse orange to dark
orange brown, wrinkled towards the edges; YES, 25 C:
Weak sporulation on YES; exudate absent; soluble pigments absent; reverse buff, two concentric rings. MEA,
25 C: Sporulation on MEA strong; mycelium white turns
pale yellow at the margins; exudate absent; soluble pigments absent; reverse pale brown or orange brown towards
the centre.
Micromorphology: Sclerotia absent. Conidiophores
monoverticillate, stipe smooth, 2.5–3.5 lm wide, unbranched or with one or two branches 25–58 lm long.
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98
Fig. 64 Jahnula queenslandica (MFLU 18-1692, holotype). a Host.
b Appearance of ascomata on host. c Section of ascoma. d Periphysate
ostioler region. e Peridium. f, g Asci. h Apex of an ascus.
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i Pseudoparaphyses. j, k Apiosporous ascospores. Scale bars: c, f,
g, i = 100 lm, d = 50 lm, e = 20 lm, h, j, k = 10 lm
Fungal Diversity (2019) 96:1–242
99
Fig. 65 Phylogram generated from maximum likelihood analysis
based on combined LSU and ITS sequence data of selected taxa.
Related sequences are obtained from GenBank. Forty-nine strains are
included in the analyses, which comprise 1159 characters including
gaps. Single gene analyses are carried out and compared with each
species, to compare the topology of the tree and clade stability. The
tree is rooted with Farlowiella carmichaeliana (JX 43). Tree topology
of the ML analysis is similar to the MP and BI. The best scoring
RAxML tree with a final likelihood value of - 9016.535047 is
presented. The matrix had 607 distinct alignment patterns, with
27.68% of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.244145, C = 0.229207, G = 0.302247,
T = 0.224401; substitution rates AC = 0.875220, AG = 1.547245,
AT = 1.078795, CG = 0.865551, CT = 4.551536, GT = 1.000000;
gamma distribution shape parameter a = 0.372555. The maximum
parsimonious dataset consisted of constant 604, 385 parsimonyinformative and 170 parsimony-uninformative characters. The parsimony analysis of the data matrix resulted in the maximum of two
equally most parsimonious trees with a length of 1569 steps
(CI = 0.556, RI = 0.754, RC = 0.420, HI = 0.444) in the first tree
Maximum parsimony bootstrap (MPBT, blue) values [ 65%, Bayesian posterior probabilities (PP, green) [ 0.80% and maximum
likelihood bootstrap (ML, black) values [ 65%) are given above
the nodes. The scale bar indicates 0.07 changes. The ex-type strains
are in bold and black. The newly generated sequence is indicated in
bold and blue
Phialides ampulliform to cylindrical shaped, 3–10 per
metula,
7.5–11.5 9 2.5–3.5 lm.
Conidia
globose
2.5–3.5 lm.
Culture characteristics: The isolate grew over a wide
range of temperatures with varying growth rates on MEA,
CYA, and YES. The average growth rates of CNUFCGEU220-1 on MEA, CYA, and YES medium at 25 C
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100
Fig. 66 Kamalomyces mangrovei (MFLU 18-1691, holotype). a, b
Ascomata. c. Section of ascoma. d Section through peridium. e–g
Asci and pseudoparaphyses. h–k Ascospores. l Germinating
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ascospore. m, n Cultures on PDA (m from above view, n from
below view). Scale bars: a = 500 lm, c = 200 lm, b = 100 lm, e–
g = 50 lm, d, h–l = 20 lm
Fungal Diversity (2019) 96:1–242
were 12, 26, and 10.5 mm per 7 days, respectively. Optimal growth was observed around 25 C, slow growth was
observed at below 10 C, and no growth at 37 C.
Material examined: REPUBLIC OF KOREA, Geumsan
Park, Jeju Island, from forest soil, 27 March 2018
(CNUFC-GEU220-1, holotype), ex-type living culture
preserved as glycerol stock at Rhizophydium koreanum
80 C in the CNUFC.
GenBank numbers: CAL: MK481061, RPB2:
MK481053, MK481054, TUB2: MK481057, MK481058.
Notes: Penicillium circulare belongs to section Sclerotiora and is phylogenetically related to P. jacksonii, but
differs by the production of two concentric rings on YES.
In addition, it produces monoverticillate conidiophores
with one or two branches, in contrast to a single branch of
P. jacksonii.
Penicillium geumsanense Hyang B. Lee, P.M. Kirk &
T.T.T. Nguyen sp. nov.
Index Fungorum number: IF555412; Facesoffungi
number: FoF05794; Fig. 75
Etymology: geumsanense, referring to its collection
location, Geumsan Park, Jeju Island from where the species
was isolated (Korea).
Holotype: CNUFC-GEU2229-1
Macromorphology: CYA, 25 C: Sporulation on CYA
moderate; dark grey green; velvety floccose in centre;
mycelium white; soluble pigment absent; reverse orange to
dark orange brown; MEA, 25 C: Colonies grey green;
velvety; floccose in centre; strongly fasciculate; soluble
pigment absent; reverse dark orange brown and light yellow; YES, 25 C: Sporulation on YES strong; mycelium
white, radially sulcate; soluble pigments absent; reverse
white to dark orange brown.
Micromorphology: Sclerotia absent. Conidiophores
mostly terverticillate, sometimes mono-or biverticillate,
occasionally with a divaricate branch, branches 22 lm
long, stipe 3.0–4.5 lm wide. Metulae mostly 2–3,
9.0–17.5 9 2–4 lm. Phialides ampulliform shaped, 1–5
per metula, 7.5–10.5 9 2–3.5 lm. Conidia ellipsoidal,
3.5–4.0 9 3.0–3.5 lm.
Culture characteristics: The isolate was observed to
grow over a wide range of temperatures with varying
growth rates on MEA, CYA, and YES. The average growth
rates of CNUFC-GEU2229-1 on MEA, CYA, and YES
medium at 25 C were 6.5, 11.5, and 13.5 mm per 7 days,
respectively. Optimal growth was observed around 25 C,
slow growth was observed at below 10 C, and no growth
at 37 C.
Material examined: REPUBLIC OF KOREA, Geumsan
Park, Jeju Island, from forest soil, 27 March 2018
101
(CNUFC-GEU2229-1, holotype), ex-type living culture
preserved as glycerol stock at - 80 C in the CNUFC.
GenBank numbers: CAL: MK481062, MK481063,
RPB2: MK481055, MK481056, TUB2: MK481059,
MK481060.
Notes: Penicillium geumsanense belongs to section
Robsamsonia and is phylogenetically related to P. glandicola, a new species described below. It differs from P.
glandicola by producing terverticillate, monoverticillate or
biverticillate conidiophores, in contrast to the terverticillate
conidiophores of P. glandicola, in addition, grows more
slowly than P. glandicola on all media.
Penicillium mali-pumilae Hyang B. Lee, T.T.T. Nguyen &
Houbraken, sp. nov.
Index Fungorum number: IF555410; Facesoffungi
number: FoF05790; Fig. 76
Etymology: mali-pumilae, referring to Malus pumila, the
type was isolated from rotten fruit of apple.
Holotype: CBS H-22503
Macromorphology: CYA, 25 C: Colonies plane;
sporulation on CYA strong; colony texture fasiculate;
mycelium white; exudate present as large clear or pale
yellow droplets; soluble pigment absent; radial sulcate;
margin entire; conidia green to dull blue green; reverse
pale brown with (dark) brown in centre. YES, 25 C:
Sporulation absent, mycelium (pale) yellow in centre,
white near edge; exudate present as yellow pigments; soluble pigments absent; reverse pale brown, sometimes with
brown centre. MEA, 25 C: Sporulation on MEA strong;
colony texture fasiculate; mycelium white; exudate present
as large pale yellow or brown droplets; soluble pigments
absent; conidia green occasionally blue-green; reverse
yellow with pale brown centre or pale brown with brown
centre. DG18, 25 C: Sporulation absent or poor; texture
fasiculate; mycelium white or pale yellow, conidia dull
green; reverse (pale) yellow.
Micromorphology: Sclerotia absent. Conidiophores
mostly long, with rough walled stipes, non-vesiculate,
predominantly terverticillate, branches 20 lm long, stipe
2.5–3.5 lm wide. Metulae 4–10, 9–13 9 2.5–4 lm.
Phialides
flask
shaped,
3–8
per
metula,
8–12 9 2.5–3.5 lm. Conidia in long, distorted chains,
smooth walled or very finely roughened, globose to subglobose, 3–3.5 lm.
Colony diameters: 7 d, in mm: CYA 22–32; CYA15 C
26–31; CYA30 C 4–15; CYA37 C no growth; DG18
21–31; MEA 19–28; YES 21–32; CYAS 20–28; creatine
agar 10–20, weak growth, poor acid production, base formation absent.
Material examined: NETHERLANDS, Lilium bulb,
Aalsmeer, CBS 127.90 = IBT 12175; REPUBLIC OF
KOREA, rot apple (Malus pumila), stored in refrigerator
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102
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Fungal Diversity (2019) 96:1–242
b Fig. 67 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and TEF1-a sequence data of selected
taxa. Related sequences are obtained from GenBank. Data set
comprises 2530 characters including gaps. Single gene analyses are
carried out and compared with each species, to compare the topology
of the tree and clade stability. Tree is rooted with Botryosphaeria
agaves (MFLUCC 10-0051) and B. dothidea (CBS 115476). Tree
topology of the ML analysis is similar to the MP and BI. The best
scoring RAxML tree with a final likelihood value of - 27950.184903
is presented. The matrix had 1137 distinct alignment patterns, with
26.87% of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.234676, C = 0.258513, G = 0.266669,
T = 0.240142; substitution rates AC = 1.053687, AG = 4.072971,
AT = 3.048439, CG = 0.739670, CT = 8.539737, GT = 1.000000;
gamma distribution shape parameter a = 0.712704. The maximum
parsimonious dataset consisted of constant 1587, 768 parsimonyinformative and 175 parsimony-uninformative characters. The parsimony analysis of the data matrix resulted in the maximum of two
equally most parsimonious trees with a length of 5615 steps
(CI = 0.273, RI = 0.584, RC = 0.160, HI = 0.727) in the first tree.
Maximum likelihood bootstrap (ML, left), maximum parsimony
bootstrap (MP, middle) values [ 65%, and Bayesian posterior
probabilities (BYPP, right) [ 0.80% are given above the nodes.
The scale bar indicates 0.05 changes. The ex-type strains are in bold
and black. The newly generated sequence is indicated in bold and
blue
(4 C), Yesan, 2012, CBS 140671 = DTO 327-D1 =
EML-MP6080-1.
GenBank numbers: CAL: KP900992, ITS: KP900991,
RPB2: MK033522, TUB2: KP900993.
Notes: Penicillium mali-pumilae resembles P. psychrotrophicum and detailed notes are given in the
description of that species.
Penicillium psychrotrophicum Hyang B. Lee, H.Y. Mun,
J.C. Frisvad & Houbraken, sp. nov.
Index Fungorum number: IF555409; Facesoffungi
number: FoF05789; Fig. 77
Etymology: psychrotrophicum, refers to its ability to
grow at low temperature conditions.
Holotype: CBS H-22504
Macromorphology: CYA, 25 C: Colonies plane;
sporulation on CYA strong; colony texture fasiculate;
mycelium white; exudate present as large clear droplets;
soluble pigment absent; radial sulcate; margin entire;
conidia green; reverse pale brown, brown in centre. YES,
25 C: Sporulation poor, mycelium white; exudate absent;
soluble pigments absent; conidia dull green; reverse brown.
MEA, 25 C: Sporulation on MEA strong; colony texture
fasiculate; mycelium white; exudate present as large clear
droplets; soluble pigments absent; conidia green; reverse
yellow with pale brown centre. DG18, 25 C: Sporulation
moderate; texture fasiculate; mycelium white, conidia dull
green; reverse yellow-orange.
Micromorphology: Sclerotia absent. Conidiophores
100–200 lm long, with rough walled stipes, non-
103
vesiculate, terverticillate, branch up to 20 lm long, stipe
2.5–3.5 lm wide. Metulae 4–10, 9–15 9 2.5–4 lm.
Phialides flask shaped, 3–8 per metula, 7.5–9.5 9 2–3 lm.
Conidia in long, distorted chains, finely rough to rough,
often with spiral striations, ovoidal to fusiform, often with
a connective on one side, 2.5–3.5 lm.
Colony diameters: 7 d, in mm: CYA 27–31; CYA15 C
33–37; CYA30 C 8–12; CYA37 C no growth; DG18
31–36; MEA 23–28; YES 34–39; CYAS 29–34; creatine
agar 20–25, weak growth, poor acid production, base formation absent.
Material examined: REPUBLIC OF KOREA, Deodeok
root (Codonopsis lanceolata), Cheongyang, 2011, (CBS
H-22504, holotype), ex-type living culture (CBS
140670 = DTO 327-C9 = EML-COD3 = IBT 33673).
GenBank numbers: CAL: KP900994, ITS: KP941754,
RPB2: MK033521, TUB2: KP900995.
Notes: Penicillium psychrotrophicum belongs to series
Corymbifera and is phylogenetically related to P. tulipae,
P. radicicola and P. mali-pumilae, a new species described
below. Penicillium psychrotrophicum differs from P. tulipae and P. radicicola (and P. hirsutum) by having smaller
colonies on CYA, CYA incubated at 30 C and MEA.
Penicillium mali-pumilae is phenotypically similar but
differs by having smaller colonies on CYAS (29–34 vs
20–28 mm), DG18 (31–36 vs 21–31 mm) and CYA incubated at 15 C (33–37 vs 26–31 mm). Furthermore, P.
psychrotrophicum produces (pale) yellow mycelium on
YES, while P. mali-pumilae has white coloured mycelium
and the reverse of P. psychrotrophicum on DG18 has a
shade of orange and this is lacking in P. mali-pumilae.
The taxonomy of P. tulipae and P. radicicola needs
attention. Both species can be recognized using TUB2
sequences; however, these species can not be differentiated
using CAL sequences only. Three groups of strains are
present in the combined phylogram, though the relationship
is not resolved. A further complication is the position of a
strain (presumably) representing the type of P. virescens
Bainier (non Sopp, 1912), CBS 109553 (= NRRL 974).
This strain belongs to the P. tulipae-P. radicicola clade and
the name might thus have priority over the other two. CBS
109553 differs from both P. tulipae and P. radicicola by
having a (dark) brown reverse on CYA and CYAS and
brown soluble pigment production. P. hirsutum, another
member of series Corymbifera, also has a brown reverse on
CYA; however, these reverses are in shades of yellow–
brown or orange-brown. Sporulation on YES is poor and
this feature is shared with P. radicicola, while P. tulipae
colonies sporulate moderately or strong sporulation on
YES. Base production on CREA is shared with P. tulipae
and P. virescens; a feature absent in P. radicicola. More
research is needed to resolve the taxonomy of this clade.
123
104
Fig. 68 Thysanorea uniseptata (MFLU 18-1723, holotype). a, b
Colonies on natural substrate. c Conidiophore with conidia. d, e
Conidiogenous cells with conidia. f Conidia. g Geminated conidium.
123
Fungal Diversity (2019) 96:1–242
h, i Colony on PDA (h from above view, i from below view). Scale
bars: a = 250 lm, b = 100 lm, c = 50 lm, d, e = 10 lm, f,
g = 5 lm
Fungal Diversity (2019) 96:1–242
105
Fig. 69 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU and SSU sequence data representing
Herpotrichiellaceae. Related sequences are taken from Liu et al.
(2015), Tian et al. (2016) and Dong et al. (2018). Twenty-nine strains
are included in the combined analyses which comprise 3634
characters (674 characters for ITS, 1361 characters for LSU, 1599
characters for SSU) after alignment. Cyphellophora oxyspora (CBS
698.73) and C. sessilis (CBS 243.85) (Cyphellophoraceae,
Chaetothyriales) are used as the outgroup taxa. Single gene analyses
were carried out and the topology of each tree had clade stability.
Tree topology of the maximum likelihood analysis is similar to the
maximum parsimony analysis and the Bayesian analysis. The best
RaxML tree with a final likelihood value of - 11580.668860 is
presented. The matrix had 664 distinct alignment patterns, with
28.94% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.255728, C = 0.221018, G = 0.274376,
T = 0.248878; substitution rates AC = 1.635663, AG = 2.578277,
AT = 1.796079, CG = 1.025171, CT = 6.468570, GT = 1.000000;
gamma distribution shape parameter a = 0.513924. Maximum parsimony analysis of 3080 constant characters and 182 informative
characters resulted in one equally most parsimonious tree (CI =
0.610, RI = 0.675, RC = 0.411, HI = 0.390). Bootstrap values for
maximum likelihood (ML) and maximum parsimony (MP) equal to or
greater than 75% are placed above and below the branches
respectively. Branches with Bayesian posterior probabilities (BYPP)
equal or greater than 0.95 are in bold. The ex-type strains are in bold
and black. The newly generated sequence is indicated in bold and
blue
Penicillium wandoense Hyang B. Lee, P.M. Kirk & T.T.T.
Nguyen, sp. nov.
Index Fungorum number: IF555411; Facesoffungi
number: FoF05791; Fig. 78
Etymology: wandoense, referring to its collection location, Wan island (Wando) from where the species was
isolated (Korea).
Holotype: CNUFC-WT31-1
Macromorphology: CYA, 25 C: Sporulation on CYA
moderate; colonies radially sulcate; surface texture velutinous; mycelium white; soluble pigment absent; reverse
yellow brown to reddish brown; MEA, 25 C: Sporulation
on MEA strong; colonies pale green; margins low, wide,
irregular; velvety to floccose texture; reverse pale red or
pale yellow; YES, 25 C: Sporulation poor, mycelium
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106
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Fig. 70 Phylogram generated from maximum likelihood analysis
based on combined TUB2, CAL, and RPB2 sequence data for
Penicillium circulare and related species within the sect. Sclerotiora.
Penicillium levitum (CBS 345.48) was used as outgroup taxon.
Bootstrap values for maximum likelihood (ML) greater than 50% are
placed above the branches. Type strains are in bold. The newly
generated sequences are indicated in blue
Fig. 71 Phylogram generated from maximum likelihood analysis
based on combined TUB2, CAL, and RPB2 sequence data for
Penicillium geumsanense and related species within the sect. Robsamsonia. Penicillium turbatum (CBS 237.60) was used as outgroup
taxon. Bootstrap values for maximum likelihood (ML) greater than
50% are placed above the branches. Type strains are in bold. The
newly generated sequences are indicated in blue
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107
Fig. 72 Phylogram generated from maximum likelihood analysis
based on combined TUB2, CAL, and RPB2 sequence data for
Penicillium psychrotrophicum and P. mali-pumilae and related
species within the sect. Fasciculata series Corymbifera. Penicillium
chrysogenum (CBS 306.48) was used as outgroup taxon. Bootstrap
values for maximum likelihood (ML) equal to or greater than 70% are
placed above the branches. Branches with Bayesian posterior
probabilities (BYPP) equal to or greater than 0.95 are in bold. Type
strains are in bold. The newly generated sequences are indicated in
blue
white; surface texture velutinous; reverse yellow brown to
reddish brown.
Micromorphology: Sclerotia absent. Conidiophores
arising from agar surface, predominantly biverticillate, but
also many monoverticillate, branch up to 26.5 lm long,
stipe
2.5–3.5 lm
wide.
Metulae
2–4,
9.5–15.5 9 2.5–4.0 lm. Phialides ampulliform shaped,
2–5 per metula, 6.5–9.5 9 2–3.5 lm. Conidia subglobose
to ellipsoidal, 3–4 9 2.5–3.5 lm.
Culture characteristics: The isolate grew over a wide
range of temperatures with varying growth rates on MEA,
CYA, and YES. The average growth rates of CNUFCWT31-1 on MEA, CYA, and YES medium at 25 C were
34.5, 23, and 21 mm per 7 days, respectively. Optimal
growth was observed around 25 C, slow growth was
observed at below 10 C, and at 37 C.
Material examined: REPUBLIC OF KOREA, Wando
island (34190 1.2000 N 126450 0.0000 E), from freshwater, 24
August 2018 (CNUFC-WT31-1, holotype), ex-type living
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108
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Fig. 73 Phylogram generated from maximum likelihood analysis
based on combined TUB2, CAL, and RPB2 sequence data for
Penicillium wandoense and related species within the sect. LanataDivaricata. Penicillium glabrum (CBS 125543) was used as outgroup
taxon. Bootstrap values for maximum likelihood (ML) greater than
50% are placed above the branches. Type strains are in bold. The
newly generated sequences are indicated in blue
culture preserved as glycerol stock at - 80 C in the
CNUFC.
GenBank numbers: CAL: MK080564, MK080565,
RPB2: MK080566, MK080567, TUB2: MK080562,
MK080563.
Notes:
Penicillium
wandoense
differs
from
P. araracuarense, and P. wotroi by its pale red or pale
yellow reverse on MEA. In the phylogenetic tree based on
multiple genes, the strain formed a separate branch from
other species of the genus and is considered to represent a
new species.
the Laboulbeniales, while the two new orders include oly a
few of the known species (less than 3%).
Laboulbeniomycetes Engl.
Laboulbeniales Lindau
Notes: The order Laboulbeniales includes about 2100
described species of obligate ectosymbionts on Arthropods.
The Laboulbeniales sensu Lindau comprehended all the
Laboulbeniomycetes, but recently the class has been
splitted in three orders as a result of molecular analysis
(Goldmann and Weir 2018; Haelewaters et al. 2019).
However, the great majority of species is still classified in
123
Laboulbeniaceae G. Winter
Notes: A full taxonomic treatment of the family is more
than 30 years old (Tavares 1985), but this classification has
been recently questioned in the papers reported above.
Although a new comprehensive taxonomy the group has
not yet been advanced, the two genera considered below
can still be safely included in the family Laboulbeniaceae.
Prolixandromyces R. K. Benjamin 1970, Aliso 7: 174 [MB
4369]
= Monandromyces R. K. Benjamin 1999, Aliso 18: 72
[MB 28387] syn. nov.
Notes: The genus Monandromyces was erected by R.K.
Benjamin in 1999 to accommodate Autophagomyces
microveliae Thaxt. and ten new species. The given diagnosis (page 72) is as follows: ‘‘Receptacle consisting of
three in-line superposed cells (I, II, III) bearing distally on
one side a stalked perithecium and on the other side a
simple, free appendage; basal (I) and suprabasal (II) cells
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109
Fig. 74 Penicillium circulare (CNUFC-GEU220-1, holotype). a, d Colonies in Czapek yeast agar (CYA). b, e Colonies in malt extract agar
(MEA). c, f Colonies in yeast extract sucrose agar (YES). g–m Conidiophores. n Conidia. Scale bars: g–m = 20 lm, n = 10 lm
strongly obliquely superposed; cell II subtending the
perithecium; terminal cell (III) subtending the appendage.
Appendage consisting of three superposed cells and a
single, terminal, simple, flask-shaped antheridium; basal
cell small, broader than long to nearly isodiametric; median
cell elongate, several times longer than broad; terminal cell
small, nearly isodiametric or ± elongate, always less than
length of median cell; antheridium flask shaped with an
123
110
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Fig. 75 Penicillium geumsanense (CNUFC-GEU2229-1, holotype). a, d Colonies in Czapek yeast agar (CYA). b, e Colonies in malt extract
agar (MEA). c, f Colonies in yeast extract sucrose agar (YES). g–m Conidiophores. n Conidia. Scale bars: g–m = 20 lm, n = 10 lm
elongate, slender efferent tube. Perithecium with two stalk
cells (VI, VII), three persistent basal cells (m, n, n’), and
four vertical rows of outer wall cells of five cells
each. ….’’.
123
The recent description of four new species in the genus
Prolixandromyces (Kaishian and Weir 2018) and the consequent emendation of the genus, now including also a
species with a single, terminal antheridium (i.e.
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111
Fig. 76 Penicillium mali-pumilae (EML-MP6080-1, ex-type culture). a 7 d old cultures, 25 C, left to right; first row, all obverse,
Czapek yeast agar (CYA), yeast extract sucrose agar (YES), dichloran
18% glycerol agar (DG18), malt extract agar (MEA); second row,
CYA reverse, YES reverse, DG18 reverse, CREA obverse. b–f
Conidiophores. g Conidia. Scale bars: b–g = 10 lm
P. bromelicola A. Weir & Kaishian), removes any difference between Monandromyces, (characterized by a single
antheridium) and Prolixandromyces. In the emended
diagnosis (page 223) is written: ‘‘The uppermost cell in the
series either producing a single, simple, terminal
antheridium in the 3-celled appendage or more frequently
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112
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Fig. 77 Penicillium psychrotrophicum (EML-COD3, ex-type culture). a 7 d old cultures, 25 C, left to right; first row, all obverse,
Czapek yeast agar (CYA), yeast extract sucrose agar (YES), dichloran
18% glycerol agar (DG18), malt extract agar (MEA); second row,
CYA reverse, YES reverse, DG18 reverse, CREA obverse. b–f
Conidiophores. g Conidia. Scale bars: b–g = 10 lm
in the 4–11-celled appendage giving rise to both a terminal
and a lateral antheridium’’. All the other features are
identical, as it is identical the family of most of the host
insects: the Veliidae (semiaquatic bugs). As a consequence
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113
Fig. 78 Penicillium wandoense (CNUFC-WT31-1, holotype). a, d Colonies in Czapek yeast agar (CYA). b, e Colonies in malt extract agar
(MEA). c, f Colonies in yeast extract sucrose agar (YES). g–m Conidiophores. n Conidia. Scale bars: g–m = 20 lm, n = 10 lm
of the new synonymy all the species of Monandromyces
are transferred to the genus Prolixandromyces with the
following new combinations.
Prolixandromyces australis (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces australis R.K. Benjamin 1999, Aliso
18: 78 [MB 459855]; Index Fungorum number: IF556540
123
114
Prolixandromyces elongatus (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces elongatus R.K. Benjamin 1999,
Aliso 18: 86 [MB 459861]; Index Fungorum number:
IF556541
Prolixandromyces falcatus (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces falcatus R.K. Benjamin 1999, Aliso
18: 77 [MB 459854]; Index Fungorum number: IF556542
Prolixandromyces longispinae (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces longispinae R.K. Benjamin 1999,
Aliso 18: 87 [MB 459862]; Index Fungorum number:
IF556543
Prolixandromyces microveliae (Thaxter) W. Rossi, comb.
nov.
: Autophagomyces microveliae Thaxter 1931, Mem.
Amer. Acad. Arts Sci. 16: 96
= Monandromyces microveliae (Thaxter) R.K. Benjamin
1999, Aliso 18: 74 [MB 459852]; Index Fungorum number:
IF556544
Prolixandromyces neoalardi (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces neoalardi R.K. Benjamin 1999,
Aliso 18: 82 [MB 459859]; Index Fungorum number:
IF556545
Prolixandromyces polhemorum (R.K. Benjamin) W.
Rossi, comb. nov.
: Monandromyces polhemorum R.K. Benjamin 1999,
Aliso 18: 79 [MB 459856]; Index Fungorum number:
IF556546
Prolixandromyces protuberans (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces protuberans R.K. Benjamin 1999,
Aliso 18: 81 [MB 459858]; Index Fungorum number:
IF556547
Prolixandromyces pseudoveliae (R.K. Benjamin) W.
Rossi, comb. nov.
: Monandromyces pseudoveliae R.K. Benjamin 1999,
Aliso 18: 84 [MB 459860]; Index Fungorum number:
IF556548
Prolixandromyces tenuistipitis (R.K. Benjamin) W. Rossi,
comb. nov.
: Monandromyces tenuistipitis R.K. Benjamin 1999,
Aliso 18: 80 [MB 459857]; Index Fungorum number:
IF556549
Prolixandromyces umbonatus (R.K. Benjamin) W. Rossi,
comb. nov.
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: Monandromyces umbonatus R.K. Benjamin 1999,
Aliso 18: 74 [MB 459853]; Index Fungorum number:
IF556550
Stigmatomyces H. Karst., Chemismus Pfl.-Zelle: 78 (1869)
Notes: After the recent description of Stigmatomyces
coronatus (Rossi and Bernardi 2018), the accepted species
are 148 distributed in all continents, all occurring on flies
(Diptera) with the only exceptions being S. caudicola Speg.
and S. euconni F. Picard, two species that certainly do not
belong to the genus Stigmatomyces. Stigmatomyces succini
is the only known fossil species of the Laboulbeniales
reported so far (Rossi et al. 2005). Very few sequences are
available for species in the genus.
Stigmatomyces chamaemyiae W. Rossi & M. Leonardi, sp.
nov.
Index Fungorum number: IF828160; Facesoffungi
number: FoF05175; Fig. 79
Etymology: From the name of the genus of the host
insects, Chamaemyia.
Holotype: FI WR3847
Receptacle, distal portion of the free appendage ad
perithecial apex almost hyaline; stalk and basal cell of the
free appendage and perithecial venter rusty or amber yellow;
the rest of the fungus is very diluted reddish or amber yellow.
Receptacle very variable in length, slightly tapering to the
foot, with the suprabasal cell longer than the basal. Stalk cell
of the appendage relatively small, distally prominent,
strongly tapered below, its outer margin almost straight. Free
appendage usually turned sidewise, with the axis consisting
of five cells (rarely four), each bearing two anteridia, the
uppermost of which is spinose. Stalk cell of the perithecium
distinctly broader than long, separated by the secondary
stalk-cell by a concave septum; the cells above it subequal in
size and shape, the two outer of which are more or less
prominent. Perithecial venter finely granular, oblong,
slightly and symmetrically inflated, passing without abrupt
transition to the spreading base of the paler and longer neck,
which is slightly curved dorsally and slightly swollen at its
junction with the tip; the latter is asymmetrical, with the
ventral side more convex; the apex short, rather abruptly
narrower, distinctly turned towards the dorsal side, ending in
four, small, paired, subtruncate lips. Total length
205–440 lm; perithecium 45–75 9 130–220 lm; free
appendage 50–65 lm; ascospores 30–35 lm.
Material examined: MALTA, Ghajn Tuffieha Bay, sea
level, 1 May 2001, B. Merz, on legs and apex abdomen of
Chamaemyia flavicornis (Strobl) (Diptera, Chamaemyidae)
(FI WR3533 & WR3534, paratypes). FRANCE, Pyrénées
Orientales, La Bastide, Col de Palomère, high grassland, 10
June 2007, J. C. Deeming, on legs of C. juncorum (Fallén)
(FI WR3142, paratype). PORTUGAL, Monte Gordo, V.
Fungal Diversity (2019) 96:1–242
115
Fig. 79 Stigmatomyces
chamaemyiae (FI WR3847,
holotype). Scale bar = 50 lm
R. S. Antonio, 11 May 2012, R. Andrade, on various parts
of C. flavipalpis (Haliday) (FI WR3848, paratype); Sintra,
Almargem do Bispo, 21 August 2013, R. Andrade, on
various parts of C. polystigma (Meigen) (FI WR3847,
holotype).
Notes: The thalli on the tergites of C. flavipalpis from
Portugal are very long (up to 590 lm) because of a very
long receptacle (up to 350 lm). Stigmatomyces chamaemyiae is the first species of the Laboulbeniales reported on
Chamaemyidae. Among the described species, the species
most similar to S. chamaemyiae seems to be S. aphaniosomae W. Rossi & A. Weir occurring on Chyromyidae. In
the latter species, however, the perithecial apex is very
different, the free appendage is shorter, with the axis
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consisting of three cells, and the basal cell of the receptacle
is usually longer than the suprabasal (Rossi and Weir
2011). Twenty-four species of Stigmatomyces are recorded
from Portugal, while eleven are from France; to date, none
of the species of Stigmatomyces have been reported from
Malta (Castaldo et al. 2004; Rossi et al. 2013).
Stigmatomyces cocksii W. Rossi & M. Leonardi, sp. nov.
Index Fungorum number: IF828161; Facesoffungi
number: FoF05176; Fig. 80
Etymology: Named after the Australian entomologist
Graeme Cocks, who collected the new species.
Holotype: CAMB WR4074a
Thallus grayish yellow: the perithecial venter, the stalk
cell and lower cell of the appendage are darker, the basal
cell of the receptacle (cell I) and the free appendage are
paler, while the suprabasal cell of the receptacle (cell II) is
hyaline or almost so. Receptacle very variable in length;
basal cell regularly tapering to the foot and rather abruptly
constricted at the septum with the suprabasal, which is of
nearly uniform diameter although variably elongate. Stalk
cell of the appendage stout, broadly heart-shaped, distally
prominent. Free appendage reaching the summit of the
perithecial venter, with the axis consisting of four cells, the
lower of which is about twice longer than broad, the others
slightly broader than long and gradually smaller; each axial
cell bears two antheridia except the uppermost, which bears
a third, terminal antheridium. Perithecial stalk and basal
cells relatively small, not strongly unequal in size, distinctly prominent externally. Perithecial venter regularly
elliptical in outline, its surface distinctly warty, the wartlike elevations disposed transversely forming darker ridges;
the neck abruptly narrower, longer than the venter, almost
isodiametric, transversely striate, slightly enlarged at its
junction with the tip, which tapers gradually to a subtruncate apex. Total length 175–345 lm; perithecial venter
38–62 9 40–58 lm; perithecial neck, up to155 lm; free
appendage 48–57 lm; ascospores 29–31 lm.
Material examined: AUSTRALIA, Queensland, Hermit
Park, 44 Marks St., Malaise trap, 23 December 2015, G.
Cocks, on Notiochyromya sp. nova (Diptera, Chyromyidae)
(CAMB WR4074a, holotype; FI WR4074a, isotype),
Same data as the type, 20 November 2015 (FI WR4073,
paratype).
Notes: Because of the transverse ridges formed by wartlike elevations, Stigmatomyces cocksii bears a superficial
resembles to S. micrandrus Thaxt. and S. rugosus Thaxt.,
both parasitic on shore flies (Ephydridae). However, in the
two latter species these ridges are smaller and more
numerous, the appendage is distinctly shorter than the
perithecial venter, the apex of perithecium bearing relatively large with diverging lips (Thaxter 1908).
123
Fig. 80 Stigmatomyces cocksii (CAMB WR4074a, holotype). Scale
bar = 50 lm
The only previously described species of Stigmatomyces
on flies of the family Chyromyidae is S. aphaniosomae.
The latter is very different from S. cocksii: among other
distinguishing features, it has a smooth surface of the
perithecial venter or almost so, a shorter perithecial neck, a
shorter free appendage, the perithecial apex bearing
Fungal Diversity (2019) 96:1–242
unequal and diverging lips (Rossi and Weir 2011). The
identified species of Stigmatomyces reported from Australia thus far are three: S. limnophorae Thaxt., S. rugosus
Thaxt. (Rossi et al. 2013), and S. zaleae W. Rossi & A.
Weir (Rossi and Weir 2007).
Stigmatomyces papei W. Rossi & M. Leonardi, sp. nov.
Index Fungorum number: IF828162; Facesoffungi
number: FoF05178; Fig. 81
Etymology: Named after the Danish dipterologist Thomas Pape, who collected the new species.
Holotype: FI WR4044
Perithecium pale reddish brown; the rest of the thallus is
almost hyaline or faintly tinged with reddish brown, with
the exception of the lower cell of the free appendage which
stands out with brilliant rusty red. Receptacle rather short
and stocky, tapering throughout, the basal cell subtriangular with its content contracted above, the suprabasal
distinctly longer. Stalk-cell of the appendage relatively
long and narrow, distally prominent. Free appendage
slender and attenuate, straight or sigmoid, its axis usually
117
consisting of 9 cells, each bearing a pair of slightly
diverging antheridia arranged in two rows, except for the
uppermost cell, which bears three superimposed antheridia.
Stalk-cell of the perithecium large, longer than broad, its
outer margin overlapped for most of its length by the
secondary stalk-cell, which is only slightly smaller.
Perithecial basal cells subequal, large, longer than broad.
Perithecium longer than the rest of the thallus, slender and
elongate, the venter oblong, the neck hardly distinguished,
about as long as the venter, very slightly tapering towards
the tip and ending with two large and unequal outgrowths,
of which the shorter is broadly ovoid, the other beanshaped and bending outwards. Total length 575–640 lm;
perithecium 60–70 9 340–380 lm; free appendage
105–140 lm; ascospores 55 lm.
Material examined: TANZANIA, Morogoro Region,
Udzungwa Mts. Natl. Park, Mizimu Camp, 769 m, 30–31
July 2015, T. Pape, N. Scharff & al., on tergites of Plagiostenopterina submetallica (Loew) (Diptera, Platystomatidae) (FI WR4044, holotype).
Fig. 81 Stigmatomyces papei
(FI WR4044, holotype).
a Whole thallus. b Upper
portion of a syntype. Scale bars:
a, b = 50 lm
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Notes: A few thalli occurring on the upper surface of the
right wing of the host are smaller and stockier, with the
larger of the perithecial outgrowths longer and slenderer.
The two large perithecial outgrowths are unique in the
genus and make Stigmatomyces papei distinguishable at
first sight. It is likely allied with S. ortalidanus Thaxt.
described 100 years ago on Plagiostenopterina vicaria
Hendel (= P. nigripes Enderlein) from the Cameroons and
never reported since, with which it shares the structure of
the free appendage, the shape of the perithecial venter and
the elongation of the perithecial basal cells. However, the
latter fungus lacks the two peculiar perithecial outgrowths,
it is much longer (up to 1 mm), with very long suprabasal
cell and perithecial stalk-cell, wall cells of the perithecium
spirally twisted, and surface of the perithecial neck distinctly corrugated (Thaxter 1931).
Stigmatomyces papei is the first species of Stigmatomyces reported thus far from Tanzania.
Stigmatomyces tschirnhausii W. Rossi & M. Leonardi, sp.
nov.
Index Fungorum number: IF828163; Facesoffungi
number: FoF05177; Fig. 82
Etymology: Named after the German dipterologist
Michael von Tschirnhaus.
Holotype: FI WR3542
Receptacle, distal portion of the free appendage ad
perithecial apex almost hyaline; the rest of the fungus is
unequally colored with diluted rusty red, with the
perithecial venter and lower cell of the free appendage
distinctly darker. Receptacle tapering throughout, the basal
cell abruptly narrow just above the foot, the suprabasal
longer, the horizontal septum dividing the two cells associated with a slight constriction. Stalk-cell of the appendage rather short and stout, broadly heart-shaped, very
prominent distally. Axis of the appendage consisting of
four cells, the lower distinctly larger and darker, about
twice as long as broad, the others gradually smaller, each
bearing a pair of relatively large antheridia with strongly
curved necks, the uppermost of which is spinose. Stalk-cell
of the perithecium flattened and subtriangular; the cells
above distinctly prominent externally. Perithecial venter
relatively short and stout, symmetrically inflated, the spiral
wall cells separated by thin furrows; the neck distinctly
longer, almost isodiametric above the slightly spreading
base; the tip very slightly bent and tapering; the lips
strongly unequal, two being short, flattened and divergent,
the other two elongate, forming together a tooth-like,
suberect prolongation. Total length 280–400 lm; perithecium 50–70 9 175–225 lm; free appendage 55–70 lm.
Material examined: KENYA, Western Region, Kakamega District, Kakamega Forest, Kisere Forest, about
20 km NE’ Kakamega, about 90 km SSE’ Mount Elgon
123
Fig. 82 Stigmatomyces tschirnhausii (FI WR3542, holotype). Scale
bar = 50 lm
summit, 1597 m asl, forest, Malaise trap, 00230 0700 N,
34530 1600 E, 16 April–16 May 2003, M. Kraemer, on
sternites near apex abdomen, antennae and legs of Dasiops
sp. (Diptera, Lonchaeidae) (FI WR3542, holotype).
Notes: The perithecial apex of Stigmatomyces tschirnhausii bearing two pairs of lips of very unequal length is
reminiscent of the apex of S. dacinus Thaxt. The latter,
occurring on Tephritidae, shares with the new species also
the spirally twisted wall-cells of the perithecial venter, but
is quite different in other features, such as the very long
Fungal Diversity (2019) 96:1–242
stalk-cell of the appendage, the free appendage longer with
the axis consisting of a higher number of cells, the basal of
which is sterile, the twist of the perithecial venter extending to the neck (Thaxter 1931). Stigmatomyces tschirnhausii is the first species of this genus reported on flies of
the family Lonchaeidae. The species of Stigmatomyces
reported from Kenya to date are four: S. rugosus (Rossi
119
1988), S. limnophorae (Rossi et al. 2013), S. limosinae
Thaxt. and S. dacinus (Rossi and Leonardi 2018).
Stigmatomyces vikhrevii W. Rossi & M. Leonardi, sp. nov.
Index Fungorum number: IF828164; Facesoffungi
number: FoF05179; Fig. 83
Fig. 83 Stigmatomyces
vikhrevii (FI WR3694,
paratype). Scale bars = 50 lm
123
120
Etymology: Named after the Russian dipterologist Nikita
Vikhrev, who collected the new species.
Holotype: FI WR3691
Receptacle almost hyaline; the rest of the fungus is
unequally colored with diluted rusty red, with the
perithecial venter and lower cell of the free appendage
distinctly darker. Receptacle rather stout, nearly uniform,
with the basal cell abruptly tapered at the base and the
suprabasal longer, sometimes slightly broader above. Stalkcell of the appendage short and stout, its outer margin more
or less convex. Free appendage also short and stout, with
the axis composed of three cells, the lower distinctly darker, each producing a pair of large antheridia, the uppermost of which bears a tiny spine. Stalk-cell of the
perithecium much compressed, the secondary stalk-cell
larger, with lower and outer convex margins. Perithecial
venter elliptical in outline, its wall cells very slightly spiral
with a verruculose surface, the verrucosity disposed more
or less transversely; passing without abrupt transition to the
base of the paler, smooth and shorter neck; the latter erect,
its margins very slightly concave, distally broader at its
junction with the stout, tapering tip; the subtruncate apex
ends in four lips, three of which are very small, subequal
and flattened, the fourth rounded and more prominent.
Total
length
210–415 lm;
perithecium
45–60 9 140–220 lm; free appendage 50–55 lm; ascospores 40–45 lm.
Material examined: MALAYSIA, Borneo, Sabah, Kota
Kinabalu, 5.99 N, 116.09E, 26–30 December 2011, N.
Vikhrev, on the tergites near the apex of the abdomen of
Discomyza maculipennis (Wiedemann) (Diptera, Ephydridae) (FI WR3691, holotype; FI WR3692, WR3693,
WR3694, paratype).
Notes: Stigmatomyces vikhrevii bears a superficial
resembles to S. compressus Thaxt. reported on Jamaican
shore-flies, which however has a more slender habit, a
different perithecial apex, and a much longer free appendage (Thaxter 1931).
The species of Stigmatomyces reported thus far from
Malaysia are seven. These are S. ilytheae Thaxt. (Thaxter
1917), S. dacinus Thaxt., S. limosinoides Thaxt., S. tortimasculus Thaxt. (Thaxter 1918), S. sygmoideus Thaxt.,
S. subinflatus Thaxt. (Thaxter 1931) and S. neurochaetae
W. Rossi et A. Weir (Rossi and Weir 2007). Actually,
S. subinflatus was described twice: in the first description
the host (Drosophila sp.) is reported from Cameroon
(Thaxter 1918, p. 740), while in the second the collecting
place was transferred to ‘‘Sarawak, Borneo’’ (Thaxter
1931, p. 134). The right place seems the second because
this is the one reported on the label of the type slide of
S. subinflatus (Thaxter no. 2180 = FH 6409).
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Lecanoromycetes O.E. Erikss. & Winka
Notes: This class represents most of the lichenized
members within Ascomycota (Lücking et al. 2017). However, it also includes some non-lichenized ascomycetes
(Kirk et al. 2008). Most of the taxa are characterised by
apothecial ascomata, while a few form perithecia (Ekanayaka et al. 2017).
Subclass Lecanoromycetidae
Notes: The subclass Lecanoromycetidae with in Lecanoromycetes was established by Hibbett et al. (2007). This
subclass comprises Caliciales, Lecanorales, Lecideales,
Leprocaulales, Peltigerales, Rhizocarpales and Teloschistales (Jaklitsch et al. 2016).
Caliciales Bessey
Notes: This order includes both mazaediate and nonmazaediate genera of both crustose, fruticose and foliose
lichenized genera in various habitats (Jaklitsch et al. 2016;
Ekanayaka et al. 2017). However the taxa in this order are
highly varied in their morphological characters.
Caliciaceae Chevall.
Notes: This family was established by Chevallier (1826)
and currently comprises 29 genera and 630 species (Jaklitsch et al. 2016; Wijayawardene et al. 2018a). Taxa are
mostly lichenized and form crustose to squamulose,
foliose, or fruticose lichen thalli. Ascomata are mostly
apothecial (Jaklitsch et al. 2016). Species are widely distributed in temperate, subtropical, and tropical regions,
especially on bark, rocks and wood (Jaklitsch et al. 2016).
Buellia De Not.
Notes: The lichen genus Buellia was introduced by De
Notaris (1846). Currently this genus includes around 400
species. The genus is characterized by black lecideine
apothecia, septate, oblong to ellipsoid, rarely citriform,
hyaline to brownish ascospores and a deep reddish brown
to yellow or yellowish green to rarely hyaline hypothecium
(Joshi et al. 2010).
Buellia viridula Ekanayaka & K.D. Hyde, sp. nov.
Index Fungorum number: IF555467; Facesoffungi
number: FoF05082; Fig. 84
Etymology: The specific epithet viridula refers to the
pigment colour in hymenium layer of apothecia.
Holotype: MFLU 16-0587
Saprobic on dead stems. Sexual morph Apothecia 300–
350 9 110–160 lm (
x = 331.7 9 132.6 lm, n = 10), arising singly, erumpent, sessile, black when fresh. Receptacle
pulvinate, black. Disc flat to slightly convex, black. Margins
black. Excipulum 22–38 lm (
x = 31.5 lm, n = 10) wide at
margins and flanks, composed of black cells of textura
angularis. Hymenium blackish green. Paraphyses 1–3 lm
wide (
x = 2.1 lm, n = 20), numerous, branched, septate.
Fungal Diversity (2019) 96:1–242
121
Fig. 84 Buellia viridula (MFLU 16-0587, holotype). a Substrate. b,
c Ascomata on wood. d Cross section of an ascoma. e Close up of the
cross section of ascoma at margins. f Cylindrical paraphyses. g–j
Cylindric-clavate asci. k–n Obvoid ascospores. Scale bars:
b = 500 lm, d = 150 lm, c = 100 lm, e = 60 lm, f = 20 lm, g–
j = 10 lm, k–n = 5 lm
Asci 24–50 9 8–14 lm (
x = 34.3 9 10.4 lm, n = 30), 8spored, globose, long stipitate, apex rounded, inoperculate,
amyloid ring absent at the apex. Ascospores 7–11 9 3–4 lm
(
x = 9.5 9 3.7 lm, n = 40), multiseriate, obovoid, immature spore hyaline, greenish brown at maturity, smooth-walled, 1-septate, guttules present. Asexual morph
undetermined.
Material examined: THAILAND, Chiang Rai Province,
Doi Mae Suai, on dead stems, 22 June 2015, A.H. Ekanayaka, HD0026 (MFLU 16-0587 holotype; HKAS
104247, isotype).
GenBank numbers: ITS: MK075947 (MFLU 16-0587a), MK075948 (MFLU 16-0587-b).
Notes: Buellia viridula is characterized by sessile black
apothecia, a blackish excipulum, a blackish green
123
122
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Fungal Diversity (2019) 96:1–242
b Fig. 85 Phylogram generated from a maximum likelihood analysis
based on ITS sequence data. The newly generated nucleotide
sequences are compared against the GenBank (http://www.ncbi.nlm.
nih.gov/) database using the Mega BLAST program. Related
sequences were obtained from GenBank. Fifty-eight strains were
included in the sequence analyses, which comprised 655 characters
including gaps. Calicium abietinum (MP40) is used as the outgroup
taxon. The best scoring RAxML tree with a final likelihood value of
- 9417.010515 is presented. The matrix had 460 distinct alignment
patterns, with 21.50% of undetermined characters or gaps. Estimated
base frequencies were as follows; A = 0.223, C = 0.287, G = 0.258,
T = 0.232; substitution rates AC = 1.600208, AG = 2.778296,
AT = 1.882342, CG = 0.827224, CT = 6.521407, GT = 1.000000;
gamma distribution shape parameter a = 0.407728. Bootstrap support
values for ML equal or greater than 50% are given above the nodes.
The newly generated sequences are in blue
hymenium, globose asci and ellipsoid greenish ascospores.
Buellia viridula is similar to B. ocellata and B. jugorum by
having coloured hymenium, but the latter two species have
ornamented ascospores (Scheidegger 1993). The phylogenetic analysis clearly establishes B. viridula as a new
species (Fig. 85).
Lecanorales Nannf.
Notes: This order was established by Nannfeldt (1932).
Taxa form lichen thalli with protococcoid green photobionts (Crespo et al. 2010; Miadlikowska et al. 2014). The
members are widely distributed (Kirk et al. 2008; Ekanayaka et al. 2017).
Lecanoraceae Körb.
Notes: This family was established by Körber (1855).
The members of the family are widely distributed and
commonly found on rocks, soil or bark (Kalb et al. 2011).
Species are characterised by apothecial ascomata, sparingly
branched paraphyses, semifissitunicate cylindrical to clavate asci, and ellipsoid, subglobose or bacilliform, hyaline
ascospores (Kalb et al. 2011).
Lecidella yunnanensis Ekanayaka, & K.D. Hyde, sp. nov.
Index Fungorum number: IF555468; Facesoffungi
number: FoF05083; Fig. 86
Etymology: The specific epithet yunnanensis refers to
the province where type specimen collected.
Holotype: MFLU 17-1910
Saprobic on dead stems. Sexual morph 300–
500 9 150–200 lm
(
x = 368 9 181 lm,
n = 10).
Apothecia arising singly or in small groups, sessile, slightly
erumpent from the substrate, pulvinate, black with shiny
appearance. Receptacle convex. Margin distinct, black.
Hymenium upper part dark brown and lower part hyaline,
within a thick gelatinous matrix. Ectal excipulum composed of small, thin-walled, blackish cells of textura
angularis, with algal cells. Medullary excipulum composed
of narrow, long, thin-walled, hyaline cells of textura
123
intricata. Paraphyses 1–2 lm wide (
x = 1.6 lm, n = 20),
numerous, filiform, propoloid, septate, slightly swollen at
the apices, apices are glued together to develop a pseudoepithecium. Asci 40–50 9 9–13 lm (
x = 45 9 12 lm,
n = 30), 8-spored, cylindric–clavate, narrowed to the base,
short pedicellate, rounded at the apex, amyloid ring absent
at the ascus apex, croziers present at the base of asci. Ascospores 10–15 9 5–6 lm (
x = 12 9 5.4 lm, n = 40),
hyaline when immature, greenish brown at maturity,
smooth walled, ellipsoid, 1-septate, guttulate.
Material examined: CHINA, Kunming, Yunnan Province, 25 April 2016, A.H. Ekanayaka, HC 001, HC004
(MFLU 17-1910, holotype).
GenBank numbers: ITS: MK075945, LSU: MK075949
(MFLU 17-1910-a); ITS MK075946, LSU MK075950
(MFLU 17-1910-b).
Notes: Lecidella yunnanensis is characterized by black,
sessile, pulvinate apothecia, with a black disk with a shiny
appearance, propoloid septate paraphyses, cylindric–clavate asci and ellipsoid ascospores. Lecidella yunnanensis is
similar to Lecidella enteroleucella, however L. enteroleucella differs from L. yunnanensis by having black, greenish
to reddish brown to olive disks and a hyaline hymenium
layer (Nash et al. 2004). In the phylogenetic analysis, L.
yunnanensis groups clearly distinct from known species of
Lecidella (Fig. 87).
Pilocarpaceae Zahlbr.
Notes: Pilocarpaceae is a family of mainly crustose
lichens, including mostly foliicolous lineages, plus the bulk
of species currently classified in the genus Micarea, previously placed in a separate family Micareaceae (Andersen
and Ekman 2004, 2005; Miadlikowska et al. 2014).
Micarea Fr.
Notes: The genus Micarea itself is highly heterogeneous, with several lineages falling outside the genus in its
proper sense and even outside the Pilocarpaceae; the
monophyletic core group also possibly represents more
than one genus (Czarnota and Guzow-Krzemińska 2010;
Schmull et al. 2011; Ekman and Svensson 2014). Here, we
introduce a new species with squamulose thallus that forms
an isolated lineage within the monophyletic Micarea core
clade (Fig. 89). It might well represent a separate genus,
but further data are required to resolve this entire clade
taxonomically.
Micarea squamulosa Aptroot, Lücking & M. Cáceres, sp.
nov.
Index Fungorum number: IF555463; Facesoffungi
number: FoF05966; Fig. 88
Etymology: Referring to the squamulose thallus.
Holotype: M. E. S. Cáceres & A. Aptroot ISE 40719
(ISE)
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Fig. 86 Lecidella yunnanensis (MFLU 17-1910, holotype). a Substrate. b Ascomata on wood. c Cross section of an ascoma. d Close up
of a vertical section of the ascoma at margin. e Aseptate paraphyses.
f–i Short pedicellate asci. j–m Ovate ascospores. Scale bars: c,
d = 100 lm, e = 20 lm, f–i = 15 lm, j–m = 6 lm
Terricolous squamulose Micarea with loose to agglutinated, olive-green lobes which are flat to terete and solid to
hollow, without rhizines; apothecia immarginate, dark
brown, ascospores hyaline, ellipsoid, 11–12 9 6–6.5 lm.
Thallus squamulose, terricolous, extending up to 2 cm
diam., consisting of loose to crowded and occasionally
agglutinated, simple to plicate or lobate flat to ascending
squamules of c. 0.4–1.2 mm diam. Squamules entire to
shallowly incised, partly saddle-shaped to erect, olive
green, not pruinose, smooth but not shiny; margins rather
thick and rounded, of the same colour as the upper surface;
lower surface dull, pale olive green, without rhizines;
medulla whitish, often becoming hollow, up to 300 lm
thick; marginal lobes more often flat, solid and elongate
than the central lobes which are more often erect and
hollow. Photobiont green. Ascomata apothecia, superficial
on the thallus, solitary and centrally on the squamules,
round to irregular in outline, 0.2–0.6 mm diam.; disc dark
brown to dark grey, convex, dull; margin c. 0.1 mm wide,
dark brown (paler inside), glossy, not visible from above.
Hymenium hyaline, c. 60 lm high; epihymenium olive
brown; excipulum dark brown outside, hyaline inside;
hypothecium contiguous with the excipulum, hyaline. Paraphyses rather agglutinated. Asci cylindrical, with 8
ascospores.
Ascospores
hyaline,
ellipsoid,
11–12 9 6–6.5 lm, thin-walled, without gelatinous sheath
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Fig. 87 Phylogram generated from a maximum likelihood analysis
based on combined ITS and LSU sequence data. The newly generated
nucleotide sequences are compared against the GenBank database
using the Mega BLAST program. Related sequences were obtained
from GenBank. Fifty-eight strains were included in the sequence
analyses, which comprised 1397 (ITS-1-607, LSU-608-1397) characters including gaps. Rhizoplaca porterii 55145 is used as the
outgroup taxon. The best scoring RAxML tree with a final likelihood
125
value of - 5739.681991 is presented. The matrix had 462 distinct
alignment patterns, with 31.86% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.234, C = 0.258,
G = 0.284, T = 0.224; substitution rates AC = 1.101172, AG =
2.447560, AT = 1.586081, CG = 0.813290, CT = 8.452097, GT =
1.000000; gamma distribution shape parameter a = 0.205506.
Bootstrap support values for ML equal or greater than 50% are given
above the nodes. The newly generated sequences are in blue
Fig. 88 Micarea squamulosa
(isotype). Habitus. Scale
bar = 0.5 mm
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but usually with a central vacuole/oil droplet. Pycnidia not
observed. Secondary chemistry: Cortex and medulla K–,
P–, C–, UV–. TLC: Atranorin, gyrophoric acid and skyrin;
all apparently in concentrations which are too low for spot
reactions.
Material examined: BRAZIL. BAHIA: Chapada Diamantina, Lençóis, Cachoeira do Mosquito; 12230 S,
41220 4000 W, alt. c. 450–500 m; 22 July 2017, on siliceous
soil, M.E.S. Cáceres & A. Aptroot ISE 40719 (ISE holotype; ABL isotype).
Notes: This species looks strongly like a small species of
Toninia, especially because of the olive-green, often contorted squamules with rounded margins and often hollow
medulla. Timdal (1991) did not accept any of the ten
species originally described from Brazil in this genus; they
were partly referred to other genera by him, and partly
excluded without alternative generic dispositions. Aptroot
(2002) referred some of these to the genera Bacidiopsora
and Micarea. The first genuine Toninia, T. massata (Tuck)
Herre, was reported from Brazil only recently by Cáceres
et al. (2017), from Ceará state. The present species differs
from all previously reported toninioid species from the
country. Sequence data of the mitochondrial small subunit
rDNA (GenBank accession: MK080110) place this taxon
in the Pilocarpaceae, in the monophyletic Micarea core
clade (Fig. 89); however, no distinctly squamulose taxa
were known in this clade before. This clade remains
unresolved and may include several genera, apart from
Micarea s.str., which is represented by the strongly supported M. prasina subclade. In that case, the squamulose
taxon may represent a separate genus. The species previously reported as Toninia from Brazil or similar taxa
elsewhere also need to be resampled, to test whether some
of these are related to the M. bullata lineage.
The species was found in the Chapada Diamantina, an
area in the state of Bahia in tropical northeastern Brazil
representing the second largest conserved area in the
country outside the Amazon (Funch et al. 2009). The natural vegetation of the Chapada Diamantina includes various types of evergreen moist and semi-deciduous dry
tropical forest, belonging to the Caatinga, Cerrado and
Atlantic rain forest biomes and the Restinga-like Capitinga
(Funch et al. 2009). The siliceous bedrock, geologically
mostly of precambrian origin, surfaces in many parts in the
form of isolated inselbergs surrounded by forest. These
inselbergs are usually only up to 500 m higher than the
surrounding landscape, but the meso- and microclimate on
the often rather flat tops and the generally very steep slopes
Fig. 89 Best-scoring maximum-likelihood tree of a subset of Pilocarpaceae based on mtSSU sequence data, including the Pilocarpaceae core (as outgroup or sister clade) and the Micarea core
clade, showing the position of the new species, M. squamulosa.
GenBank accession numbers are indicated in the tree figure. The final
alignment comprised 914 columns, with 450 distinct alignment
patterns, and the final likelihood was –6458.814273. Estimated base
frequencies were as follows: A = 0.335039, C = 0.150256,
G = 0.201024, T = 0.313681, and substitution rates as follows:
AC = 1.022403, AG = 4.972129, AT = 1.612594, CG = 0.507417,
CT = 4.968323, GT = 1.000000, with the gamma distribution shape
parameter a = 0.290696. Bootstrap values based on 1000 pseudoreplicates are placed above the branches
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Fungal Diversity (2019) 96:1–242
differs markedly from that in the surrounding areas, leading
to the presence of so-called campos rupestres, to some
extent resembling elfin forest but dominated by sclerophyllous small trees and shrubs adapted to the particular
edaphic and climatic conditions of this habitat (Conceição
et al. 2007; Funch et al. 2009). So far, around 300 lichenized species have been reported from the region (Kalb
1986; Ahti and Oksanen 1990; Kashiwadani and Kalb
1993; Mendonça et al. 2016; Aptroot and Cáceres 2018a, b;
Kalb and Aptroot 2018), many from a single field trip
during which also the following enigmatic squamulose
lichen was collected, on vertical soil at a raised ledge in the
middle of a river. The new species was found on wet sand
with bryophytes and a possibly undescribed Gyalideopsis
on a vertical ledge, in crevices between sandstone bedrock,
on a small island in a broad river near a waterfall. The
location is very wet and poor in nutrients.
Ostropomycetidae V. Reeb, Lutzoni & Cl. Roux
Notes: Ostropomycetidae is the second largest subclass
in class Lecanoromycetes, including six orders with mostly
lichenized taxa but also some secondarily delichenized
lineages (Lücking et al. 2017).
Ostropales Nannf.
Notes: Ostropales is the largest order of subclass
Ostropomycetidae, with 11 families, approximately 140
genera, and well over 3200 accepted species (Lücking et al.
2017). Among the largest families are Graphidaceae and
Porinaceae, with a new genus in Graphidaceae and a new
species in Porinaceae introduced below.
Porinaceae Rchb.
Notes: Porinaceae is a mid-sized family of approximately 360 known species, currently classified in six genera (Lücking et al. 2017). The generic concept is in flux and
based on molecular data, at least 15 genus-level lineages
can be distinguished (Sobreira et al. 2018). All Porinaceae
produce perithecioid ascomata and are rather uniform in
their internal morphology; some lineages also reproduce
vegetatively by means of isidia or isidioid structures and
rarely soredia. The latter are characteristic of the genus
Myeloconis (McCarthy and Elix 1996) but otherwise
unknown in the family. Here we describe a new species
with exposed, black perithecia and a sorediate thallus
(Fig. 90) which based on ITS sequence data belongs in the
Porina byssophila clade (Fig. 91), representing an
unnamed genus (Sobreira et al. 2018) corresponding to
Pseudosagedia subgen. Limosagedia Hafellner & Kalb
(1995).
Porina sorediata Aptroot, Lücking & M. Cáceres, sp. nov.
Index Fungorum number: IF555464; Facesoffungi
number: FoF05967; Fig. 90
127
Etymology: Referring to the sorediate thallus, an unusual
feature in the family except in the genus Myeloconis.
Holotype: M. E. S. Cáceres & A. Aptroot ISE 42451
(ISE)
Porina with capitate soralia and black semiglobose
perithecia with 3-septate, somewhat curved ascospores
with rounded ends, of 14–15 9 3.5–4 lm. Thallus covering araes of bark of up to 20 cm diam., mostly smooth and
often somewhat glossy, greyish green, not surrounded by a
differentiated prothallus; photobiont tentepohlioid, globiose to usually ellipsoid, c. 7–12 9 6–8 lm. Soralia
numerous, round, globose, somehwat paler than the thallus,
especially when abraded, c. 0.2–0.5 mm diam. Soredia
farinose,
globose
to
usually
ellipsoid,
c.
17–27 9 10–18 lm, consisting of one trentepohlioid algal
cell of c. 7–12 9 6–8 lm, surrounded by dozens of hyaline, semiglobose to somewhat angular hyphal cells with
lumina of c. 2–3 lm. Ascomata uncommon, perithecia,
black, semiglobose, c. 0.2 mm diam., c. 0.1 mm high. Wall
clearly differentiated into a proper ascoma wall and an
involucrellum which covers the exposed half of the ascoma
with a separate layer; both layers purple-brown, unchanged
in KOH. Ascospores 8/ascus, hyaline, 3-septate, somewhat
curved with rounded ends, 14–15 9 3.5–4 lm.
Material examined: Brazil, Alagoas, Quebrangulo,
Reserva Biólogica de Pedra Talhada, private area, 9150 S,
36250 3500 W, on tree bark, c. 600 m alt., 21–23 Oct 2017,
M.E.S. Cáceres & A. Aptroot ISE 42451 (ISE holotype;
ABL isotype); Same locality and datails, ISE 42328, 42450
& 42675 (ISE, ABL).
Notes: Only three species in the family Porinaceae are
known to form soredia, all in the genus Myeloconis
(McCarthy and Elix 1996). Species of the latter genus have
brighly pigmented soredia and medulla, perithecia with
pale walls, and large, muriform ascospores, thus quite
different from the new species. Species with exposed,
black perithecia occur in several lineages of Porinaceae,
including the genera Pseudosagedia and Trichothelium and
the Porina byssophila clade (Sobreira et al. 2018). Unfortunately, we were unable to generate mtSSU data for the
present taxon, and the nuLSU primers sequenced a fungal
contaminant. However, the ITS data (GenBank accession:
MK080109) place the new species in the P. byssophila
clade, together with P. guentheri (Fig. 91). This is notable,
as the P. byssophila clade, in spite of its very different
morphology, is closely related to the genus Myeloconis
based on mtSSU data (Sobreira et al. 2018), and hence the
molecular phylogeny brings the lineages with soredia
rather close together. Hafellner and Kalb (1995) established subgenus Limosagedia for a subgroup of species
within Pseudosagedia, with Porina linearis (Leight.)
Zahlbr. as type but including also P. byssophila. The stated
difference was the K ? bluish outer perithecial wall. Thus,
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Fig. 90 Porina sorediata (isotype). a Thallus with ascomata and soralia. b Thallus with soralia. c Section through ascoma. d Soredia. Scales
bars: a, b = 1 mm, c = 0.2 mm, d = 50 lm
Fig. 91 Best-scoring maximum-likelihood tree of available subset of
Porinaceae based on ITS sequence data, showing placement of P.
sorediata in the P. byssophila clade which, based on mtSSU sequence
data, is closely related to the genus Myeloconis (Sobreira et al. 2018).
The final alignment comprised 534 columns, with 210 distinct
alignment patterns, and the final likelihood was –1903.538679.
Estimated base frequencies were as follows: A = 0.187911,
C = 0.338406, G = 0.295727, T = 0.177956, and substitution rates
as follows: AC = 1.312730, AG = 3.551014, AT = 2.797834, CG =
1.299677, CT = 6.511396, GT = 1.000000, with the gamma distribution shape parameter a = 0.330010. Bootstrap values based on
1000 pseudoreplicates are placed above the branches
Limosagedia could potentially be a name available for this
clade when raised to genus level; however, in the present
species no such K-reaction was observed. The new taxon
was found in the Pedra Talhada Biological Reserve in the
state of Alagoas, northeastern Brazil, where it was locally
quite common on shaded bark of various trees, but it has
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Fungal Diversity (2019) 96:1–242
not been observed in any other site in this locality, a
phenomenon also known from other species that appear to
represent local endemics. Pedra Talhada Biological
Reserve is ecologically quite unique, representing one of
the larger remnants of Atlantic rain forest in northeastern
Brazil, with various phytophysiognomies formed on
inselbergs built from precambrian gneiss (Nusbaumer et al.
2015; Studer et al. 2015). The new species is thus far only
known only from Brazil.
Graphidaceae Dumort.
Notes: Graphidaceae is the second largest family of
lichenized Ascomycota, containing containing over 2000
known species and about 80 genera (Lücking et al. 2017).
Although most lineages have been sequenced, odd taxa that
do not fit any of the genera now recognized continue to be
encountered on a regular basis (Lumbsch et al. 2014). Here
we describe a new genus based on sequenced material
found in the same area as Porina sorediata, described
elsewhere in this paper. The species was identified as the
rare Thelotrema cryptotrema Nyl., later recombined as
Ocellularia cryptotrema (Nyl.) Kalb and more recently as
Schizotrema cryptotrema (Nyl.) Rivas Plata & Mangold
(Rivas Plata et al. 2010). Sequence data show that the taxon
is unrelated to Thelotrema or Schizotrema and instead
forms a novel lineage in tribe Ocellulariae, related to the
Ocellularia profunda and O. praestans clades (Fig. 93).
The latter two are not members of Ocellularia sensu stricto
and will require new genera placement as well, but here we
formally describe a new genus for Thelotrema cryptotrema,
which in addition to its phylogenetic placement also
exhibits some unusual features within tribe Ocellularieae
(Fig. 92).
Cryptoschizotrema Aptroot, Lücking & M. Cáceres, gen.
nov.
Index Fungorum number: IF555465; Facesoffungi
number: FoF05968
Etymology: Referring to the anatomical resemblence
with Schizotrema species, due to the partially carbonized
and concentrically layered, fissured excipulum, and the
notion that the Schizotrema-like excipulum is hidden
beneath a covering layer.
Type species: Cryptoschizotrema cryptotrema (Nyl.)
Aptroot, Lücking & M. Cáceres.
Cryptoschizotrema cryptotrema (Nyl.) Aptroot, Lücking &
M. Cáceres, comb. nov.
Index Fungorum number: IF555466; Facesoffungi
number: FoF05969; Fig. 92
= Thelotrema cryptotrema Nyl., Ann Sci Nat Bot Sér 5,
7: 318 (1867).
= T. secoligella Müll. Arg., Hedwigia 34: 31 (1895).
129
= T. annulatum Müll. Arg., J Linn Soc Bot 30: 453
(1895).
New genus in Graphidaceae with myriotremoid to
porinoid, immersed-erumpent ascomata with a narrow
pore, dark brown to partially (apically) carbonized, fissured
and becoming concentrically layered excipulum lacking
periphysoids, muriform, subdistoseptate, I-negative (nonamyloid) ascospores with thin walls and septa, and psoromic acid chemistry. Thallus light greenish grey, unevenverrucose, with dense, prosoplectenchymatous cortex. Ascomata immersed-erumpent, rounded, disc covered by
narrow pore, invisible; proper margin covered by whitish
thalline layer, not visible, when abraded seen as brown to
black, fissured lobules forming concentric layers in older
ascomata. Columella absent. Excipulum prosoplectenchymatous, brown to partially (apically) carbonized, becoming
layered (striate) in older ascomata; periphysoids absent.
Hymenium clear; paraphyses unbranched. Ascospores
1–2(–4)/ascus, richly muriform, up to 100 9 30 lm,
oblong-ellipsoid, distoseptae with rather thin walls and
septa and angular lumina, colorless, I– (non-amyloid).
Secondary chemistry: psoromic acid (thallus P ? yellow).
Material examined: Brazil, Alagoas, Quebrangulo,
Reserva Biólogica de Pedra Talhada, private area, 9150 S,
36250 3500 W, on wooden pole, c. 550 m alt., 21–23 Oct
2017, M.E.S. Cáceres & A. Aptroot ISE 42655 (ABL, B,
ISE).
Notes: Tribe Ocellularieae largely comprises the taxa
previously classified in the collective genera Myriotrema
and Thelotrema (Hale 1980; Lumbsch et al. 2014). Both
are highly polyphyletic, as shown by Frisch et al. (2006)
and Rivas Plata et al. (2012), and even after formally
segregating several lineages as new genera, quite a number
of orphaned lineages remain unnamed. Our results show
that Thelotrema cryptotrema (nuLSU Genbank accession:
MK080108) represents an additional lineage in this
heterogeneous assembly, related to the O. profunda and O.
praestans clades. This topology would theoretically allow
to include the latter two clades together with T. cryptotrema in a single genus, for which the here established
name would then be available. However, at this time we
refrain from placing additional species in Cryptoschizotrema, since the long branches leading to each
clade (Fig. 93) do not suggest a single genus to be involved
and the differences between Cryptoschizotrema and the
other two clades are substantial, including the massive,
fully carbonized and non-striate excipulum and the distoseptate, I ? violet-blue (amyloid) ascospores in O. profunda and O. praestans and relatives.
This monospecific new genus is thus far only known
from Central and South America. It is tropical lowland to
montane species growing on bark or wood; the sequenced
specimen was found in Brazilian Atlantic Rain Forest, on a
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Fungal Diversity (2019) 96:1–242
Fig. 92 Cryptoschizotrema cryptotrema. a–d Cáceres & A. Aptroot
ISE 42655. e, f isotype of T.annulatum in BM). a Thallus with
ascomata. b Ascomata enlarged. c Hymenium with asci and
ascospores. d Ascospores. e Abraded ascoma showing schizotremoid
proper margin (photograph A. Mangold). f Section through ascoma
showing striate, apically carbonized excipulum (photograph A.
Mangold). Scale bars: a = 1 mm, b, e = 0.5 mm, c, d = 10 lm,
f = 100 lm
wooden pole of a fence dividing different parts of the
reserve.
Wijayawardene et al. (2018a) accepted 11 orders and 44
families in Leotiomycetes.
Leotiomycetes O.E. Erikss. & Winka
Notes: Leotiomycetes is the largest the largest class of
inoperculate discomycetes, a group of nonlichenized
ascomycetes (Eriksson and Winka 1997; Lumbsch et al.
2005; Zhang and Wang 2015; Ekanayaka et al. 2017).
Helotiales Nannf.
Notes: Helotiales species are abundant in humid areas as
saprobes on dead leaves and shoot on herbaceous and
woody plants (Webster 2007). In addition, they occur as
plant pathogens and endophytes. Helotiales is a highly
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131
Fig. 93 Best-scoring maximum-likelihood tree of subset of Graphidaceae tribe Ocellularieae based on three markers (mtSSU, nuLSU,
RPB2), showing placement of Cryptoschizotrema cryptotrema near
the O. profunda and O. praestans clades. The final alignment
comprised 2618 columns, with 983 distinct alignment patterns, and
the final likelihood was –14732.747014. Estimated base frequencies
were as follows: A = 0.289260, C = 0.202949, G = 0.260731,
T = 0.247061, and substitution rates as follows: AC = 0.929088,
AG = 3.966253, AT = 2.046683, CG = 0.790669, CT = 7.857784,
GT = 1.000000, with the gamma distribution shape parameter
a = 0.231216. Bootstrap values based on 1000 pseudoreplicates are
placed above the branches
diversified order and accommodates 27 families (Ekanayaka et al. 2017; Wijayawardene et al. 2018a).
Saprobic on dead flower petals and leaves of Rosa sp.
Sexual morph Undetermined. Asexual morph Conidiomata
sporodochial, 175–200 lm high (
x = 185 lm, n = 10), 122–
140 lm diam. (
x = 133.5 lm, n = 10), 48–56.5 lm
(
x = 52.5 lm, n = 5) wide at the base, superficial, scattered,
sessile, relatively sphaerical, initially yellowish brown,
turning brownish or almost black, initially convex cap turning
to concave. Peridium thick, textura angularis. Conidiophores
11.2–19.8 9 1.0–1.4 lm (
x = 15.6 9 1.2 lm, n = 10), a
dense palisade, hyaline to pale brown, tapering distally.
Conidiogenous cells 2.2–3.3 9 0.8–1.1 lm (
x = 2.6 9 0.9
lm, n = 10), enteroblastic, phialidic, hyaline, tapering
towards the apex, terminal and lateral. Conidia 4.8–
7.4 9 1.5–2.4 lm (
x = 6.1 9 1.9 lm, n = 40), L/W 3.2,
1-celled, hyaline, fusoid to allantoid, slightly falcate, apex
acute, base obtuse, thin-walled, guttules at both ends, formed
singly on the conidiogenous cells.
Culture characteristics: Colonies reaching to 36.5 mm
after 2 weeks at 28 C on PDA with two zones. Inner zone
dirty yellow to light brown, flat, slightly clock wise rotate
sparse mycelia, reverse light brown; outer layer dirty white,
flat, smooth, lobate margin, reverse dirty white. Later,
zonate, dirty white, flat surface, smooth with light brown
patches, reverse dark brown centre and light brown margin.
Chaetomellaceae Baral, P.R. Johnst. & Rossman
Notes: Chaetomellaceae was introduced by Baral et al.
(2015) with its type genus Chaetomella Fuckel to accommodate non-lichanized discomycetes as a family insertae
sedis in Leotiomycetes. ITS and LSU based phylogenetic
study revealed that the family consists of four genera viz.
Chaetomella (= Zoellneria), Pilidium Kunze (= Discohainesia), Sphaerographium Sacc. and Synchaetomella
Decock & Seifert. Wijayawardene et al. (2018a) accepted
the family in the Helotiales (Leotiomycetes).
Pilidium Kunze
Notes: The genus Pilidium was introduced with P.
acerinum (Alb. & Schwein.) Kunze as the type species
with a Hainesia synasexual, and Discohainesia sexual
morphs (Rossman et al. 2004; Johnston et al. 2014). Currently, 22 accepted Pilidium species have been reported as
saprobes and pathogens on various hosts (Wijayawardene
et al. 2017a).
Pilidium concavum (Desm.) Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 124: 148 (1915)
Facesoffungi number: FoF05212; Fig. 94
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Fig. 94 Pilidium concavum (MFLU 18-0106, new geographical
record). a Host. b, c Appearance of conidiomata on host surface.
d Vertical section through the conidioma. e–h Conidiophores and
conidiogenous cells. i, j Culture on PDA (i from above view, j from
below view). k Conidiospores. Scale bars: b–d = 100 lm, e–h,
k = 5 lm
Material examined: THAILAND, Chiang Rai Province,
Muang District, Tha Sut Sub-district, (20 030 24.700 N, 99
520 23.500 E), dead petals and leaves of Rosa sp. (Rosaceae),
20 August 2017, MC. Samarakoon, SAMC017 (MFLU
18-0106, HKAS 102341), living culture (MFLUCC
17-2671).
GenBank numbers: ITS: MK100327, LSU: MK108192,
SSU: MK108189.
Notes: Pilidium concavum (synanamorph Hainesia
lythri (Desm.) Höhn.; sexual morph Discohainesia oenotherae (Cooke and Ellis) Nannf.), is an interesting species
with morphological differentiation due to genetic control
(Sutton 1980; Palm 1991; Rossman et al. 2004). The species is often pathogenic and occasionally saprobic on
flowers, fruits and oil and forest plants (Cardin et al. 2009;
Geng et al. 2012; Lopes et al. 2010; Ayoubi et al. 2016).
Pilidium concavum has been recorded mainly in temperate
regions and on many hosts. Palm (1991) reports P. concavum on Rosa leaves from the United Kingdom and
Connecticut (USA). The strain isolated from petals and
leaves of Rosa sp. in this study is similar in morphology
and phylogeny to P. concavum. The ITS-LSU multi-locus
phylogenetic studies reveal that strain MFLUCC 17-2671
is closely related (86% ML/0.99 BYPP) to P. concavum
and the P. lythri clade (Fig. 95). Morphological characterizations, especially the conidia, overlap with the consistency measurements (5.5–7.2 9 1.4–1.8 lm) described
by Palm (1991). Thus, we report this as P. concavum on
petals and leaves of Rosa sp. as a saprobe from Thailand, a
new geographical record.
123
Pezizomycetes O.E. Erikss. & Winka
Notes: Pezizomycetes is the representative operculate
discomycete of Ascomycota. It is characterized by
apothecial ascomata or deformed apothecia and asci with
an apical slit or operculum (Hansen and Pfister 2006).
Presently, 21 families are accepted in this class (Ekanayaka
et al. 2018). Phylogenetic studies based on multiple genes
Fungal Diversity (2019) 96:1–242
133
Fig. 95 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data for Pilidium species.
Related sequences are taken from Marin-Felix et al. (2017). Thirteen
strains are included in the combined analyses which comprise 1250
characters (465 characters from ITS, 785 characters for LSU) after
alignment. Chaetomella raphigera (BPI 843551) (Chaetomellaceae,
Helotiales) is used as the outgroup taxon. Single gene analyses are
carried out and the topology of each tree had clade stability. Tree
topology of the maximum likelihood analysis is similar to the
Bayesian analysis. The best RaxML tree with a final likelihood value
of - 3008.149666 is presented. The matrix had 128 distinct
alignment patterns, with 1.3% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.258502,
C = 0.211927, G = 0.278401, T = 0.251171; substitution rates AC =
3.685184, AG = 2.311128, AT = 2.325507, CG = 0.504055, CT =
5.692118, GT = 1.000000; gamma distribution shape parameter
a = 0.02. Bootstrap values for maximum likelihood (ML) equal to or
greater than 70% and Bayesian posterior probabilities (BYPP) equal
or greater than 0.95 are placed respectively on the branch. Type
strains are bold. The newly generated sequence is indicated in blue
indicate the Pezizomycetes is a monophyletic group
(Ekanayaka et al. 2018).
(Korf & W.Y. Zhuang) Y.J. Yao & Spooner, B. luteola
Svrček, B. nicholsonii (Massee) Spooner & Y.J. Yao, B.
ovalispora (Boud.) Van Vooren, B. subprolata (Korf &
W.Y. Zhuang) Y.J. Yao & Spooner and B. vermiphila
Brumm. & R. Kristiansen are known. Three species have
available molecular data (Perry et al. 2007; Hansen et al.
2013; Kušan et al. 2018).
Pezizales J. Schröt.
Notes: Phylogenetic analyses by Ekanayaka et al. (2018)
show six clades in this order. All operculate taxa and some
non-apothecial taxa are included in Pezizales. In addition
to genera from 21 families with a resolved placement, 15
genera are unresolved and listed as Pezizales incertae sedis
(Jaklitsch et al. 2016; Wijayawardene et al. 2017a).
Ascodesmidaceae J. Schröt. [as ‘Ascodesmidacei’]
Notes: The family Ascodesmidaceae was introduced by
Schröter (1893) and is typified by Ascodesmis Tiegh. with
Ascodesmis aurea Tiegh. as type species. This family is
characterized by small ascomata, straight or curved paraphyses and smooth or ornamented ascospores (Güngör
et al. 2014; Ekanayaka et al. 2018). Currently, twelve
genera are listed in this family, but just four have sequence
data. (Ekanayaka et al. 2018; Wijayawardene et al. 2018a).
Phylogenetically, this family is close to Pyronemataceae
Corda and Otideaceae Eckblad within Pezizales (Ekanayaka et al. 2018).
Boubovia Svrček
Notes: The genus Boubovia was established by Svrček
(1791) for Boubovia luteola Svrček which is characterized
by glabrous to setaceous ascomata, inamyloid asci and
smooth or ornamented ascospores with a cyanophilic
sheath (Korf and Zhuang 1984; Yao and Spooner 1996;
Kristiansen 2016). Six species, Boubovia ascoboloides
Boubovia gelatinosa M. Zeng, Q. Zhao & K.D. Hyde, sp.
nov.
Index Fungorum number: IF555488; Facesoffungi
number: FoF05174; Fig. 96
Etymology: The epithet refers to ascospores surrounded
by gelatinous sheath.
Holotype: HKAS 102407
Saprobic on sheep dung. Sexual morph Ascomata
superficial, scattered. Disc pulvinate, yellowish to orange,
shiny, up to 1 mm high, 0.5 mm broad, with 1–2 mm long
setaceous hairs, exceeding the disc. Medullary excipulum
of textura intricata. Ectal excipulum, hyaline, J-, comprised of 8–12 9 6–8 lm cells of textura angularis to
globulosa. Paraphyses 2–3 lm broad, filiform, septate,
straight, hyaline, J-. Asci 108–160 9 18–23 lm, 8-spored,
subcylindrical to clavate, operculate, inamyloid. Ascospores
[20/1/1,
in
H2O]
(16.3–)16.6–18.9(–
21.1) 9 (7.7–)8.9–10.4(–11.0)
(Q = 1.65–2.30,
Q = 1.84 ± 0.15), ellipsoid to subglobose, uniseriate to
biseriate or irregular, smooth-walled, enveloped by gelatinous sheath or over 2/3 part surrounded by sheath, deBarybubbles present. Asexual morph Undetermined.
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Fungal Diversity (2019) 96:1–242
Fig. 96 Boubovia gelatinosa (HKAS 102407, holotype). a–c Typical
mature specimens. d Squash of ascomata in Melzer’s reagent.
e Receptacle surface of pileus. f Asci and paraphyses in Melzer’s
reagent. g–i Asci (h, i Asci in Melzer’s reagent). j–m Ascospores.
Scale bars: d = 100 lm, f–i = 30 lm, e, j = 20 lm, k–m = 15 lm
Material examined: CHINA, Jiuzhaigou, Sichuan, on
sheep dung, 22 August 2018; Ming Zeng, ZM1d (HKAS
102407, holotype).
GenBank numbers: ITS: MK093850, LSU: MK093851,
SSU: MK093852, TEF1-a: MK093853.
Notes: This species is distinguished by orange shiny
ascomata with long hairs, straight paraphyses and ellipsoid,
smooth wall ascospores with sheath. The phylogenetic tree
shows a close relationship with Coprotus ochraceus (P.
Crouan & H. Crouan) J. Moravec (Fig. 97), but morphologically, this species is more similar to Boubovia species.
Boubovia may yet prove to be synonym of Coprotus or
more than two genera may exist here, but further collections are needed. Boubovia vermiphila differs in its
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Fungal Diversity (2019) 96:1–242
135
Fig. 97 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU, SSU and TEF1-a sequence data for
Boubovia species and several closely related genera in Ascodesmidaceae. Related sequences are taken from Perry et al. (2007), Hansen
et al. (2013), Lindemann et al. (2015) and Kušan et al. (2018). Sixteen
strains are included in the combined analyses which comprise 3584
characters (669 characters for ITS, 890 characters for LSU, 1043
characters for SSU, 982 characters for TEF1-a) after alignment.
Pseudoboubovia benkertii (V.K. 3202) is used as the outgroup taxon.
Single gene analyses are carried out and the topology of each tree had
clade stability. Tree topology of the maximum likelihood analysis is
similar to the maximum parsimony analysis and the Bayesian
analysis. The best RaxML tree with a final likelihood value of
- 12041.925019 is presented. The matrix had 891 distinct alignment
patterns, with 51.43% undetermined characters or gaps. Estimated
base frequencies were as follows: A = 0.246462, C = 0.237728,
G = 0.271829, T = 0.243980; substitution rates AC = 0.807419,
AG = 1.258958, AT = 1.238671, CG = 0.599522, CT = 3.934755,
GT = 1.000000; gamma distribution shape parameter a = 0.201448.
Maximum parsimony analysis of 2717 constant characters and 600
informative characters resulted in two equally most parsimonious tree
of 1614 steps (CI = 0.778, RI = 0.704, RC = 0.547, HI = 0.222).
Bootstrap values for maximum likelihood (ML) and maximum
parsimony (MP) equal to or greater than 75 are placed above and
below the branches respectively. Branches with Bayesian posterior
probabilities (BYPP) equal or greater than 0.90. The newly generated
sequence is indicated in blue
glabrous ascomata, curved paraphyses, ornamented ascospores with 1–2 oil drops and large apical ascospore caps
(Brummelen and Kristiansen 1999; Kristiansen 2016).
Boubovia ascoboloides and B. ovalispora differ in their
glabrous ascomata and curved paraphyses (Korf and
Zhuang 1984; Kristiansen 2016). Boubovia subprolata
123
136
differs in its pinkish glabrous ascomata and curved paraphyses (Korf and Zhuang 1991).
Sordariomycetes O.E. Erikss. & Winka
Diaporthomycetidae Senan., Maharachch. & K.D.
Hyde
Notes: The subclass Diaporthomycetidae was introduced
to accommodate ten orders and 26 families (Maharachchikumbura et al. 2015). Diaporthales, initially
placed in the subclass Sordariomycetidae, where it formed
a sister group with Magnaporthaceae, has been transferred
and established as type order of the subclass Diaporthomycetidae (Zhang et al. 2006). Species of this subclass are saprobic, endophytic, parasitic or pathogenic in
plants, animals or humans (Maharachchikumbura et al.
2015). The sexual morphs comprise stromata made up of
orange, brown or black parenchymatous tissues and seldom
pyriform ascomata. The ascospores are 2–3 seriate, aseptate or septate, of different shapes, and may or may not
have appendages or sheaths. The asexual morphs are ceolomycetes, where conidiomata are acervuli or pycnidial
with annellidic or phialidic proliferating conidiogenous
cells, or as hyphomycetes, where conidiophores originate
from hyphae with occasionally coloured terminal or lateral
conidiogenous cells (Maharachchikumbura et al. 2015).
Diaporthales Nannf.
Notes: Diaporthales is a well-supported monophyletic
clade within the Diaporthomycetidae and members of this
order are saprobes and soil inhabitants, pathogens, parasites
or endophytes in plants (Zhang et al. 2006; Rossman et al.
2007; Maharachchikumbura et al. 2015). Some are also
pathogenic to humans and animals (Rossman et al. 2007).
Diaporthalean microfungi are currently accommodated
within 21 families. Among all the species in this order,
Cryphonectria parasitica (Murrill) M.E. Barr is probably
the most well-known chestnut blight fungus which
destroyed the whole population of American chestnut trees
[Castanea dentata (Marsh.) Borkh.] within a short period
of time (Anagnostakis 1987). Taxa belonging to this order
have yellowish brown to black valsoid or diatrypoid
pseudo- or asco-stromata, brown to black perithecial
fruiting bodies, regularly immersed in stromata of leafy or
woody substrates with papillate ostioles swollen at the
apex. The mature 2–32 spored asci with a J- refractive
apical apparatus, float freely and become dehiscent at
maturity (Barr 1978; Samuels and Blackwell 2001; Senanayake et al. 2017). The asexual states are usually
ceolomycetous, with conidiophores arising from the topmost cell layer of basal or parietal tissue or from under the
developing scutellum; otherwise, they are reduced to
enteroblastic, holoblastic, phialidic or annellidic hyaline or
olivaceous conidiogenous cells (Sogonov et al. 2008;
Senanayake et al. 2017).
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Diaporthaceae Höhn.
Notes: The family Diaporthaceae currently comprises 14
genera (Maharachchikumbura et al. 2016; Senanayake
et al. 2017, 2018; Dissanayake et al. 2017b).
Diaporthe Nitschke
Notes: Diaporthe includes economically important plant
pathogens, endophytes, and saprobes (Udayanga et al.
2014; Hyde et al. 2016). They have a broad range of hosts
from cultivated crops to ornamental plants (Dissanayake
et al. 2017b). Rossman et al. (2015a) prioritized the older
name Diaporthe over Phomopsis, while the latest account
is provided in Fig. 100. An updated phylogenetic tree for
Diaporthe species with new records and new species is
presented in Fig. 100.
Diaporthe italiana Chethana, Camporesi & K. D. Hyde,
sp. nov.
Index Fungorum number: IF555376; Facesoffungi
number: FoF04934; Fig. 98
Etymology: The specific epithet ‘italiana’ was given
after the country, where the fungus was collected.
Holotype: MFLU 17-0311
Saprobic on dead aerial branch of Morus alba L. Sexual
morph Perithecia 0.2–0.42 mm (
x = 0.311 mm, n = 10),
solitary, scattered, immersed to semi-immersed, partially
erumpent, globose, black, with tapering perithecial necks.
Peridium multi-layered, 20 lm wide at the top, 10–12 lm
wide in sides, outer layer composed of 4–5 layers of thick,
brown cells, inner 3–4 layers of hyaline cells of textura
angularis, cells towards inner layer lighter. Asci 45–
55 9 6–8.5 lm (
x = 51 9 7.7 lm, n = 20), 8-spored,
unitunicate, sessile, cymbiform to clavate. Ascospores
10.5–13 9 2.7–4.5 lm (
x = 12.1 9 3.5 lm, n = 40),
overlapping biseriate, hyaline, fusiform to allantoid,
slightly curved, 2-celled, widest near the centre, with acute
rounded ends, 1-median septate, rarely bi-guttulate, often
tetra-guttulate, with larger guttules at centre and the
smaller towards the ends, smooth-walled. Asexual morph
not observed.
Culture characteristics: Colonies on the PDA, circular,
fluffy, aerial, white mycelium with fimbriate margin, olivaceous to grey olivaceous reverse, relatively slow growing, reach 3 cm diam. after 7 days at 25 C.
Material examined: ITALY, Forlı̀-Cesena Province,
Forlı̀, via Friuli, on dead, aerial branch of Morus alba L.
(Moraceae), 19 January 2017, Erio Camporesi, IT 3217
(MFLU 17-0311, holotype), ex-type living culture
(MFLUCC 18-0090, KUMCC 18-0002); (HKAS 101460,
isotype), ex-isotype living culture (MFLUCC 18-0091,
KUMCC 18-0003).
GenBank numbers: CAL: MH853690, ITS: MH846237,
TEF1-a: MH853686, TUB2: MH853688 (MFLUCC
Fungal Diversity (2019) 96:1–242
137
Fig. 98 Diaporthe italiana (MFLUCC 18-0090, holotype). a Appearance of perithecia on dead branch of Morus alba. b Longitudinal
section of a perithecium. c Longitudinal section of a perithecium wall
showing cell organization. d Bi-seriate asci. e–j Ascospores. k, l
Colony on PDA (k from above view, l from below view). Scale bars:
a = 1 mm, b = 100 lm, c = 20 lm, d = 10 lm, e–j = 5 lm
18–0090); CAL: MH853691, ITS: MH846238, TEF1-a:
MH853687, TUB2: MH853689 (MFLUCC 18-0091).
Notes: Diaporthe italiana is phylogenetically closely
related to D. rudis, but is clearly distinguished based on the
morphology and phylogeny. In our phylogenetic analysis
of combined ITS, TEF1-a, TUB2, and CAL sequence data
of Diaporthe species (Fig. 100), our strains (18-0090 and
18-0091) cluster together with the ex-type strain of Diaporthe rudis (AR3422) with relatively high bootstrap and
Bayesian probabilities (100% ML/1.00 BYPP). Sequence
comparison for ITS region between D. rudis (AR3422) and
D. italiana (MFLUCC 18–0091) showed 2.11% base pair
differences. Our collection is distinct from D. rudis in
having smaller asci (45–55 9 6–8.5 lm), and smaller
ascospores (10.5–13 9 2.7–4.5 lm), in contrast to larger
asci (53.5–58.5 9 10.6–12 lm), and larger ascospores
(12–14.2 9 3.5–3.7 lm) of D. rudis (Udayanga et al.
2014). A Diaporthe species named Diaporthe mori Berl.
(current name Phomopsis moricola (Sacc.) Grove) has
been identified from Morus alba and M. nigra (BargagliPetrucci 1915). However, they differ from our species in
having smaller, immersed perithecia (0.18–0.2 mm),
smaller asci (36–40 9 7 lm), and smaller, bi-guttulate
ascospores (7–8 9 3–3.5 lm). Since, this species were
introduced in 1915, DNA sequences were unavailable in
123
138
the databases. Therefore, they were not included in the
phylogenetic analysis.
Diaporthe rumicicola Manawasinghe, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555379; Facesoffungi
number: FoF04940; Fig. 99
Etymology: Name reflects the host genus.
Holotype: MFLU 18-0739
Saprobic on dead leaves of woody plants. Sexual
morph Undetermined. Asexual morph Conidiomata on
host pycnidial, 98–280 lm diam. (
x = 208 lm, n = 10),
ampulliform, scattered, immersed, ostiolate, with elongate
Fig. 99 Diaporthe rumicicola (MFLU 18-0739, holotype). a Appearance of conidiomata on dead branch of Rumex spp. b Cross section of
conidioma on host. c Pycnidial wall on host. d, e Conidiogenous cells
with developing conidia. f Conidia on host. g Pycnidia on PDA.
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black neck. Pycnidial wall comprising 2–3 layers, heavily
pigmented, thick-walled, comprising dark brown cells of
textura angularis, with lighter cells towards the inside,
with inner layer composed of 2–5 layers, hyaline, thinwalled cells of textura angularis. Conidiophores on host
reduced to conidiogenous cells. Pycnidia in culture black,
immersed, solitary or merged, exudated. Paraphyses
observed on culture. Conidiogenous cells enteroblastic
with percurrent annellations, integrated, clustered, hyaline,
smooth-walled. Alpha conidia, on host 3–5 9 2–3 lm
(
x = 3.5 9 2.5 lm, n = 30), hyaline, smooth-walled,
mono or bi-guttulate. Beta conidia not observed.
h Appearance of paraphysis. i, j Colony on PDA (i from above view,
j from below view). Scale bars: a = 2000 lm, b = 200 lm,
g = 100 lm, c–e, h = 20 lm, f = 10 lm
Fungal Diversity (2019) 96:1–242
Culture characteristics: Colonies on PDA 50 mm diam.,
after 7 days at 25 C, cream to white cottony mycelium,
irregular and lobate at the margins, white at the centre;
reverse yellowish and concentric lines on conidial formation can be observed when colony getting old.
Material examined: ITALY, near Castrocaro Terme
(province of Forlı́-Cesena), on dead aerial stem of Rumex
sp. (Polygonaceae), 19 February 2018, Camporesi Erio IT
(MFLU 18-0739, holotype), ex-type living culture
(MFLUCC 18-1566, JZB320005).
GenBank numbers: ITS: MH84623, TEF1-a:
MK049554, TUB2: MK049555.
Notes: Diaporthe rumicicola was isolated from dead
branch of Rumex sp. from Forı́-Cesena of Italy. Morphologically, this species is similar to genus Diaporthe, and the
combined gene sequence analysis of ITS, TEF1-a, CAL
and TUB2 genes placed the taxon in Diaporthe (Fig. 100).
The current species develop a sister clade with Diaporthe
foeniculina with 100% bootstrap value in ML analysis and
1.0 Bayesian probabilities in Bayesian analysis (Fig. 99).
Morphologically the current species is differing with
developing smaller alpha conidia (3.5 9 2.4 lm) compared to D. foeniculina (8.5–9 9 2.3–2.5 lm) (Udayanga
et al. 2014). In addition to that this is the first time that
Diaporthe species recorded on Rumex sp. (Farr and Rossman 2019).
Gnomoniaceae G. Winter
Notes: The family Gnomoniaceae, typified by the genus
Gnomonia Ces. & De Not., encompasses fungi with
immersed black perithecia which are solitary or aggregated
with or without a stromatic tissue (Sogonov et al. 2008).
The neck of the perithecial bodies extends beyond the host
epidermis and may be short or long with an upright, oblique or parallel positioning to the host surface (Sogonov
et al. 2008). The asexual morphs of species of Gnomoniaceae produce acervular or pycnidial fruiting bodies
(Monod 1983). Species belonging to this family have
mainly been reported from north America and Europe, with
only few originating from Asia (Walker et al. 2014). Most
of the species are host-specific (Senanayake et al. 2017)
and commonly occur as endophytes, pathogens or saprobes, even though, they sometimes behave as parasites or
opportunistic pathogens (Sieber 2007; Senanayake et al.
2018).
Plagiostoma Fuckel
Notes: Plagiostoma (Gnomoniaceae, Diaporthales)
encompasses microfungal members of pathogenic, endophytic or saprobic nature inhabiting stems, branches, twigs
and leaves of woody and herbaceous plants belonging to
various families which occur mostly temperate regions of
the northern hemisphere as well as some parts of Asia
(Mejı́a et al. 2011; Walker et al. 2014). The genus, typified
139
by P. euphorbiae (Fuckel) Fuckel, currently accommodates
32 species (Mejı́a et al. 2011; Walker et al. 2014; Senanayake et al. 2017). These species produce perithecial and
pycnidial fruiting bodies on overwintered leaves and twigs
(Walker et al. 2014) and the genus primarily distinguishes
itself from other genera in the family by neck characters of
the perithecia along with ascospore morphology (Mejı́a
et al. 2011). Other than these, presence or absence of
stroma, geographical location or host are secondary differentiating features (Mejı́a et al. 2011). In this study, we
report Plagiostoma salicellum (Fr.) Sogonov for the first
time from a new geographical area in Europe, notably Italy.
Plagiostoma salicellum (Fr.) Sogonov, Stud. Mycol. 62: 73
(2008)
Facesoffungi number: FoF05761; Fig. 101
Saprobic on dead branches of Salix caprea. Sexual
morph
Ascomata
205–350 9 260–330 lm
(
x = 275 9 290.5 lm, n = 6), perithecial, mostly solitary,
scattered, immersed in bark, producing a faint elevation of
the periderm, each comprising of a convergent, protruding
neck, black, subglobose. Neck 110–120 lm long
(
x = 113 lm, n = 6), 76–126 lm diam. at the base
(
x = 97.3 lm, n = 6) and 88–131 lm diam. at apex
(
x = 103.8 lm, n = 6), cylindrical, eccentric to lateral,
straight to very slightly twisted, not surrounded by a
stroma. Peridium 18–29 lm wide, thick, comprising 4–6
layers, outer layer heavily pigmented, thick walled, consisting of dark brown to blackish cells of textura angularis,
cells towards inside lighter, inner layer composed of 1–2
layers, pale brown to hyaline, flattened, thin-walled cells of
textura
angularis.
Asci
32–55 9 5–15 lm
(
x = 43.7 9 8.7 lm, n = 34), 8-spored, cylindrical to clavate, clavate-elongated, straight to slightly curved, broadly
rounded apex with a conspicuous J- refractive apical ring,
1.7–4.3 lm diam., often consisting of a short but not
necessarily persistent, tapered and pointed stipe. Ascospores 12–18 9 1–4 lm (
x = 15.8 9 2.5 lm, n = 54),
obliquely parallel or irregularly seriate, hyaline, ellipsoidelongated, tapering slightly towards rounded ends, 1-septate, slightly constricted at median to sub-median septum,
with granular cytoplasm, without any sheath or appendage.
Asexual morph Undetermined.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Monte Fumaiolo, on dead aerial branch of Salix
caprea L. (Salicaceae), 27 August 2017, Erio Camporesi,
IT 3454 (MFLU 17-1601).
GenBank numbers: ITS: MK080111, TEF1-a:
MK424970, TUB2: MK424971.
Notes: Plagiostoma salicellum is characterised by a
cylindrical neck surrounded by a white stroma and narrow
ellipsoid-elongated ascospores often having short appendages (Mejı́a et al. 2011). However, in our specimen no
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140
Fig. 100 Phylogenetic tree generated by maximum likelihood analysis of combined ITS, TEF1-a, TUB2 and CAL sequence data of
Diaporthe species. Sequences were obtained from GenBank. Onehundred and ninety-two strains are included in the analyses, which
comprise 2603 characters including gaps. Single gene analyses are
carried out to compare the topology of the tree and clade stability.
Tree was rooted with Diaporthella corylina (CBS 121124). Tree
topology of the Bayesian analysis was similar to the RAxML. The
best scoring RAxML tree with a final likelihood value of =
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Fungal Diversity (2019) 96:1–242
- 60797.442221 is presented. The matrix had 2107 distinct alignment patterns, with 40.98%of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.228179,
C = 0.282550, G = 0.268109, T = 0.221162; substitution rates AC =
0.854574, AG = 3.369047, AT = 0.975280, CG = 0.800183, CT =
3.251377, GT = 1.000000; gamma distribution shape parameter
a = 0.881490. RAxML and Bayesian posterior probabilities values C 60% (ML) and 0.9 (BYPP) are shown respectively near the
nodes. The newly generated sequences are indicated in blue
Fungal Diversity (2019) 96:1–242
141
Fig. 100 continued
stroma was observed around the perithecial neck which
was also shorter x = 113 lm (this study) vs x = 177 lm
(Mejı́a et al. 2011)] and broader both at the base
x = 97.3 lm (this study) vs x = 81 lm (Mejı́a et al. 2011)]
and at the apex x = 103.8 lm (this study) vs x = 79 lm
(Mejı́a et al. 2011)] as compared to the lectotype and
epitype of P. salicellum described by Mejı́a et al. (2011).
The perithecia were mostly solitary rather than aggregated
and narrower in diameter x = 290.5 lm (this study) vs
x = 397 lm (Mejı́a et al. 2011)]. The asci also were shorter
and narrower (
x = 43.7 9 8.7 lm (this study) vs
x = 55.0 9 13.0 lm (Mejı́a et al. 2011)]. The ascospores
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142
Fungal Diversity (2019) 96:1–242
Fig. 100 continued
of P. salicellum (MFLU 17-1601) did not have an appendage unlike those of the lectotype and epitype of P. salicellum and they were smaller x = 15.8 9 2.5 lm (this
study) vs x = 18.5 9 3.5 lm (Mejı́a et al. 2011)]. However, the length of the ascospores may not be a key distinguishing factor since Mejı́a et al. (2011) observed that
123
length and width of ascospores of P. salicellum may vary,
even within an ascus. The combined data set of ITS, TUB2
and TEF1-a sequences in our phylogenetic analyses has not
clearly separated our strain (MFLU 17-1601) with significant bootstrap support from the strains of P. salicellum (AR
3828) and (LCM 449.01) (Fig. 102). The morphological
Fungal Diversity (2019) 96:1–242
Fig. 101 Plagiostoma salicellum (MFLU 17-1601, new geographical record). a Appearance of ascomata on host substrate. b Close-up
of ascoma on host substrate. c Section of an ascoma. d Section through
143
ostiole. e Peridium. f–h Asci. i–l Ascospores. Scale bars: a = 500 lm,
b, c = 200 lm, d = 100 lm, i–l = 20 lm, e–h = 10 lm
123
144
Fig. 102 Phylogram generated
from maximum likelihood
analysis based on combined
ITS, TUB2 and TEF1-a
sequence data for Plagiostoma
species in Gnomoniaceae.
Related sequences were taken
from Sogonov et al. (2008),
Mejı́a et al. (2011) and Walker
et al. (2014). Sixty-five strains
are included in the combined
analyses which comprise 1948
characters (511 characters for
ITS, 777 characters for TUB2,
660 characters for TEF1-a) after
alignment. Apiognomonia
errabunda (AR 4182)
(Gnomoniaceae, Diaporthales)
is used as the outgroup taxon.
Single gene analyses are carried
out and the topology of each
tree had clade stability. Tree
topology of the maximum
likelihood analysis is similar to
the Bayesian analysis. The best
RaxML tree with a final
likelihood value of
- 12805.418798 is presented.
The matrix had 982 distinct
alignment patterns, with 17.73%
undetermined characters or
gaps. Estimated base
frequencies were as follows:
A = 0.220722, C = 0.299844,
G = 0.226468, T = 0.252965;
substitution rates
AC = 0.958759,
AG = 3.395596,
AT = 0.951907,
CG = 1.027587,
CT = 4.491882,
GT = 1.000000; gamma
distribution shape parameter
a = 0.409235. Bootstrap values
for maximum likelihood (ML)
equal to or greater than 60 are
placed above and below the
branches. Bayesian posterior
probabilities (BYPP) equal to or
greater than 0.90 are in bold.
The newly generated sequence
is indicated in bold and blue
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Fungal Diversity (2019) 96:1–242
differences mentioned above may be the result of phenotypic plasticity arising from a need to adapt to the climatic
conditions in Italy (Jeewon & Hyde 2016). Due to lack of
significant molecular support, we therefore report our strain
as P. salicellum, collected for the first time from Italy. It is
also known from Canada, Europe (Austria, Belgium, Bulgaria, Czech Republic, Germany, Poland, Sweden,
Switzerland, UK and USA (Sogonov et al. 2008).
145
Distoseptispora K.D. Hyde, McKenzie & Maharachch.
Notes: Distoseptispora is typified by D. fluminicola
Mckenzie, H.Y. Su, Z.L. Luo & K.D. Hyde (Su et al. 2016)
and currently 16 species are recorded in Index Fungorum
(2019). In addition, D. adscendens (HKUCC 10820) and D.
leonensis (HKUCC 10822) were also included in this genus
based on evidence of phylogeny (Su et al. 2016). Members
in this genus are commonly encountered in freshwater
habitats and are hyphomycetous, which have olivaceous,
brown or yellowish/reddish brown conidia, euseptate or
distoseptate, obclavate or cylindrical with rounded apex,
and are of indeterminate length (Su et al. 2016; Luo et al.
2018; Tibpromma et al. 2018; Yang et al. 2018b).
macronematous, mononematous, unbranched, erect,
straight or flexuous, cylindrical, smooth-walled, brown to
pale olivaceous, septate, slightly constricted at septa, thinwalled. Conidiogenous cells 13–20 lm long (
x = 16 lm,
n = 10), 3–5 lm wide (
x = 4.5 lm, n = 10), monoblastic
or polyblastic, terminal, determinate, cylindrical, brown to
pale olivaceous. Conidia 17–30 lm long (
x = 25 lm,
n = 15), 7.5–10 lm wide (
x = 9.2 lm, n = 15), acrogenous, solitary, obpyriform to obclavate, broad cylindrical
or irregular, straight or curved, 3–5-distoseptate, olivaceous, thick-walled, smooth, rounded at the apex, truncate
at the base.
Culture characteristics: On PDA, colony circular,
reaching 40 mm in 45 days at 25 C, grey from above,
dark grey from below, surface rough, dry, raised, edge
entire.
Material examined: CHINA, Dehong, on submerged
wood in a stream, 25 November 2017, G.N. Wang H10A
(HKAS 101738, holotype), ex-type living culture
(KUMCC 18-0090).
GenBank numbers: ITS: MK085061, LSU: MK079662,
TEF1-a: MK087659.
Notes: Distoseptispora dehongensis is phylogenetically,
a strongly supported member in Distoseptisporaceae (100
ML/1.00 BYPP) in Fig. 105. Morphological characteristics
of D. dehongensis (such as cylindrical, unbranched, septate
conidiophores and distoseptate conidia) matches well with
those in Distoseptispora. It can be easily distinguished
from the other taxa in Distoseptispora by its fewer distosepta (3–5-distoseptate vs. [ 7-distoseptate) and shorter
length (17–30 lm vs. [ 35 lm) of conidia (Hyde et al.
2016; Su et al. 2016; Yang et al. 2018b; Luo et al. 2018).
Distoseptispora dehongensis has similar conidial dimensions to D. martinii. However, they differ in conidial shape
(Xia et al. 2017). In our multi-gene phylogenetic analyses,
D. dehongensis has a close phylogenetic affinity to D.
obpyriformis and D. rostrata with weak support (Fig. 105),
and they can be distinguished in conidial shape, size and
number of distosepta (Luo et al. 2018).
Distoseptispora dehongensis W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555406; Facesoffungi
number: FoF05057; Fig. 103
Etymology: Name reflects Dehong, from where the
species was isolated.
Holotype: HKAS 101738
Saprobic on decaying wood submerged in freshwater.
Sexual morph Undetermined. Asexual morph Colonies
gregarious or scattered, effuse, hairy, olivaceous. Mycelium
mostly immersed, consisting of branched, septate, smooth,
hyaline hyphae. Conidiophores 45–80 lm long
(
x = 59 lm, n = 10), 4–5 lm wide (
x = 4.5 lm, n = 10),
Distoseptispora palmarum S.N. Zhang, K.D. Hyde & J.K.
Liu, sp. nov.
Index Fungorum number: IF555431; Facesoffungi
number: FoF05090; Fig. 104
Etymology: The epithet reflects the host plant.
Holotype: MFLU 18-1588
Colonies on the substratum superficial, effuse, hairy or
velvety, black. Mycelium mostly immersed, composed of
branched, septate, smooth, pale brown hyphae. Sexual
morph Undetermined. Asexual morph Conidiophores 90–
165 lm long (
x = 125 lm, n = 10), 4–7 lm wide
(
x = 5.5 lm, n = 10), macronematous, mononematous,
septate, unbranched, single or in groups of 2 or 3, erect,
Diaporthomycetidae, families incertae sedis
Distoseptisporaceae K.D. Hyde & McKenzie
Notes: Distoseptisporaceae was established to accommodate a group of sporidesmium-like hyphomycetous taxa,
which are phylogenetically distinct from Sporidesmiaceae
and characterized by darker conidia with slightly paler
rounded apices and indeterminate length, and especially
with most species having relatively short conidiophores (Su
et al. 2016). Subsequently, a series of new species (Hyde
et al. 2016; Yang et al. 2018b; Luo et al. 2018) were
introduced to this monotypic family, and Yang et al.
(2018b) provided an emendation of Distoseptispora
according to much longer, percurrently proliferating conidiophores and euseptate conidia. Two new taxa are introduced to this monotypic family herein, namely
Distoseptispora dehongensis and D. palmarum, which were
found from freshwater and palm habitats, respectively.
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146
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Fungal Diversity (2019) 96:1–242
b Fig. 103 Distoseptispora dehongensis (HKAS 101738, holotype). a–
c Colonies on substrate. d, e Conidiophores. f, g Conidiophores with
conidia. h–m Conidia. n Germinated conidium. o, p Colony on PDA
(o from above view, p from below view). Scale bars: n = 20 lm, d–
m = 10 lm
straight or flexuous, smooth, brown, cylindrical, robust at
the base. Conidiogenous cells polyblastic, integrated,
determinate, terminal, pale brown to brown, cylindrical.
Conidia 35–180 lm long (
x = 76.3 lm, n = 32), 7–11 lm
wide (
x = 8.6 lm, n = 32), acrogenous, dry, oblong,
obclavate, cylindrical or rostrate, elongated, straight or
147
curved, truncate at the base, rounded at the apex, 7–27distoseptate, smooth, greenish black to brown, paler
towards the apex, thick-walled.
Culture characteristics: Colonies on PDA, 20–25 mm
diameter after 21 days at 25 C, mycelium sparse, dark
brown to black, reverse concolourous.
Material examined: THAILAND, Trat Province, Koh
Chang, on rachis of Cocos nucifera (Arecaceae), 27 April
2017, S.N. Zhang, SNT128 (MFLU 18-0588, holotype),
ex-type living culture (MFLUCC 18-1446); ibid. (HKAS
102209, isotype).
Fig. 104 Distoseptispora palmarum (MFLU 18-1588, holotype). a Colonies on substrate surface. b, c Conidiophores with conidiogenous cells
bearing condia. d Conidiophores. e Conidiogenous cells. f–i Conidia. Scale bars: a = 200 lm, b–d, i = 50 lm, f–h = 20 lm, e = 10 lm
123
148
GenBank numbers: ITS: MK085062, LSU: MK079663,
RPB2: MK087670, SSU: MK079661, TEF1-a:
MK087660.
Notes: Distoseptispora palmarum shares similar conidial
features to other species of Distoseptispora, but its conidiophores are longer than most Distoseptispora species (Luo
et al. 2018; Tibpromma et al. 2018; Yang et al. 2018b;
Phookamsak et al. 2019). Distoseptispora palmarum is
similar to D. leonensis (: Ellisembia leonensis)
(McKenzie 1995; Shenoy et al. 2006; Su et al. (2016), D.
suoluoensis and D. guttulata (Yang et al. 2018b). However,
D. palmarum is distinct because of its polyblastic conidiogenous cells. Multi-gene analysis based on LSU, ITS,
TEF1-a and RPB2 indicates that D. palmarum and D.
leonensis constitute a well-supported independent lineage
(Fig. 105). However, D. palmarum differs from D.
Fig. 105 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS, TEF1-a and RPB2 sequence data for
Distoseptisporaceae species and several related families. Related
sequences are taken from Hyde et al. (2016), Su et al. (2016), Xia
et al. (2017), Yang et al. (2018b) and Luo et al. (2018). Thirty-four
strains are included in the combined analyses which comprise 3612
characters (897 characters for LSU, 694 characters for ITS, 946
characters for TEF1-a, 1075 characters for RPB2) after alignment.
Cryphonectria parasitica (CMW 7048) (Cryphonectriaceae, Diaporthales) is used as the outgroup taxon. Single gene analyses are
carried out and the topology of each tree had clade stability. Tree
topology of the maximum likelihood analysis is similar to the
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Fungal Diversity (2019) 96:1–242
leonensis in the length and number of distosepta in the
conidia.
Hypocreomycetidae O.E. Erikss. & Winka
Notes: Eriksson and Winka (1997) introduced this subclass with four orders. In the classification of Eriksson
(2006), the subclass Hypocreomycetidae comprised four
orders (i.e. Coronophorales, Halosphaeriales, Hypocreales
and Microascales) (Maharachchikumbura et al.
2015, 2016). Most of the taxa of the Hypocreomycetidae
have light coloured perithecia, nonamyloid apical rings in
the asci when apical rings are present and the absence of
true paraphyses (Zhang et al. 2006)
Glomerellales Chadef. ex Reblova et al.
Notes: We follow the updated classification by
Wijayawardene et al. (2018a).
Bayesian analysis. The best RaxML tree with a final likelihood value
of - 15208.000555 is presented. The matrix had 1181 distinct
alignment patterns, with 45.45% undetermined characters or gaps.
Estimated base frequencies were as follows: A = 0.240242,
C = 0.262790, G = 0.284086, T = 0.212881; substitution rates AC =
0.955762, AG = 2.559225, AT = 1.264794, CG = 0.896334, CT =
7.954863, GT = 1.000000; gamma distribution shape parameter
a = 0.214652. Bootstrap values for maximum likelihood (ML) equal
to or greater than 60 and Bayesian posterior probabilities (BYPP)
equal to or greater than 0.95 are placed above the branches,
respectively. The ex-type strains are in bold and black. The newly
generated sequences are indicated in bold and blue
Fungal Diversity (2019) 96:1–242
Plectosphaerellaceae W. Gams et al.
Notes: Several asexual morph genera in this family have
verticillate conidiophores such as Acrostalagmus Corda
and Verticillium Nees. Stachylidium bicolor, the type of its
genus, produces erect, roughened, verticillate conidiophores, often with additional verticillate axes emerging
from the main stipe; this results in a more complex conidiophore than in other similar genera (Réblová et al. 2011).
Acrostalagmus Corda
Notes: Based on the phylogenetic analyses of Réblová
et al. (2011), the common tropical hyphomycete described
and illustrated by Seifert (1985) as Stilbella annulata was
revealed as a member of Plectosphaerellaceae and a sister
species to Acrostalagmus luteoalbus, the type of the genus.
Both Stilbella annulata and Acrostalagmus luteoalbus
149
produce ameroconidia in bright orange to reddish slimy
masses; in both species the reddish pigmentation sometimes also colours the phialides.
Acrostalagmus annulatus (Berk. & M.A. Curtis) Seifert,
Studies in Mycology 68: 186 (2011)
Facesoffungi number: FoF05253; Fig. 106
Saprobic on decaying fruit. Sexual morph Undetermined. Asexual morph Hyphomycetous. forming rounded
pale reddish brown slimy heads, oval. Conidiophore axis
erect, repeatedly branched, pale reddish brown at the base
and hyaline at the apex, full-grown conidiophore stipes 4–
5 lm wide at the base, tapering to 3 lm. Conidiogenous
cells phialidic 9–26 9 2–3.5 lm, narrowly flask-shaped in
the widest part, arising in whorls of 1–5 at several levels
along the main stipe and its branches. Conidia 6–8 9 3–
Fig. 106 Acrostalagmus annulatus (MFLU 16-0952, new geographical record). a Host fruit. b, c Conidiomata on host surface.
d Conidiophores. e–g Conidiogenus cells and conidiophores with conidia. h, i Conidia. Scale bars: d = 20 lm, e–i = 10 lm
123
150
Fungal Diversity (2019) 96:1–242
4 lm (
x = 7.4 9 3.6 lm, n = 30), hyaline, sub-globose to
cylindrical, single-celled.
Culture characteristics: Conidia germinated on MEA.
On MEA colonies are appressed, circular, flat surface, edge
entire, first cream then become dark brown and rise in the
centre with mycelium, reverse brown reaching 2 cm in
2 weeks at 18 C.
Material examined: THAILAND, Chiang Mai Province,
Doi Suthep, 22 December 2015, decaying fruits of
unknown species, Subashini C. Jayasiri, C 132 (MFLU
16-0952), living culture (MFLUCC 16-0612).
GenBank numbers: ITS: MK607614, LSU: MK607617,
RPB2: MK607616, SSU: MK607615.
Notes: The new isolate of Acrostalagmus annulatus fits
with the type description in having erect, roughened, verticillate conidiophores, phialidic conidiogenous cells and
oblong-ellipsoidal conidia accumulate in slime (Seifert
1985; Réblová et al. 2011). Acrostalagmus annulatus
(MFLUCC 16-0612) forms a sister clade to A. annulatus
(DAOM 212126) and an environment sample with high
statistical support (100% ML/1.00 BYPP) in Fig. 107. In
here based on morphology and phylogenetic affinity
introduced a new record of Acrostalagmus annulatus from
Thailand.
Fig. 107 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data of selected taxa of
Plectosphaerellaceae. Related sequences were obtained from GenBank. Twenty-two strains are included in the analyses, which
comprise 1299 characters including gaps. Tree was rooted to
Hypomyces aurantius (GJS74 69) and Trichoderma viride (DAOM
JBT1003). Tree topology of the ML analysis is similar to the BI. The
best scoring RAxML tree with a final likelihood value of
- 4999.737002 is presented. The matrix had 345 distinct alignment
patterns, with 41.99% of undetermined characters or gaps. Estimated
base frequencies were as follows; A = 0.222188, C = 0.278776,
G = 0.279137, T = 0.219898; substitution rates AC = 0.918516,
AG = 1.868080, AT = 2.187888, CG = 0.896478, CT = 5.118188,
GT = 1.000000; gamma distribution shape parameter a = 0.637862.
Maximum likelihood bootstrap (MLBS) values [ 70% and Bayesian
posterior probabilities (BYPP) C 0.95% are given near the nodes
respectively. The scale bar indicates 0.04 changes. The ex-type strains
are in bold and black. The newly generated sequence is indicated in
blue
123
Hypocreales Lindau
Notes: The order Hypocreales (Hypocreomycetidae)
comprise highly diverse species in the tropics, subtropics
and temperature regions (Põldmaa 2011) and compirse the
families Bionectriaceae, Clavicipitaceae, Cordycipitaceae,
Flammocladiaceae, Hypocreaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae, Stachybotriaceae and
Tilachlidiaceae with 237 genera (Kirk et al. 2008;
Maharachchikumbura et al. 2015, 2016)
Fungal Diversity (2019) 96:1–242
Cordycipitaceae Kreisel ex G.H. Sung et al.
Notes: The family Cordycipitaceae was first described
by Kreisel (1969). However, Cordycipitaceae was validly
segregated from Clavicipitaceae by Sung et al. (2007)
based on morphological and multi-gene phylogenetic
analyses. Most of the species in the family are entomogenous and produce stalked, erect, stromatic ascomata or
reduced stipes or subiculate stromata (Kepler et al. 2017).
Stroma are fleshy or subiculate and pallid or brightly
coloured (Sung et al. 2007; Kepler et al. 2017). Kepler
et al. (2017) included 11 genera in this family.
Wijayawardene et al. (2017a) included 17 genera in this
family.
Cordyceps Fr.
Notes: Cordyceps was validly published by Link (1833)
and formally moved to Cordycipitoideae by Sung et al.
(2007) with the type species Cordyceps militaris (L.) Fr.
Kepler et al. (2017) synonymised Isaria Pers., Microhilum
H.Y. Yip & A.C. Rath., Phytocordyceps C.H. Su & H.H.
Wang and Evlachovaea B.A. Borisov & Tarasov under
Cordyceps using phylogentically-based results.
Cordyceps succavus Y.P. Xiao, T.C. Wen & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555420; Facesoffungi
number: FoF05065; Fig. 108
Etymology: The specific epithet refers to the feature of
the stromata (inside hollow after mature).
Holotype: MFLU 18-1890
Parasitic in an unknown insect buried in the upper 1 cm
of soil, forming yellowish to yellow stromata. Sexual
morph: Stromata 4–5 cm long, 3–6 mm wide, yellowish
to yellow, fleshly, mostly solitary, stipitate, inside hollow
after mature. Stipe 2–3 cm long, 3-5 mm wide, fleshly,
yellowish to white, clavate, with stromata on the top.
Fertile head 1.5–2 cm long, 4–5 mm wide, fleshly, solitary, cylindrical, yellow to yellowish. Stroma 3–3.4 9 2.4–
2.7 mm (
x = 3.2 9 2.6 lm, n = 30) in vertical section,
yellow to yellowish. Perithecia 534–655 9 179–278 lm
(
x = 594 9 229 lm, n = 30), half-embedded, ampulliform
to ovoid, yellow to yellowish, thick-walled, with ostiole on
the top. Peridium 23–36 lm wide (
x = 29 lm, n = 60),
two layers, textura porrecta in first layer, textura angularis
in secondary layer. Asci 486–600 9 3.6–4.9 lm
(
x = 543 9 4.3 lm, n = 60), 8-spored, hyaline, narrow
cylindrical, with a thick apex. Apical cap 3.1–4.1 9 3.3–
4.5 lm (
x = 3.6 9 3.9 lm, n = 60) lm diam., hyaline.
Ascospores 466–594 9 0.9–1.2 lm (
x = 530 9 1.1 lm,
n = 60), filiform, smooth, hyaline, septate, breaking into
secondary spores. Secondary spores 2.8–4.9 9 0.9–1.2 lm
(
x = 3.8 9 1.1 lm, n = 60), cylindrical, smooth, hyaline,
one-celled. Asexual morph: Undetermined.
151
Material examined: THAILAND, Chiang Mai Province,
Samoeng, on larvae (Lepidopteran), 10 July 2017, Y.P.
Xiao, CM0752 (MFLU 18-1890, holotype); ibid.
CM0752b, HKAS 96376 (MFLU 18-1891, isotype).
GenBank numbers: ITS: MK086060, LSU: MK086062,
RPB1: MK084616, RPB2: MK079353, SSU: MK086058.
Notes: According to morphology and phylogenetic
analysis (Fig. 110), the new collection is closely related to
Cordyceps grylli, which was collected from adults of
Gryllidae in China (Teng 1936; Mains 1959), but the
phylogenetic analyses supports two different species.
Cordyceps succavus is characterized by yellow to yellowish, longer stromata, not becoming grayish when dry, yellow ampulliform to ovoid perithecia, longer asci and
shorter ascospores (Table 3). Cordyceps grylli has yellow
stromata, becoming grayish when dry, with half-embeded,
light brown or chestnut brown, ovoid perithecia, cylindrical
asci and filiform ascospores breaking into 1-celled secondary spores (Mains 1959) (Table 3). Therefore, we
propose Cordyceps succavus as a new species.
Blackwellomyces Spatafora & Luangsa-ard
Notes: This genus was introduced by Kepler et al.
(2017) with two species with ascospores not disarticulating
into secondary spores at maturity and with irregularly
spaced septa.
Blackwellomyces lateris Y.P. Xiao, T.C. Wen & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555421; Facesoffungi
number: FoF05066; Fig. 109
Etymology: The specific epithet refers to the feature of
the stroma (unilateral on the stipe).
Holotype: MFLU 18-0663
Parasitic in a larvae (Lepidoptera) buried in the upper
2 cm of soil, dark to dark-brown host, forming yellow to
yellowish stromata. Sexual morph Thallus within host,
without mycelium cover. Stromata 5–6 cm long, 1.5–
2 mm wide, fleshly, mostly branched, yellow to yellowish,
stipitate arising from head of the host. Stipe 3.5–4 cm long,
1.5 mm wide, fleshly, yellow to yellowish, clavate, with
stromata on the side face. Fertile head 0.5–2.5 cm long,
1.5–2 mm wide, fleshly, solitary or branched, cylindrical,
yellow to yellowish, unilateral on the stipe. Stroma 713–
822 9 939–1106 lm (
x = 767 9 1023 lm, n = 30) in
vertical section, yellow to yellowish, unilateral. Perithecia
287–343 9 138–184 lm (
x = 315 9 161 lm, n = 30),
embedded, ampulliform to ovoid, yellow to yellowish,
thick-walled, with ostiole on the top. Peridium 17–29 lm
wide (
x = 23 lm, n = 60), three layers, textura porrecta in
first layer, textura angularis in middle layer, textura
intricata in third layer. Asci 166–223 9 3.7–5.1 lm
(
x = 195 9 4.4 lm, n = 60), 8-spored, hyaline, narrow
cylindrical, with a thin apex. Apical cap 2.2–3.0 9 3.3–
123
152
Fungal Diversity (2019) 96:1–242
Fig. 108 Cordyceps succavus (MFLU 18-1890, holotype). a Habitat
of Cordyceps succavus. b Stromata without infected insect. c Vertical
section of stroma. d Vertical sections showing the semi-immersed
perithecia. e Perithecium wall. f, g Asci. h Part of ascospores. i Apical
cap of asci. j, k Secondary ascospores. Scale bars: c, d = 500 lm, f,
g = 200 lm, h = 50 lm, e = 20 lm, i–k = 5 lm
4.4 lm (
x = 2.2 9 3.3 lm, n = 60), hyaline. Ascospores
160–217 9 1.0–1.5 lm
(
x = 188 9 1.2 lm,
n = 60)
cylindrical, smooth-walled, hyaline, without septa, not
breaking into secondary part spores. Asexual morph
Undetermined.
Material examined: THAILAND, Chiang Mai Province,
The Mushroom Research Centre, on larvae (Lepidopteran),
12 June 2017, Y.P. Xiao, MRC170617 (MFLU 18-0663,
holotype); ibid. MFLU 18-0664, MRC170617b, (HKAS
96376, isotype).
GenBank numbers: ITS: MK086059, LSU: MK086061,
RPB1: MK084615, RPB2: MK079354, SSU: MK086057,
TEF1-a: MK069471.
Notes: According to morphological and phylogenetic
analysis (Fig. 110), Blackwellomyces lateris is closely
related to B. cardinalis and B. pseudomilitaris, and characterized by yellow to yellowish, longer stromata; yellow
to yellowish, longer fertile head with stromata in side face;
embedded, ampulliform to ovoid perithecia; shorter asci;
aseptate ascospores (Table 4). Molecular data indicate that
Blackwellomyces lateris has 16 bp in ITS region, 2 bp in
nrSSU, 10 bp in nrLSU, 11 bp in TEF1-a, 12 bp in RPB1,
123
This study
1-celled,
2.8–4.9 9 0.9–1.2
Cylindrical
486–600 9 3.6–4.9
15–20 long, 4–5
wide, fleshly,
solitary,
cylindrical, yellow
to yellowish
20–30
long,
3–5
wide
Half-embedded,
ampulliform to ovoid,
yellow to yellowish
534–655 9 179–278
Filiform,
multiseptate,
466–594 9 0.9–1.2
Zang and
Kinjo
(1998)
1-celled, 4–5 9 1
Cylindric,
300–400 9 4
1–2
thick
Ovoid, partly embedded
(up to one-half), light
brown or chestnut-brown
400–700 9 300–350
Filiform, break into
fragments
References
Ascospores (lm)
Asci (lm)
Perithecia (lm)
Stipe
(mm)
Yellow when
fresh
becoming
grayish
Yellowish to
yellow
4–5 cm
long,
3–6 mm
wide
Gryllidae
(Orthoptera)
Unknown
insect
C. grylli
C.
succavus
Fertile head (mm)
Stromata
Host
Species
Table 3 Synopsis of Cordyceps species discussed in the paper
153
Secondary spores
(lm)
Fungal Diversity (2019) 96:1–242
20 bp in RPB2 that are different from B. cardinalis (OSC
93610), and has 30 bp in ITS region, 17 bp in nrSSU,
10 bp in nrLSU, 35 bp in RPB1 that are different from B.
pseudomilitaris (NBRC 101509). Blackwellomyces lateris
has good support in the phylogenetic tree and, therefore,
we propose it as a new species.
Hypocreales genera incertae sedis
Sarocladium W. Gams & D. Hawksw.
Notes: Sarocladium was erected by Gams and Hawksworth (1975) and S. oryzae selected as the type species.
Sarocladium is classified in Hypocreales, family incertae
sedis (Maharachchikumbura et al. 2015, 2016). This genus
encompasses 22 species according to Index Fungorum
(2019). Sarocladium attenuatum was however, confirmed
as synonym of S. oryzae (Bills et al. 2004). Therefore, 20
species are accepted. Sarocladium species resembles
Acremonium species, but differ in their elongated, solitary
phialides, sparsely or repeatedly branched conidiophores,
abundant adelophialides and elongated conidia (Giraldo
et al. 2015). Colony and conidial morphology can be the
primary characters to identify species. Phylogenetic analysis based on a LSU-SSU and LSU-ITS dataset resolves
intraspecific identification (Summerbell et al. 2011; Giraldo et al. 2015; Liu et al. 2017). Species of this genus are
rice plant pathogens (Saravanakumar et al. 2009), saprobes,
mutualistic endophytes (Yeh and Kirschner 2014) and
human opportunistic pathogens (Giraldo et al. 2015). Some
endophytes even have a potential as biological control
agents against plant pathogens (Kelemu et al. 2001).
Sarocladium kiliense (Grütz) Summerb, Stud. Mycol. 68:
158 (2011)
: Acremonium kiliense Grütz, Dermatol. Wochenschr.
80:774. 1925.
Facesoffungi number: FoF05816; Fig. 111
Holotype: CBS 122.29
Culture characteristics: Colonies attaining a diam of
2.6 cm in 12 days at 22 C, entire margin, circular, dense,
moist, with radial wrinkles on the surface, pink from above
and reverse. Mycelium strands form at the central of
colonies from which the phialides are produced. Phialides
20–46 9 1–2 lm (
x = 34 9 1.7 lm, n = 50) phialidic,
slender, tapering towards the apex. Conidia 2–6 9 1–2 lm
(
x = 4 9 1.5 lm, n = 100), abundant, cylindrical to elliptical, asepate, hyaline, smooth-walled, straight to curved,
with round end, tending to gather in slimy head at the tip of
phialide.
Material examined: CHINA, Yunnan Province, Kunming Institute of Botany, on scarred leaves of an unidentified plant, 10 September 2017. Rungtiwa Phookamsak,
Dry culture (HKAS 101476), living culture (KUMCC
18-0031).
123
154
Fungal Diversity (2019) 96:1–242
Fig. 109 Blackwellomyces lateris (MFLU 18-0663, holotype).
a Habitat of Blackwellomyces lateris. b Overview of the host and
stromata. c Host. d, e Stromata. f Vertical section of stroma.
g Vertical sections showing the immersed perithecia. h Perithecium.
i–k Asci. l Apical cap of asci. m, n Ascospores. Scale bars: b–
d = 1 cm, e = 2 mm, f, g = 200 lm, i–k = 100 lm, m = 50 lm, h, l,
n = 20 lm
GeneBank numbers: ITS: MK616348, LSU: MK616474.
Notes: Our isolate has a close affinity with Sarocladium
kiliense (CBS 122.29) based on the analysis of a combined
LSU and ITS sequence data (Fig. 112). Comparison of ITS
sequence showed 1/519 bp nucleotides differ from the type
strain of S. kiliense (CBS 122.29 = MUCL 9724). Sarocladium kiliense is a combined species which initially
proposed by Summerbell et al. (2011) based on the
phylogenetic analysis of a combined LSU and SSU
sequences. The morphological descriptions of Sarocladium
kiliense such as strain MUCL 9724 (Perdomo et al. 2011)
and strain IR5 (Ebrahimi and Fotouhifar 2016) were given.
Our isolates morphologically resemble type strain MUCL
9724 by having cylindrical, smooth, hyaline, conidia with
rounded ends which is in slimy heads and dirty white to
pale orange colony, but differ in absent of adelophialides
123
Fungal Diversity (2019) 96:1–242
155
Fig. 110 Phylogram generated from maximum likelihood analysis
based on combined ITS, SSU, LSU, TEF1-a, RPB1 and RPB2
sequence data for Ophiocordyceps species and Blackwellomyces.
Related sequences are taken from Sung and Spatafora (2004), Sung
et al. (2007) and Kepler et al. (2017). Fifity-seven strains are included
in the combined genes sequence analyses which comprise total 4558
characters (1002 characters for SSU, 716 characters for LSU, 494
characters for ITS, 920 characters for TEF1-a, 570 characters for
RPB1 and 856 characters for RPB2) after alignment. Purpureocillium
lilacinum (CBS 284.36) and P. lilacinum (CBS 431.87) are used as
outgroup taxa. Tree topology of the maximum likelihood analysis is
similar to the Bayesian analysis. The best sorting RaxML tree with a
final likelihood value of - 37381.826537 is presented. The matrix
had 1832 distinct alignment patterns, with 29.69% undetermined
characters or gaps. Estimated base frequencies were as follows:
A = 0.244626, C = 0.262958, G = 0.266932, T = 0.225484; substitution rates AC = 1.386955, AG = 3.678399, AT = 1.009103, CG =
1.080113, CT = 7.048074, GT = 1.000000; gamma distribution
shape parameter a = 0.209990. Bootstrap values for maximum
likelihood and Bayesian equal to or greater than 75 are placed above
and below the branches respectively. The newly generated sequences
are indicated in bold and blue
and chlamydospores. Our isolates also similar as strain IR5
in colonial characters as well as the shape and size of
phialides (tapering toward apex, 19–48 9 1–2 lm) and
conidia (cylindrical with round end, 3–7 9 1–2 lm), but
differ in absenting of chlamydospores (Ebrahimi and
Fotouhifar 2016). Historically, Sarocladium kiliense can be
an endophytes (strain CanL-10b, China) on Brassica napus
(Zhang et al. 2014b), a pathogen such as strain IR5 (Iran)
on Malus domstica leaf (Ebrahimi and Fotouhifar 2016),
strain CBS 400.52 (England) on Ficus carica (Lombard
et al. 2015), strain CBS 122.29 (Germany) on skin infection of man (Herrera et al. 2013), strain FMR 10426 (USA)
on blood of human or animal (Irinyi et al. 2015). Sarocladium kiliense have close association with human
infections and plant scab, the detailed morphological
description is necessary for rapid diagnosis of agent which
causes disease (Perdomo et al. 2011). Herewith we introduce our isolate as a new collection of Sarocladium
kiliense.
Pleurotheciales Réblová & Seifert
Notes: Réblová et al. (2016) introduced the order Pleurotheciales with a single family Pleurotheciaceae Réblová
& Seifert based on morphological data and phylogenetic
analyses.
Pleurotheciaceae Réblová & Seifert
Notes: Members of the family Pleurotheciaceae share
the following morphological characters: perithecial, astromatic, immersed to superficial ascomata, a hamathecium of
paraphyses, unitunicate asci, with a non-amyloid apical
ring and hyaline or versicolorous, transversely multi-septate ascospores with polar cells hyaline or brown central
123
This study
Aseptate,
160–217 9 1.0–1.5
166–223 9 3.7–5.1
Embedded, ampulliform to
ovoid, 287–343 9 138–184
Reddish orange
to reddish,
Orange
15–30 9 0.9–3
Yellow to
yellowish,
50–60 9 1.5–2
Lepidoptera
Lepidoptera
Lepidoptera
B. cardinalis
B.
pseudomilitaris
B. lateris
Reddish orange to
reddish,
cylindrical,
elliptical to
fusiform,
2–9 9 1–4
Orange to red,
cylindrical to
enlarging apically
4–50 9 0.5–1.5
Yellow to yellowish,
clavate, with
stromata on the
side face
35–40 9 1.5
Yellow to
yellowish,
cylindrical,
elliptical to
fusiform,
5–25 9 1.5–2
HywelJones
(1994)
Multiseptate,
280–390 9 1
290–410 9 5–6
Superficial with one-quarter to
one-third immersed, elongate
ellipsoid to elongate ovoid,
290–570 9 120–245
Orange, cylindrical,
2–8 9 1.2–4
Sung and
Spatafora
(2004)
175–330 9 3–5
Embedded, elliptical to
fusiform to obclavate,
230–540 9 110–240
Irregularly
multiseptate,
160–320 9 1
Asci
(lm)
Fertile head (mm)
Stromata
Host
Species
Table 4 Synopsis of Blackwellomyces species discussed in this paper
Stipe (mm)
Perithecia (lm)
Reference
Fungal Diversity (2019) 96:1–242
Ascospores (lm)
156
123
cells. The asexual morph is characterized by macronematous or semi-macronematous conidiophores, loosely fasciculate or aggregated in indeterminate synnemata, and
conidiogenous cells producing conidia holoblastically,
conidial secession rhexolytic on short denticles or rachis on
sympodially extending polyblastic conidiogenous cells and
conidia hyaline, brown or versicolorous, septate or aseptate
(Réblová et al. 2016).
Phaeoisaria Höhn.
Notes: Phaeoisaria was introduced by von Höhnel (1909)
for a collection on Gigantochloa sp. (Bambusae) and is
typified by P. bambusae Höhn. This dematiaceous hyphomycete genus presently includes 19 species that produce
indeterminate synnemata with septate or aseptate ellipsoidal,
obovoidal, fusiform-cylindrical or falcate conidia formed on
a sympodially extending rachis, occurring on decaying
wood, plant debris or soil sediments (Sutton 1973; Deighton
1974; Castañeda et al. 2002; Seifert et al. 2011; Mel’nik
2012; Cheng et al. 2014; Liu et al. 2015; Réblová et al. 2016).
Réblová et al. (2012) described the genus Pleurotheciella for
two new species, P. rivularia and P. centenaria.
Phaeoisaria siamensis Jayasiri & K.D. Hyde, sp. nov.
Index fungorum number: IF555710; Facesoffungi number: FoF05324; Fig. 113
Etymology: With reference to country (‘Siam’ earlier
name of Thailand) where the specimen was collected.
Holotype: MFLU 16-0953
Saprobic on decaying fruits. Sexual morph Undetermined. Asexual morph Conidiomata scattered, indeterminate, erect, rigid, superficial, dark brown composed of
compact appressed conidiophores. Conidiophores 330–
380 lm high, 20–25(–30) lm (
x = 360 9 24.5 lm,
n = 20) wide, macronematous, in synnematous conidiomata, scattered, synnemata subulate or cylindrical,
indeterminate, at the base 13–15 lm beneath the fertile
portion with conidiogenous cells, composed of medium to
dark brown, smooth, septate parallel hyphae, 2–2.5(–3) lm
wide, splaying out at the middle to apex. Conidiogenous
cells 8–12 9 2–2.5 lm (
x = 10.5 9 2.3 lm, n = 20),
integrated, terminal and intercalary, poly-blastic, sympodial, cylindrical or somewhat attenuated toward the tip,
pale brown near base, subhyaline at the apex, denticulate,
denticles conspicuously cylindrical, 2–2.5 9 1–1.5 lm.
Conidia 5–8 9 3–4 lm (
x = 6.2 9 3.5 lm, n = 30), globose to subglobose, hyaline, aseptate.
Culture characteristics: Conidia germinated on MEA.
Colonies are appressed, circular, flat surface, edge entire,
first cream then become dark brown and rise in the centre
with mycelium, reverse brown reaching 10 mm in 2 weeks
at 18 C.
Material examined: THAILAND, Chiang Mai Province,
Doi Suthep, 22 December 2015, decaying fruits of
Fungal Diversity (2019) 96:1–242
Fig. 111 Sarocladium kiliense (HKAS 101476, new record). a, b
Upper and reverse view of culture on PDA after 17 days. c Mycelium
strand form at centra of colony. d Phialides arising from mycelium
157
strand. e, f Phialides bearing conidia. g–j Conidia. Scale bars:
d = 100 lm, e, f = 15 lm, g–j = 3 lm
123
158
Fungal Diversity (2019) 96:1–242
Fig. 112 Phylogram generated from maximum likelihood analysis
based on combined LSU and ITS sequence data for Sarocladium
species. Related sequences were referred to Liu et al. (2017). Twentytwo strains are included in the combined genes sequence analyses
which comprise total 1086 characters (517 characters for LSU, 569
characters for ITS) after alignment. Acremonium curvulum (CBS
430.66) (Incertae sedis, Hypocreales) is used as the outgroup taxon.
Single gene analyses are carried out and the topology of each tree
with clade stability. Tree topology of the maximum likelihood
analysis is similar to the maximum parsimony analysis and the
Bayesian analysis. The best sorting RaxML tree with a final
likelihood value of - 4666.270461 is presented. The matrix had
300 distinct alignment patterns, with 3.01% undetermined characters
or gaps. Estimated base frequencies were as follows: A = 0.232286,
C = 0.271554, G = 0.293432, T = 0.202727; substitution rates AC =
1.861270, AG = 2.197180, AT = 2.893484, CG = 0.997016, CT =
6.880061, GT = 1.000000; gamma distribution shape parameter
a = 0.179594. Maximum parsimony analysis of 791 constant characters and 142 informative characters resulted in two equally most
parsimonious tree of 153 steps (CI = 0.614, RI = 0.643, RC = 0.395,
HI = 0.386). Bootstrap values for maximum likelihood (ML) and
maximum parsimony (MP) equal to or greater than 50 are placed
above and below the branches respectively. Branches with Bayesian
posterior probabilities (BYPP) equal or greater than 0.90 are in bold.
The ex-type strains are in bold and black. The newly generated
sequence is indicated in bold and blue
Meliaceae sp., Subashini C. Jayasiri, C 128 (MFLU
16-0953, holotype), ex-type living culture (MFLUCC
16-0607).
GenBank numbers: ITS: MK607610, LSU: MK607613,
RPB2: MK607611, SSU: MK607612.
Notes: Phaeoisaria siamensis fits with the generic
description of Phaeoisaria in having indeterminate synnemata with aseptate ellipsoidal, obovoidal, fusiformcylindrical or falcate conidia formed on a sympodially
extending rachis. Phaeoisaria siamensis (MFLUCC
16-0607) forms a sister clade to P. aquatica (MFLUCC
16-1298) with high statistical support (99% MLBS/1.0
BYPP) in Fig. 114. Phaeoisaria aquatica differs from P.
siamensis in having compactly and parallelly adpressed
conidiophores, with flared conidiogenous cells in the above
half (Luo et al. 2018). A comparison of the ITS and RPB2
nucleotides of these two strains reveals 8 (1.6%) and 13
(1.5%) nucleotide differences which justifies the new species following the guidelines of Jeewon and Hyde (2016).
123
Savoryellales Boonyuen, Suetrong, Sivichai, K.L. Pang &
E.B.G. Jones
Fungal Diversity (2019) 96:1–242
Notes: The order Savoryellales was introduced by
Boonyuen and Pang (2011) to accommodate three sexual
genera viz. Ascotaiwania Sivan. & H.S. Chang, Ascothailandia Sri-indr., Boonyuen, Sivichai & E.B.G. Jones (now
Canalisporium Nawawi & Kuthub.), Savoryella E.B.G.
Jones & R.A. Eaton and an asexual genus Canalisporium
based on multi-gene phylogenies. It was placed in subclass
Hypocreomycetidae (Sordariomycetes) (Boonyuen and
Pang 2011), and subsequently raised to subclass
Savoryellomycetidae (Sordariomycetes) by Hongsanan
et al. (2017) based on phylogenetic analyses and its stem
age (267 MYA) reported in Hyde et al. (2017a). This was
also confirmed by Dayarathne et al. (2019a).
Savoryellaceae Jaklitsch & Réblová
Notes: The family Savoryellaceae has a worldwide
distribution in freshwater, marine and brackish water
habitats (Ranghoo 1998; Abdel-Wahab and Jones 2000;
Cai et al. 2003; Jones et al. 2015). The family was
reviewed by Dayarathne et al. (2019a) with morphomolecular and molecular clock analyses. It comprises three
genera Ascotaiwania Sivan. & H.S. Chang, Canalisporium
Nawawi & Kuthub. and Savoryella E.B.G. Jones & R.A.
Eaton (Dayarathne et al. 2019a).
Canalisporium Nawawi & Kuthub.
Notes: Canalisporium was proposed to accommodate
Berkleasmium caribense Hol.-Jech. & Mercado, B. pulchrum Hol.-Jech. & Mercado and C. elegans Nawawi &
Kuthub. and typified by C. caribense (Hol.-Jech. & Mercado) Nawawi & Kuthub. Twelve species are accepted in
the genus, nine of which have molecular data in GenBank.
Canalisporium is characterized by muriform conidia and
with sexual morph formerly known in Ascothailandia (Sriindrasutdhi et al. 2010).
Canalisporium dehongense W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555407; Facesoffungi
number: FoF05058; Fig. 115
Etymology: Name reflects Dehong, from where the
species was isolated.
Holotype: MFLU 18-1189
Saprobic on decaying wood submerged in freshwater.
Sexual morph Undetermined. Asexual morph Conidiomata sporodochial, scattered, punctiform, pulvinate,
granular, black, shiny. Mycelium immersed in natural
substrate, consisting of branched, septate, thin-walled,
smooth, hyaline to pale brown hyphae. Conidiophores up
to 65 lm long, micronematous, mononematous, vesiculate,
consisted of 1–9 subglobose, smooth, hyaline cells,
unbranched, septate, constricted at the septa. Conidiogenous cells 7–16 9 7–14 lm (
x = 12 9 10 lm, n = 15),
holoblastic,
monoblastic,
terminal,
determinate,
159
subglobose, ellipsoidal, sometimes swelling to globose,
hyaline, smooth, thin-walled. Conidia 20–30 9 12–19 lm
(
x = 25 9 16 lm, n = 40), solitary, acrogenous, ellipsoidal to obovoid, muriform, smooth, brown, 1 straight
column of vertical septa and 3–5 rows of transverse septa,
slightly constricted at the septa, darkened and thickly
banded at the septa, canals in the septa obscured by dark
pigmentation; basal cell single, cuneiform, sometimes
swollen, pale brown to hyaline.
Culture characteristics: On PDA, colony circular,
10 mm in 40 days at 25 C, brown to grey from above,
black from below, raised, felty wooly, fairly dense.
Material examined: CHINA, Yunnan Province, on
submerged wood in a small river, 25 November 2017, G.N.
Wang, H158D (MFLU 18-1189, holotype), ex-type living
culture, (MFLUCC 18-1396), ibid.; H158D (HKAS
101725, isotype), ex-isotype living culture (KUMCC
18-0075).
GenBank numbers: ITS: MK051033, LSU: MK051034,
SSU: MK051035.
Notes: Canalisporium dehongense is morphologically
similar to C. caribense (Hol.-Jech. & Mercado) Nawawi &
Kuthub. and C. exiguum Goh & K.D. Hyde in having
muriform conidia with one longitudinal septum, two cells
at the apex and darkened bands at the septa (Goh et al.
1998b; Sri-indrasutdhi et al. 2010). However, it differs
from
C.
caribense
by
its
smaller
conidia
(20–30 9 12–19 lm vs. 24–51 9 15–29 lm) and from C.
exiguum by having more rows of transverse septa (3–5 vs.
2–3). A comparison of nucleotides of C. dehongense with
C. caribense SS03839, C. caribense SS03683 and C. exiguum SS00809 also reveals a difference of 94, 101 and
82 bp in the ITS gene region, respectively. The species
clusters as a distinct clade within the genus Canalisporium
(Fig. 116) with high statistical support (0.99 BYPP).
Sordariomycetidae O.E. Erikss. & Winka
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A.
Fernández
Notes: The order Chaetosphaeriales was introduced by
Huhndorf et al. (2004) to accommodate Chaetosphaeriaceae Réblová, M.E. Barr & Samuels. Maharachchikumbura et al. (2015) indicated that Chaetosphaeriaceae and
Helminthosphaeriaceae Samuels, Cand. & Magni were
sister families within Chaetosphaeriales clade. Subsequently, Konta et al. (2017) introduced another two families, Leptosporellaceae Konta & K.D. Hyde and
Linocarpaceae Konta & K.D. Hyde into Chaetosphaeriales
based on LSU and ITS sequence data. Besides these four
families, Chaetosphaeriales also comprises some genera
incertae sedis, e.g. Caudatispora J. Fröhl. & K.D. Hyde,
Erythromada Huhndorf, A.N. Mill., F.A. Fernández &
123
160
Fungal Diversity (2019) 96:1–242
Fig. 113 Phaeoisaria siamensis (MFLU 16-0953, holotype). a Host fruit. b, c Conidiomata on host surface. d Conidiophore. e–g Conidiogenous
cells arranged on conidiophores. h Conidia. Scale bars: d = 100 lm, e–g = 10 lm
Lodge, and Lasiosphaeriella Sivan. (Maharachchikumbura
et al. 2015, 2016).
Chaetosphaeriaceae Réblová, M.E. Barr & Samuels
Notes: Thirty-eight genera are presently accepted in
Chaetosphaeriaceae (Maharachchikumbura et al. 2015,
2016; Wijayawardene et al. 2018a; Yang et al. 2018a).
Recent studies are those of Liu et al. (2016), Lu et al.
(2016), Luo et al. (2016, 2019), Wei et al. (2018) and Yang
et al. (2016, 2018).
123
Sporoschisma Berk. & Broome
Notes: Most Sporoschisma species have been reported
from freshwater habitats (Goh et al. 1997; Ho et al. 2002;
Luo et al. 2016, 2019; Yang et al. 2016a). Thirteen species
are accepted within this genus (Luo et al. 2016, 2019).
Sporoschisma
resides
in
Chaetosphaeriaceae
(Chaetosphaeriales)
(Maharachchikumbura
et
al.
2015, 2016).
Sporoschisma chiangraiense N.G. Liu & K.D. Hyde, sp.
nov.
Index Fungorum number: IF555386; Faceoffungi number: FoF04950; Fig. 117
Fungal Diversity (2019) 96:1–242
161
Fig. 114 Simplified phylogram showing the best RAxML maximum
likelihood tree obtained from the combined SSU, ITS, LSU and RPB2
matrix of 55 taxa of the orders Pleurotheciales, Savoryellales,
Conioscyphales and Fuscosporellales. The matrix comprised 3920
characters including alignment gaps. The tree was rooted to Leotia
lubrica (AFTOLID 1) and Microglossum rufum (AFTOL-ID 1292).
The best scoring RAxML tree with a final likelihood value of
- 32124.131818 is presented. The matrix had 1958 distinct alignment
patterns, with 38.37% of undetermined characters or gaps. Estimated
base frequencies were as follows; A = 0.237765, C = 0.258447,
G = 0.293741, T = 0.210047; substitution rates AC = 1.443319,
AG = 2.967467, AT = 1.388359, CG = 1.234302, CT = 7.043503,
GT = 1.000000. MLBS equal to or greater than 70% (first set) and
BYPP values equal to or greater than 0.95 (second set) are given at
the nodes. The ex-type strains are in bold and black. The newly
generated sequence is indicated in bold and blue
Etymology: The name refers to the place that fungus was
collected, Chiang Rai, Thailand.
Holotype: MFLU 18-1722
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies on natural substrate effuse, black, hairy, with long chains of conidia.
Mycelium immersed, composed of hyaline, guttulate,
branched, 1.5–3 wide hyphae. Setae 85–150 9 3.5–8 lm,
scattered or in groups mixed with conidiophores, capitate,
sometimes surrounded by hyaline mucilage at the swollen
apex, smooth, median brown, paler towards the apex,
straight or flexuous, septate. Conidiophores 100–220 lm
long, 5–8 lm wide below venter, 11–13.5 lm wide above,
and 13–16 lm wide at venter, macronematous, mononematous, smooth, dark brown to black, paler at the torn
apex, straight or slightly flexuous, solitary or in groups of
123
162
123
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b Fig. 115 Canalisporium dehongense (MFLU 18-1189, holotype). a,
b Colonies on submerged wood. c–i Conidia with conidiogenous
cells. j, k Conidia. l Conidiophore. m, n Colony on PDA (m from
above view, n from below view). Scale bars: c, d, g, h, l = 20 lm, e,
f, i–k = 10 lm
2–3, arising from dark brown to black bulbous base,
composed of a cylindrical stipe and a swollen venter with a
long cylindrical neck, erect, sometimes proliferating percurrently. Conidiogenous cells monophialidic, percurrent,
integrated, brown, lageniform, frayed at the apex. Conidia
15.5–30 9 8.5–12.5 lm (
x = 23.8 9 10.7 lm, n = 30),
formed in chains, cylindrical, aseptate, hyaline, verruculose, smooth-walled, with a conspicuous guttule at centre
when young, 1-septate, brown, with conspicuous, circular
guttules in each cell when mature, conspicuously darkened
and slightly constricted at the septa, thickened at both ends
(Fig. 117).
Culture characteristics: Conidia germinating on water
agar media within 48 h. One or two germ tubes produced
randomly from conidium wall. Colonies growing on PDA
white, irregular, circular, edge entire, mycelium fluffy,
dense in the centre, becoming sparse at the edge.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Ban Nang Lae Nai, on decaying wood
submerged in a freshwater stream, 6 March 2018, N.G. Liu,
CR066 (MFLU 18-1722, holotype), ex-type living culture
(MFLUCC 18-0703).
GenBank numbers: ITS: MH883032, LSU: MH883030,
SSU: MH883048.
Notes: Sporoschisma chiangraiense is phylogenetically
related to S. aquaticum and S. palauense (Fig. 118).
Comparisons of ITS sequences showed that there are
106 bp differences with gaps of total 558 bp between S.
chiangraiense and S. aquaticum, and 58 bp differences
with gaps of total 563 bp between S. chiangraiense and S.
palauense. Sporoschisma chiangraiense differs from S.
aquaticum and S. palauense in having 1-septate conidia
when mature, while the latter two are 3-septate, and 1–3septate respectively (Luo et al. 2016; Yang et al. 2016a).
Conidia of S. chiangraiense have conspicuous guttules, a
feature absent in S. aquaticum.
Ellisembia Subram., Proc. Indian Natn Sci. Acad., Part B.
Biol. Sci. 58(4): 183 (1992)
Notes: Ellisembia, based on E. coronata, was segregated
from the widely circumscribed Sporidesmium by Subramanian (1992) and delimited to species with brown, distoseptate conidia and conidiophores with none or
percurrent and irregular extension. It comprises saprobic
species on wood, bamboo culm or plant remnants in terrestrial and freshwater habitats. Based on the evidence
from molecular DNA data, the difference between
163
euseptate vs. distoseptate conidia among Sporidesmiumlike species does not seem significant (Su et al. 2016; Yang
et al. 2018). The genus Ellisembia is polyphyletic, it
comprises 65 species but only some of them could be
positioned in the system based on DNA sequence data.
Currently, members of Ellisembia are placed in four clades
in Sordariomycetes, i.e. in Chaetosphaeriaceae under the
name Ellisembia, Distoseptisporaceae as Distoseptispora,
Sporidesmiaceae as Sporidesmium and Xylariales as Ellisembia calyptrata (Réblová and Winka 2001; Shenoy
et al. 2006; Su et al. 2016; this study). Until the systematic
positions of the type species of Ellisembia and Sporidesmium are revealed, we recommend using the name
Ellisembia for species grouped in a strongly supported
monophyletic clade in Chaetosphaeriaceae, experimentally
linked with sexual morphs having multiseptate, versicolorous ascospores in asci with a non-amyloid apical annulus, persistent paraphyses and immersed ascomata.
Ellisembia aurea Réblová & J. Fourn., sp. nov.
MycoBank number: MB827909; Facesoffungi number:
FoF05783; Figs. 119, 120
Etymology: Aureus (L) golden, referring to the colour of
the golden-yellow pigment covering necks and surrounding
substrate.
Holotype: PRA-14744
Saprobic on decaying wood partly submerged in freshwater. Sexual morph Ascomata immersed with protruding
necks or becoming superficial, solitary or in small groups.
Venter 600–750 lm diam., 320–400 lm high, subglobose,
upright or decumbent. Neck central, 170–200 lm wide,
300–600(–1000) lm long, upright or decumbent, cylindrical, sometimes tapering towards the apex, with a distinct
pore at the apex. Neck often basally swollen up to
320–400 lm, conical, surrounded by a dark brown to black
stroma formed of densely interwoven, dark brown, septate,
thick-walled hyphae 2.5–3.5 lm wide of textura intricata.
Stroma covered by erect, dark brown conidiophores. Neck
covered irregularly by a layer of golden-yellow granules
which disappear with age; granules of the same pigment
are also staining the surrounding substrate. Ostiole periphysate. Ascomatal wall leathery, 20–27 lm, two-layered;
outer layer consisting of brown, thick-walled, polyhedral
cells with opaque walls of textura angularis; towards the
interior cells more flattened of textura prismatica. Inner
layer consisting of several rows of thin-walled, hyaline,
flattened cells. Paraphyses hyaline, branched, anastomosing, septate, 2.5–3.5 lm wide, tapering to ca. 2.0 lm. Asci
(165–)170–205 9 23–25.5(–27.5) lm (mean ± SD =
190.4 ± 14.2 9 24.6 ± 1.6 lm),
(150–)154–175 lm
(mean ± SD = 163.1 ± 8.5 lm) long in the sporiferous
part, truncate at the apex, cylindrical to clavate, stipitate;
with 8 obliquely uniseriate or biseriate ascospores; apical
123
164
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Fig. 116 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU and SSU sequence data for Canalisporium species and several related genera. Related sequences are taken
from Sri-indrasutdhi et al. (2010) and Boonyuen and Pang (2011).
Twenty-five strains are included in the combined sequence analyses
which comprise total 2757 characters after aligned. Pleurothecium
semifecundum (CBS 131271) (Pleurotheciaceae, Pleurotheciales) is
used as the outgroup taxon. Tree topology of the maximum likelihood
analysis is similar to the Bayesian analysis. The best scoring RaxML
tree with a final likelihood value of - 11890.841233 is presented.
The matrix had 828 distinct alignment patterns, with 10.21%
undetermined characters or gaps. Estimated base frequencies were
as
follows:
A = 0.225725,
C = 0.259821,
G = 0.307021,
T = 0.207433; substitution rates AC = 1.748411, AG = 3.032505,
AT = 1.758208, CG = 1.095962, CT = 6.405200, GT = 1.000000;
gamma distribution shape parameter a = 0.244920. Bootstrap values
for maximum likelihood (ML) equal to or greater than 70 and
Bayesian posterior probabilities (BYPP) equal to or greater than 0.95
are placed above the branches, respectively. The ex-type strains are in
bold and black. The newly generated sequence is indicated in bold
and blue
annulus 6.5–7.5(–8) lm wide, 2.0–2.5 lm high, nonamyloid. Outer wall of the ascus fragile, often rupturing
basally resulting in a prolonged stipe and apically causing a
release of the apical content with the annulus. Ascospores
(33–)36–38(– 40) 9 9.5–11.5(– 12) lm (mean ± SD =
37.1 ± 2.0 9 10.6 ± 0.6 lm), ellipsoidal, sometimes
inequilateral, 5(–7)-septate, not constricted at the septa,
with a large globule in each cell obscuring the septa, versicolorous prior to discharge, middle cells brown to olivaceous brown with hyaline polar cells, smooth-walled,
without mucilaginous sheath or appendages. Asexual
morph Conidiophores macronematous, mononematous,
92–155 lm long, 6.5–7.5 lm wide, unbranched, upright or
slightly bent, seldom with a percurrent extension, septate,
dark brown, paler and tapering towards the tip, (3.5–
)5.0–5.5 lm wide at the apex, truncate. Conidia holoblastic, secession schizolytic, (65–)70–97(–103) 9 11–16(–
17.5) lm (mean ± SD = 85.0 ± 10.2 9 14.8 ± 2.3 lm),
(3.5–)4.5–5.0 lm wide at the base, obclavate to fusiform to
lanceolate, truncate at the base, tapering apically with a
terminal extension which might be up to 26 lm long,
brown to reddish brown, with the darkest basal cell, paler at
the apex, 11–13(–15)-distoseptate, smooth-walled.
Culture characteristics: Colonies on Modified Leoniańs
Agar (MLA) 7–8 mm diam. after 28 d, convex, circular to
slightly irregular. Aerial mycelium abundant, lanose, floccose, partially decumbent, colony yellow, surrounded by a
yellow-orange pigment diffused into the agar; margin
entire to slightly undulate; reverse yellow-orange. Sporulation after 60 d; conidiophores arise at the margin of the
colony. Colonies on potato-carrot agar (PCA) 12–13 mm
diam. after 28 d, convex, circular. Aerial mycelium abundant around the centre and on the inoculation block, velvety, loose to almost cobwebby towards the margin, colony
yellow-orange, surrounded by a yellow zone of submerged
growth and deep yellow pigment diffused into the agar;
margin filiform; reverse yellow. Sporulation absent.
Vegetative mycelium hyaline, some hyphae pale brown,
1.5–2.5 lm wide. Conidiophores 24–40 lm long, 6–10 lm
wide in the broadest part, seldom macronematous, mostly
semimacronematous reduced to a conidiogenous cell
sometimes with 1–2 supporting cells, conidiogenous cell
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Fig. 117 Sporoschisma chiangraiense (MFLU 18-1722, holotype).
a, b Colonies on natural substrates. c, d Conidia and conidiophores
with setae. e Conidiophores and conidia. f Portion of phialide
165
producing conidium. g–i Conidia. j Germinated conidium. Scale bars:
a = 200 lm, b = 100 lm, c–e = 50 lm, f–h = 25 lm, i, j = 20 lm
123
166
Fungal Diversity (2019) 96:1–242
Fig. 118 Phylogram generated from maximum likelihood analysis
based on combined LSU and ITS sequence data representing
Chaetosphaeriaceae. Related sequences are taken from Liu et al.
(2016), Lu et al. (2016), Luo et al. (2016) and Yang et al. (2016a).
Thirty-five strains are included in the combined analyses which
comprise 1635 characters (1028 characters for LSU, 607 characters
for ITS) after alignment. Lasiosphaeria ovina (SMH4605) (Lasiosphaeriaceae, Sordariales) is used as the outgroup taxon. Single
gene analyses are carried out and the topology of each tree had clade
stability. Tree topology of the maximum likelihood analysis is similar
to the maximum parsimony analysis and the Bayesian analysis. The
best RaxML tree with a final likelihood value of - 8025.509056 is
presented. The matrix had 607 distinct alignment patterns, with
24.38% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.228162, C = 0.260607, G = 0.308790,
T = 0.202441; substitution rates AC = 1.613577, AG = 2.168568,
AT = 1.347260, CG = 0.833554, CT = 8.019859, GT = 1.000000;
gamma distribution shape parameter a = 0.534284. Maximum parsimony analysis of 1148 constant characters and 117 informative
characters resulted in nine equally most parsimonious trees (CI =
0.591, RI = 0.751, RC = 0.443, HI = 0.409). Bootstrap values for
maximum likelihood (ML) and maximum parsimony (MP) equal to or
greater than 75 are placed above and below the branches respectively.
Branches with Bayesian posterior probabilities (BYPP) equal or
greater than 0.95 are in bold. The ex-type strains are in bold and
black. The newly generated sequence is indicated in bold and blue
swelled in the middle, dark brown, with a cup-shaped
percurrent
extension.
Conidia
(95–)105–128(–
137) 9 11.5–13.5(–16.5) lm (mean ± SD = 124.0 ±
11.8 9 12.9 ± 1.5 lm), 3.5–4.5(–5) lm wide at the base,
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Fungal Diversity (2019) 96:1–242
long-fusiform, truncate at the base, tapering apically,
brown, with the darkest cell at the base, paler towards the
apex, (12–)15–18-distoseptate, smooth-walled, apex
sometimes extended into a hyaline filiform appendage
56–85 lm long, 2.0–2.5 lm wide, conidia with an appendage 186–230(–268) lm long (mean ± SD = 221 ± 36.5).
Material examined: FRANCE, Ariège, Rimont, La
Maille brook, 42.97706, 1.307167, ca. 550 m a.s.l., on
partly submerged wood of a decorticated branch of Sambucus nigra, 4 August 2017, J. Fournier, J.F. 17058 (PRA14744, holotype), ex-type living culture (CBS 144403).
GenBank numbers: ITS: MH836375, LSU: MH836376.
Notes: Conidia that are formed on nature substrate are
shorter, usually fusiform to lanceolate, tapering apically
with a short terminal extension and with less distosepta
compared to conidia formed in MLA culture, which are
mostly long-fusiform, wider at the distal end, which
sometimes extends into a hyaline filiform appendage. The
conidiophores formed in vivo are macronematous and
cylindrical compared to usually semimacronematous and
swollen conidiophores with a frequent percurrent extension
formed in vitro. In the ITS-LSU phylogeny (Fig. 121), E.
aurea is positioned in Chaetosphaeriaceae in a strongly
supported clade together with Ellisembia brachypus, E.
foliiculata and Pyrigemmula aurantiaca.
Besides E. foliiculata (Réblová and Winka 2001), E.
aurea is another Ellisembia species experimentally linked
with a sexual morph with versicolorous ascospores. Ellisembia aurea resembles E. foliiculata in the morphology
of immersed ascomata, size of ascospores and morphology
of brown, distoseptate conidia but the latter species differs
from E. aurea in having flask-shaped ascomata, regularly
7-septate ascospores, shorter and narrower asci and cylindrical to clavate or obclavate, shorter conidia without a
rostrum or appendage at the distal end. Hughes and Illman
(1974a) reported the characteristic yellowish colour of the
substrate surrounding the colonies of E. foliiculata, similar
to the golden-yellow pigment released in vitro and found in
wood surrounding ascomata of E. aurea. Among Ellisembia species having conidia with a distal rostrate end, E.
aurea resembles E. brachypus, which differs in having
somewhat shorter conidia (as E. deightonii 50–90 lm fide
Ellis 1958, 45–65 lm fide Hughes and Illman 1974b) with
fewer, 5–8 distosepta. The conspecificity of E. aurea and
E. brachypus (HKUCC 10555, Shenoy et al. 2006) is not
supported in our phylogeny.
Subramanian (1992) segregated Ellisembia, typified by
E. coronata, from the widely circumscribed Sporidesmium
to accommodate species with distoseptate conidia and
conidiophores with none or percurrent and irregular
extension. However, such delimitation based purely on
asexual morphological characters is not supported by
molecular DNA data (Su et al. 2016; Yang et al. 2018b). Su
167
et al. (2016) resurrected Sporidesmiaceae (Fries 1849) with
the single genus Sporidesmium to accommodate taxa with
euseptate or distoseptate conidia and accepted Ellisembia
as its synonym. However, the type species of neither
Sporidesmium nor Ellisembia were included. In the same
publication, a new family Distoseptisporaceae and the
genus Distoseptispora were introduced (Su et al. 2016),
delimiting a group of Sporidesmium-like taxa in having
exclusively distoseptate conidia, and two former Ellisembia species were included. However, Yang et al. (2018b)
expanded the generic concept of Distoseptispora by
including species with euseptate conidia. At present, species formerly classified in Ellisembia are accommodated in
four different clades in the Sordariomycetes, i.e. E.
adscendens and E. leonensis in Distoseptispora in Distoseptisporaceae, E. bambusicola and E. minigelatinosa in
Sporidesmium in Sporidesmiaceae (Shenoy et al. 2006; Su
et al. 2016), E. aurea, E. foliiculata and E. brachypus in
Chaetosphaeriaceae (Réblová and Winka 2001; Shenoy
et al. 2006; this study) and E. calyptrata in Xylariales
(Shenoy et al. 2006). Moreover, various sexual morphs
have been linked with Ellisembia. Those with hyaline,
septate ascospores and superficial or immersed ascomata
are currently placed in Distoseptisporaceae, i.e. D.
adscendens with Miyoshiella triseptata sexual morph (as
Lasiosphaeria, Shoemaker & White 1985), and in
Sporidesmiaceae, i.e. S. bambusicola with Miyoshiella
fusispora sexual morph (Kawamura 1929) and Sporidesmium thailandense of which only the sexual morph is
known (Yang et al. 2018b). The positon of Sporidesmium
larvatum with Miyoshiella larvata sexual morph (Reblova
1999) is unknown. Furthermore, Ellisembia sexual morphs
with versicolorous ascospores and immersed ascomata
were so far exclusively placed in Chaetosphaeriaceae.
Despite the proposed synonymy of Ellisembia with Sporidesmium (Su et al. 2016) and also with similar Imicles
(Shoemaker and Hambleton 2001; Wu and Zhuang 2005),
in the absence of molecular DNA data of their type species,
their relationship is unclear and these genera should be
retained as separate taxa following Seifert et al. (2011).
For the time being, we use the name Ellisembia for taxa
attributed to Chaetosphaeriaceae and characterized by
pigmented, distoseptate conidia, macronematous conidiophores with or without percurrent extension and multiseptate, versicolorous ascospores in asci with a nonamyloid apical annulus, persistent paraphyses and
immersed ascomata. Members of the Ellisembia clade in
Chaetosphaeriaceae are congeneric with the morphologically similar Pyrigemmula typified by P. aurantiaca
(Magyar et al. 2011). The generic name Pyrigemmula is
available and recommended for use to accommodate Ellisembia p. p. and related fungi if its separate position from
E. coronata is confirmed in the future.
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b Fig. 119 Ellisembia aurea (PRA-14744, holotype). a–c Ascomata.
d Vertical section of ascoma (white arrow indicates stroma, black
arrow indicates ascomal wall). e Vertical section of the lower part of
the neck with visible stroma. f Stroma consisting of interwoven
hyphae in detail g Vertical section of the ascomatal wall. h Ascal apex
with apical annulus (arrows indicate ruptured outer wall and released
ascal content). i, j Asci. k Ascospore. l Paraphyses. m Ascus with
immature ascospores. n, o Conidia and conidiophores on nature
substrate. Scale bars: a–c = 200 lm, d = 100 lm, e = 50 lm,
f = 20 lm, g–o = 10 lm
Linocarpaceae Konta & K.D. Hyde
Notes: Konta et al. (2017) introduced this family to
accommodate Linocarpon Syd. & P. Syd. and Neolinocarpon K.D. Hyde and placed it in Chaetosphaeriales.
Senwanna et al. (2018) introduced the novel species, Neolinocarpon phayaoense Senwanna & K.D. Hyde.
Neolinocarpon K.D. Hyde
Notes: Neolinocarpon was included in Linocarpaceae
based on DNA sequence data (Konta et al. 2017) and this
was confirmed by Senwanna et al. (2018).
Neolinocarpon rachidis Konta & K.D. Hyde
Facesoffungi number: FoF05114; Fig. 122
Saprobic on petiole of Cocos nucifera L. Sexual morph
Ascomata 320–460 lm high 9 230–290 lm diameter
(
x = 415 9 265 lm, n = 10), solitary, deeply immersed,
with a central raised, black, globose-subglobose papilla,
with a central ostiole. Papilla 157–223 lm high 9 127–
198 lm diameter at the base (
x = 160 9 140 lm, n = 5),
black, shiny, with hyaline periphyses. Peridium 14–30 lm
wide (
x = 22 lm, n = 10), outer cells merging with the
host epidermal cells, composed of dark brown to black
cells of textura angularis. Hamathecium comprising
numerous, 1–3 lm diameter (
x = 2 lm, n = 20), hyphalike, septate, unbranched, paraphyses, longer than asci.
Asci 95–180 9 9–17 lm (
x = 145 9 13 lm, n = 20), 8spored, unitunicate, cylindrical, long pedicellate, with a
wedge-shaped, J-, subapical ring. Ascospores 123–
140 9 2–4 lm (
x = 106 9 3 lm, n = 20), parallel in
ascus, becoming spiral when mature, filiform, straight or
curved, hyaline, aseptate, containing numerous refringent
septum-like bands, rounded at the apex, pointed at the base,
lacking appendage sheath, smooth-walled. Appressoria 3–
5 9 2–5 lm (
x = 4 9 3.5 lm, n = 20), globose to subglobose, hyaline. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 24 h and produced appressoria-like structures
at each tips of germ tube. Colonies on MEA reaching
2.5–3 cm diameter after two weeks at 25–28 C, white at
the edge, brown in the middle with strong radiations outwards. After 3 weeks of incubation, colonies smooth, flat,
margin undulate, white to gray in the centre, white at the
169
margin, mycelium becoming red-brown when they produced into media, hyphae septate, branched, smoothwalled.
Material examined: THAILAND, Prachaupkhirikhan
Province, on dead of petiole, Cocos nucifera (Arecaceae),
30 July 2015, Sirinapa Konta, PJK04i (MFLU 15-2347,
isotype), ex-isotype living culture (MFLUCC 15-0814).
GenBank numbers: ITS: MK106342, LSU: MK106353,
MK106354, SSU: MK106367, MK106368.
Notes: Neolinocarpon rachidis was introduced as a new
species by Konta et al. (2017) and in this study N. rachidis
was found on Cocos nucifera (Arecaceae) in Prachaupkhirikhan Province, Thailand. Appressorium structures
were produced by germ tubes indicating that are endophytes. Phylogenetically, N. rachidis strain MFLUCC
15-0814 clustered with N. rachidis MFLUCC 15-0332 with
high support (Fig. 123). Therefore, a record of N. rachidis
on coconut host is reported.
Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Notes: Eriksson and Hawksworth (1986) introduced the
order Amphisphaeriales. However, it was synonymized
with Xylariales by Eriksson and Hawksworth (1987).
Based on molecular and morphological data, these orders
were separated and Amphisphaeriales was resurrected
(Senanayake et al. 2015). Currently, this order includes six
families, namely: Amphisphaeriaceae, Clypeosphaeriaceae, Bartaliniaceae, Discosiaceae, Phlogicylindriaceae
and Pestalotiopsidaceae (Senanayake et al. 2015).
Amphisphaeriaceae G. Winter
Notes: Winter (1884–1886) introduced the family
Amphisphaeriaceae to accommodate the type genus Amphisphaeria. For an account of Amphisphaeriaceae G.
Winter see Senanayake et al. (2015). In this manuscript we
introduce a new monotypic genus Trochilispora.
Trochilispora VP Abreu, AWC Rosado & OL Pereira, gen.
nov.
Index Fungorum number: IF555484; Facesoffungi
number: FoF04859
Etymology: From the family Trochilidae (hummingbirds) and –spora.
Type species: Trochilispora schefflerae Abreu VP,
Rosado AWC & Pereira OL
Asexual morph Associated with leaf spot disease on
Schefflera morototoni (Fig. 125a). Conidiomata acervular
(Fig. 127c), epiphyllous, scattered and occasionally confluent, subepidermal in origin, erumpent, rounded to oval in
outline, unilocular, brown or black, basal stroma thick, of
textura angularis, cells thick-walled and almost colourless;
lateral walls thick, cells thick-walled, pale brown to brown.
Conidiophores cylindrical to subcylindrical, formed in the
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170
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b Fig. 120 Ellisembia aurea (CBS 144403, ex-holotype). a Conidium
with appendage. b, c Conidia. d, e Conidiophores. f, g Conidiophores
with visible percurrent extension. h Conidia formed on aerial
mycelium. i, j Colonies on MLA and PCA after 28 d. Scale bars:
a–g = 10 lm, h = 100 lm, i, j = 0.5 cm
concavity of the conidioma, unbranched, hyaline, smoothwalled. Conidiogenous cells discrete, annellidic with 2
annellations, hyaline, thin- and smooth-walled. Conidia
fusiform, straight or slightly curved, 3–4-septate, concolourous, smooth, bearing apical appendage tubular, filiform, single, not centric, unbranched and basal appendage
absent; basal cell hyaline to subhyaline, obconic to conic,
smooth and thin-walled; 2–3 median cells doliiform,
smooth, concolourous, brown, septa darker than the rest of
the cell; apical cell hyaline to subhyaline, subconical to
hemisphaerical, thin- and smooth-walled. Sexual morph
Undetermined.
Fig. 121 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data for Ellisembia aurea
and members of Chaetosphaeriaceae. Related sequences are taken
from Réblová and Winka (2000, 2001), Réblová (2004), Réblová and
Seifert (2008), Schoch et al. (2009), Magyar et al. (2011), Réblová
et al. (2011), Yang et al. (2016a) and Vu et al. (2019). Seventeen
strains are included in the combined gene sequence analysis, which
comprises total 1837 characters including gaps (628 for ITS, 1209 for
LSU) and 528 unique characters. Cylindrotrichum clavatum (CBS
125296) and C. oligospermum (CBS 101319) are used as the outgroup
taxa. The best RAxML tree with a final likelihood value of
171
Trochilispora schefflerae VP Abreu, AWC Rosado & OL
Pereira, sp. nov.
Index Fungorum number: IF555485; Facesoffungi
number: FoF04860; Fig. 124
Etymology: Name derived from its host genus,
Schefflera.
Holotype: VIC 44384
Asexual morph Conidiomata acervular, epiphyllous,
scattered and occasionally confluent, subepidermal in origin, erumpent, rounded to oval in outline,
49–88 9 79–235 lm diam., unilocular, brown or black,
basal stroma thick, of textura angularis, cells thick-walled
and almost colourless; lateral walls 3–5 cells thick, cells
thick-walled, pale brown to brown. Conidiophores cylindrical to subcylindrical, (8.5–15.5 9 1.5–2 lm), formed in
the concavity of the conidioma, unbranched, hyaline,
smooth-walled. Conidiogenous cells discrete, annellidic
with 2 annellations, (3.5–11.5 9 1.5–3 lm), hyaline, thinand smooth-walled. Conidia fusiform, straight or slightly
curved, concolourous, smooth, bearing apical appendage,
and basal appendage absent; 3-septate (13–19 9 3.5–5),
bearing: [basal cell obconic to conic, hyaline to subhyaline,
- 7216.460156 is presented. Estimated base frequencies for the two
partitions were as follows: the ITS partition A = 0.214700,
C = 0.306167, G = 0.262251, T = 0.216881; substitution rates AC =
4.616604, AG = 5.194154, AT = 5.725980, CG = 3.570615, CT =
13.113689, GT = 1.000000; the LSU partition A = 0.242254,
C = 0.240274, G = 0.317255, T = 0.200218; substitution rates AC =
0.504038, AG = 1.058491, AT = 0.427775, CG = 0.207895, CT =
3.855638, GT = 1.000000. Branch support in nodes C 50% maximum likelihood bootstrap support is indicated above branches. The
ex-type strains are in bold and black. The newly generated sequence
is indicated in bold and blue
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Fig. 122 Neolinocarpon rachidis (MFLU 15-2347, isotype). a Appearance of ascomata on host substrate. b, c Close up of ascomata.
d Yellowish ascospore mass. e Section of ascoma. f Papilla.
g Peridium. h Paraphyses. i–k Asci. l J- reaction of apical ring. m–
p Ascospores. q Germinated ascospore (appressoria at red arrow). u–
y Appressoria. Scale bars: a = 1000 lm, b, d = 500 lm, c = 200 lm,
e = 100 lm, f, i–s = 50 lm, g = 20 lm, h = 10 lm, u–y = 5 lm
smooth and thin-walled, 2–4 lm long; two median cells
doliiform, 8.5–12.5 lm long, smooth, concolourous,
brown, septa darker than the rest of the cell (second cell
from base brown, 3.5–5.5 lm long; third cell brown,
4.5–7 lm long); apical cell 2–3.5 lm long, hyaline to
subhyaline, subconical to hemispherical, thin- and smoothwalled; with 1 tubular apical appendage, arising from the
apical crest, not centric, unbranched, filiform, 2–6.5 lm
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long;
basal
appendage
absent],
or
4-septate
(15.5–21 9 4–5), bearing: [basal cell obconic to conic,
hyaline to subhyaline, smooth and thin-walled, 2–5 lm
long; three median cells doliiform, 10–13 lm long,
smooth, concolourous, brown, septa darker than the rest of
the cell (second cell from base brown, 4–6 lm long; third
cell brown, 2.5–4 lm long; fourth cell brown, 2.5–4.5 lm
long); apical cell 2.5–3.5 lm long, hyaline to subhyaline,
subconical to hemisphaerical, thin- and smooth-walled;
with 1 tubular apical appendage, arising from the apical
crest, not centric, unbranched, filiform, 2.5–7.5 lm long;
basal appendage absent. Sexual morph Undetermined.
Culture characteristics: Colonies cultured on PDA
reaching 38 mm diam. after 1 wk at 25 C with a photoperiod of 12 h; regularly margins; with dense aerial
mycelium; white; colonies fertile (Fig. 124b). Colonies
cultured on MEA reaching 40 mm diam. after 1 wk at
25 C with a photoperiod of 12 h; regularly and submerged
margins; with scarce and sebaceous aerial mycelium; pale
yellowish; colonies fertile.
Material examined: BRAZIL, Minas Gerais, Paraopeba,
Floresta Nacional de Paraopeba (FLONA-Paraopeba), on
leaves of Schefflera morototoni (Araliaceae), 30 January
2016, V.P. Abreu & O.L. Pereira (VIC 44384, holotype),
ex-type living culture (COAD 2371).
GenBank numbers: ITS: MH128360, LSU: MH084761,
TEF1-a: MH231216, TUB2: MH231215.
Notes: Trochilispora is introduced as a new genus based
on morphology and phylogenetic support (LSU and ITS
sequence data). Based on phylogenetic analyses, Trochilispora schefflerae COAD 2371 grouped in a well-supported clade including Hymenopleella hippophaeicola CBS
140410 (Fig. 125), but different genera can be grouped in
the same clade as for example, Morinia and Zetiasplozna;
Truncatella and Broomella; among others. Unfortunately,
Jaklitsch et al. (2016) did not observed the asexual morph
of Hymenopleella hippophaeicola, but the authors cite the
Hymenopleella sollmannii species reported by Shoemaker
and Müller (1965). The phylogenetic position of the Trochilispora family is still unclear. Trochilispora schefflerae
COAD 2371 differs from Hymenopleella sollmannii by
having conidia formed in conidiomata acervular with lateral walls 3–5 cells thick of brown hyphae; conidiophores
smaller; conidiogenous cells discrete, annellidic with 2
annellations; conidia fusiform, straight or slightly curved,
3–4-septate, with medium brown central cells and hyaline
to subhyaline end cells, apical cell with an appendage
tubular, filiform, single, not centric, unbranched, not septum and basal cell without appendage basal. Our phylogenetic tree built using LSU and ITS data, and
morphological features corroborated that our isolate represents a new genus and a new species belonging to
Amphisphaeriaceae (Fig. 125).
173
Sporocadaceae Corda.
Notes: Jaklitsch et al. (2016) proposed Sporocadaceae
based on morphological observations and phylogenetic
analyses of ITS-LSU sequence data. This group of fungi
comprises different asexual morph genera which are
acervular coelomycetes having hyaline, pale or dark
brown, septate conidia. The type genus is Seimatosporium
Corda (Jaklitsch et al. 2016). Jaklitsch et al. (2016)
accepted Bartalinia, Monochaetia, Neopestalotiopsis,
Pseudopestalotiopsis, Pestalotiopsis, Seiridium and many
other conidial appendage bearing asexual morph genera
(Jaklitsch et al. 2016; Wijayawardene et al. 2017a, 2018a).
Wijayawardene et al. (2018a) listed 22 genera in this
family.
Pestalotiopsis Steyaert
Notes: Based on conidial features, Steyaert (1949)
divided Pestalotia De Not., into three genera, Pestalotia De
Not., Pestalotiopsis Steyaert, and Truncatella Steyaert.
Species with 5-celled conidia (4-septate) was grouped
within Pestalotiopsis. Based on a multilocus phylogenetic
and morphological analysis of pestalotiopsis-like species,
Maharachchikumbura et al. (2014) divided the complex
into three genera: Pestalotiopsis Steyaert, Neopestalotiopsis Maharachch., K.D. Hyde & Crous and Pseudopestalotiopsis Maharachch., K.D. Hyde & Crous based on
morphology of conidial median cells and LSU sequence
data. 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 analysis based on the
combination of three gene regions (ITS, TUB2 and TEF1a) (Jeewon et al. 2003; Maharachchikumbura et al.
2011, 2012, 2013; Geng et al. 2013).
Pestalotiopsis aggestorum F. Liu & L. Cai, Nature Scientific Reports 7: 870 (2017)
Facesoffungi number: FoF05991; Fig. 126
Saprobic on dead leaves of unknown host. Sexual
morph Undetermined. Asexual morph Conidiomata
acervuli, 130–150 lm diam., pycnidial, superficial to semiimmersed, visible as black spots on the host. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2–4 9 1–2 lm (
x = 2.4 9 1.3 lm), phialidic, discrete, cylindrical, hyaline. Conidia 17–22 9 7–9 lm
(
x = 21.0 9 8.2 lm), fusiform, tapering at both ends, 4septate; basal cell 3–5 lm long (
x = 4.1 lm), conic, hyaline and smooth-walled; three median cells together 11–
15 lm long (
x = 13.7 lm), doliiform, light brown, roughwalled; second cell from base 3–5 lm long (
x = 4.4 lm);
third cell 3–5 lm long (
x = 4.6 lm); fourth cell 3–5 lm
long (
x = 4.0 lm); apical cell 3.2–4.7 lm long
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Fig. 123 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU, SSU and TEF1-a sequence data for
Neolinocarpon species and several closely related genera in
Linocarpaceae and all families in Chaetosphaeriales. Related
sequences were referred from Konta et al. (2017) and Senwanna
et al. (2018). Twenty-eight strains are included in the combined
analyses which comprise 3673 characters (666 characters for ITS,
1005 characters for LSU, 1043 characters for SSU, 946 characters for
TEF1-a) after alignment. Sordaria fimicola and Gelasinospora
tetrasperma (AFTOL-ID 1287) (Sordariaceae, Sordariales) are used
as the outgroup taxa. Single gene analyses are carried out and the
topology of each tree had clade stability. Tree topology of the
maximum likelihood analysis is similar to the maximum parsimony
analysis and the Bayesian analysis. The best RaxML tree with a final
likelihood value of - 14985.889953 is presented. The matrix had
1082 distinct alignment patterns, with 55.45% undetermined characters or gaps. Estimated base frequencies were as follows:
A = 0.231817, C = 0.269309, G = 0.304861, T = 0.194013; substitution rates AC = 0.889100, AG = 1.588862, AT = 0.616066, CG =
1.035174, CT = 4.437165, GT = 1.000000; gamma distribution
shape parameter a = 0.281924. Maximum parsimony analysis of
3673 constant characters and 649 informative characters resulted in
two equally most parsimonious tree of 2562 steps (CI = 0.702,
RI = 0.690, RC = 0.484, HI = 0.298). Bootstrap values for maximum
likelihood (ML) and maximum parsimony (MP) equal to or greater
than 50, Bayesian posterior probabilities (BYPP) equal to or greater
than 0.9 are placed above the branches respectively. The ex-type
strains are in bold and black. The newly generated sequences are
indicated in bold and blue
(
x = 4.0 lm), conic, hyaline and smooth-walled; 2–3 apical appendages 9–17 lm long (
x = 15 lm), tubular, filiform; basal appendage 5–8 lm long (
x = 6.2 lm), single,
central, tubular, filiform.
Culture characteristics: Colonies on PDA 35 mm
diameter after 7 days at 25 C, circular, raised, dense aerial
mycelia with fluffy appearance with circular edge, white;
reverse white.
Material examined: THAILAND, Chiang Rai Province,
Mae Fah Luang District, Doi Mae Salong, on dead leaves
of unknown host, 23 September 2016, N. I de Silva, NI104
(MFLU 17-0674), living culture (MFLUCC 17-0869).
GenBank numbers: ITS: MK334672.
Notes: Our strain clusters with the type of Pestalotiopsis
aggestorum in the combined ITS, TUB2 and TEF1-a
Pestalotiopsis
phylogenetic
analyses
(Fig. 127).
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175
Fig. 124 Trochilispora schefflerae (VIC 44384, holotype). a Leaves
of Schefflera morototoni in Floresta Nacional de Paraopeba, state of
Minas Gerais, Brazil (the arrows indicate the reproductive structures
of the fungus). b Colony on PDA after 1 wk at 25 C with a
photoperiod of 12 h in the dark in Petri dishes (90 9 15 mm) (COAD
2371). c Cross section of the conidioma. d–g Conidia. Scale bars:
c = 50 lm, d–g = 10 lm
aggestorum was isolated from leaves of tea (Camellia
sinensis) in Yunnan, China. Our strain has fusiform,
4-sepatate conidia with overlap dimensions. However, our
strain has smaller apical appendages (9–17) lm than P.
aggestorum (18–28 lm; Liu et al. 2017). The current collection of P. aggestorum was isolated from dead leaves of
unknown forest plant in Chiang Mai, as a new collection
from Thailand.
Xylariales Nannf.
Castanediellaceae Hern.-Restr., Guarro & Crous
Notes: The phylogenetic position based on analysis of
LSU sequence data shows it belongs to Xylariales and is
distinct from Beltraniaceae. The typical asexual morph
features are: conidiophores macronematous, mononematous or aggregated in sporodochia, branched, brown to pale
brown. Conidiogenous cells mono or polyblastic, sympodial, discrete, solitary or in whorls, cylindrical to lageniform, hyaline to subhyaline. Conidia are unicellular or
septate transversely, cylindrical, fusiform or lunate, and are
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b Fig. 125 Phylogram generated from Bayesian Inference analysis
based on combined ITS and LSU sequence data for several closely
related genera in Amphisphaeriales. Sequence data of ex-type or exepitype cultures are taken from Senanayake et al. (2015) and the
closest hits of GenBank database were included in this study. The
combined genes sequence analysis included 40 taxa, which comprise
1405 characters (591 characters for ITS, 814 characters for LSU), and
outgroup taxon Immersidiscosia eucalypti (MAFF 242781). Bayesian
posterior probability are indicated at the nodes, and values equal or
greater than 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
hyaline. In Castanediellaceae there is single genus Castanediella (Wanasinghe et al. 2018a).
Castanediella Hern.-Restr., Crous & M.J. Wingf.
Notes: The genus Castanediella was introduced by
Hernández-Restrepo et al. (2015), with Castanediella
acaciae as type species. Species of Castenediella are differentiated on conidiophores and conidia features. Species
belonging to this genus are characterized by branched
conidiophores and falcate, cylindrical or fusiform, straight
to slightly curved, 0–1-septate conidia. Commonly, species
are isolated from wood or leaves and 12 species are currently accepted (Costa et al. 2018; Wanasinghe et al.
2018a).
While evaluating the diversity of fungi related to Melipona scutellaris substrates (honey, pollen, surface of nest)
living Atlantic Forest, Pernambuco, Brazil, a new species
was isolated and it is described based on morphological
characteristics and its phylogeny is inferred based on ITS/
LSU sequence data.
Castanediella meliponae J.E. Felinto-Santos, R.N. Barbosa
& N.T. Oliveira, sp. nov.
Index Fungorum number: IF555426; Facesoffungi
number: FoF05765; Fig. 128
Etymology: ‘‘meliponae’’ referring to substrate related
the Melipona scutellaris.
Holotype: URM 91974
Sexual morph Undetermined. Asexual morph Colonies
mycelium white, velvety, margin regular, slowly growth,
good sporulation, on PDA (potato dextrose agar), MEA
(malt extract agar) and OA (Oats Meal Agar) at 25 8C in
fourteen days. Mycelium hyaline, septate, smooth, 2–5 lm
diam. Conidiophores hyaline, erect, 35–100 9 2–3.5 lm
diam., 1-2 septate, branched apex, Conidiogenous cells
hyaline,
cylindrical,
defined
shape,
smooth
2–3 9 6–30 lm diam. Conidia hyaline, solitary, one celled, falcate, straight to slightly curved with subobtuse apex,
smooth, 3–4 9 5–10 lm diam.
Culture characteristic: Colinies growing on MEA, surface white, reverse brown with white margins, 29–31 lm
177
diam. On OA surface brown centre and white margin,
reverse in brown centre and white margin, 30–35 lm diam.
On PDA centre of the dark beige colony and white to light
beige margin, reverse with white centre and beige light
margin, 20–23 lm diam. Exudates and soluble pigment
absent in all media. Cultures incubation at 25 8C in 14 days
in incubator type Biological Oxygen Demand.
Material examined: BRAZIL, Paulista, Pernambuco
state, on surface of nest of Melipona scutellaris, native
stingless bees from to Brazilian Atlantic Forest, 29
November 2017, J.E. Felinto-Santos, FIBIO 002-B3 (URM
91974, holotype), ex-type living culture (URM 7933).
GenBank numbers: ITS: MH992664, LSU: MH988746.
Notes: Castanediella meliponae distinguishes of C.
cagnizarii, C. hyalopenicillata and C. eucalypti with
respect to conidiophores, conidiogenous cells and conidia
morphology. The conidiophores of C. cagnizarii and C.
eucalypti are pale brown to brown medium while C.
meliponae are predominantly hyaline. Castanediella
hyalopenicillata has conidiophores prostrate, while C.
meliponae is erect. The conidiogenous cells of C. meliponae distinguish of C. hyalopenicillata, C. cagnizarii, and C.
eucalypti in form, absence of denticles in apex, colour and
absent of scars. C. hyalopenicillata has conidia fusiform
with 0-1 septate. Conidia of C. cagnizarii are cylindrical to
filiform with 0-1 septate. Conidia of C. meliponae are
falcate and one celled. Furthermore, the conidia of C.
meliponae are shorter than C. cagnizarii, C. hyalopenicillata and C. eucalypti. In our phylogenetic analyses
(Fig. 129) C. meliponae is posicionated close to Synnemadiella eucalypti, however some morphological diferences were observed between those. Synnemadiella
eucalypti produces synnemata and conidia granular, ellipsoid, but inequilateral, base truncate, 0.5 lm diam. These
features are not observated in C. meliponae (Fig. 128) and
take together with ITS and LSU sequences we consider as
new species.
Diatrypaceae Nitschke
Notes: Taxa in Diatrypaceae can be distinguished by
perithecial ascomata, usually embedded in a black stroma,
cylindric-clavate to clavate, long pedicellate asci and
allantoid ascospores (Glawe and Rogers 1984; Rappaz
1987; Mehrabi et al. 2015; de Almeida et al. 2016). Both
coelomycetes (Libertella, Cytosporina) and hyphomycetes
(Phaeoisaria) have been reported as asexual morphs of
Diatrypaceae (Glawe and Rogers 1984; Wijayawardene
et al. 2017a,b; de Almeida et al. 2016; Mehrabi et al. 2016).
Currently this family comprises 18 genera (Wijayawardene
et al. 2018a; Dayarathne et al. 2019b).
Diatrypella (Ces. & De Not.) De Not.
Notes: We follow the latest treatment and updated
accounts of Diatrypella in Senwanna et al. (2017).
123
178
Fig. 126 Pestalotiopsis aggestorum (MFLU 17-0674, new geographical record). a The specimen. b, c Conidiomata on the host. d–f
Section of conidiomata. g Conidiogenous cells. h–j Conidia.
Key to the related species of Diatrypella
1. Ascospores 4–5 lm long on average……………
……………………………………Diatrypella major
1. Ascospores 6–10 lm long on average………………2
2. Entorostroma yellowish or olive-green……………..3
2. Entorostroma white………………………………….4
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k Germinating conidia. l, m Colony on PDA (l from above view,
m from below view). Scale bars: d, e = 50 lm, f, h = 20 lm, i–
k = 10 lm, g = 5 lm
3. Asci larger, 120–150 9 15.5–21.5 lm………………
…………………………………………...D. tectonae
3. Asci smaller, 40 9 8–12 lm………………D. frostii
4. Stromata small, up to 2 mm diam.…………………5
4. Stromata larger than 2 mm………………D. vulgaris
5. 1–4 ascomata per stromata, on twigs of Hevea
brasiliensis……………………………..…D. heveae
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5. 3–4 ascomata per stromata, on seed pods of Delonix
regia……………………………………..D. delonicis
Diatrypella delonicis R.H. Perera & K.D. Hyde, sp. nov.
Index Fungorum number: IF552536; Facesoffungi
number: FOF02658; Fig. 130
Etymology: Named after the host genus.
Holotype: MFLU 16-1016
Saprobic on dried seed pods of Delonix regia. Sexual
morph Stromata 0.65–1.28 mm wide, solitary or gregarious on host, immersed, with 3–4 ascomata immersed in a
single stroma, comprising of outer, dark brown to black,
small, tightly packed, thin parenchymatous cell layer, with
inner white, loosely packed, parenchymatous cells. Ascomata
243–340 lm
high,
217–380 lm
diam.,
(
x = 310 9 331 lm, n = 20), perithecial, immersed in
stromatic tissues, aggregated, globose to subglobose,
sometimes with flattened base, pale to dark brown, carbonaceous, ostioles not prominent. Peridium 16–24 lm
wide (
x = 18 lm, n = 20), 2 layered, inner 3–6 layers of
hyaline, flattened, elongate, thin-walled cells of textura
angularis, outer 4–7 layers of pale to dark brown, flattened,
thick-walled cells of textura angularis. Hamathecium
comprising 8.2–13 lm wide (
x = 9.8 lm, n = 20), septate,
paraphyses, longer than the asci, tapering towards the apex,
wider at the base. Asci 95–100 9 14–20 lm
(
x = 100 9 18 lm, n = 25), multi-spored, unitunicate,
clavate, with narrow, long, thin-walled pedicel, with
swollen upper portion, apex rounded, without an visible
apical apparatus. Ascospores 6.2–7.4 9 1.4–1.8 lm
(
x = 6.7 9 1.6 lm, n = 20), overlapping, pale brown,
allantoid to cylindrical, unicellular, with small, fat globules
at the ends, smooth-walled, without sheath or appendages.
Asexual morph Undetermined
Culture characters: Ascospores germinated on PDA
within 18 h and germ tube produced from one or both ends
of the ascospore. Reaching 6.5 cm within 14 days on PDA,
at 25 C, colonies medium dense, circular, flat, with diffuse
margin, white, below similar in color, not zonate, not
producing pigmentation on PDA media.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Mae Fah Luang University, in front of S7
building, on dried seed pods of Delonix regia (Boj. ex
Hook.) Raf. (Fabaceae), 10 December 2014, R.H. Perera,
RHP 75 (MFLU 16-1016, holotype), ex-type living culture
(MFLUCC 15-1014); ibid. RHP 77 (MFLU 16-1032).
GenBank numbers: ITS: MH812994, LSU: MH812996,
TUB2: MH847790 (MFLUCC 15-1014); ITS: MH812995,
LSU: MH812997, TUB2: MH847791 (MFLU 16-1032).
Notes: Diatrypella delonicis clusters with D. major, D.
frostii and D. vulgaris within Diatrypella sensu stricto
(Fig. 131). DNA sequences of D. delonicis and putative
strain of D. major (1058) differed in 2 nucleotides in the
179
ITS region. DNA sequences of D. delonicis and putative
strain of D. frostii (UFMGCB 1917) differed in 10
nucleotides in the ITS region. Sequence data of the TUB2
region of D. major and D. frostii are not available in the
GenBank for the comparison. Diatrypella delonicis and extype strain of D. vulgaris (HVGRF03) showed 1 nucleotide
difference in the ITS region, and 14 different nucleotides in
the TUB2 region. DNA sequences of D. delonicis and extype strain of D. tectonae (MFLUCC 12-0172a) differed in
7 nucleotide positions of the ITS region. DNA sequence
data of D. heveae (ex-type MFLUCC 17-0368) differed in
11 nucleotides in the ITS region and 33 in the TUB2. A key
to species is provided above including the phylogenetically
closely related species and the taxa were described from
Thailand, D. heveae and D. tectonae.
Peroneutypa Berl.
Notes: Peroneutypa is typfied by P. bellula (Desm.) and
currently there are 47 estimated species (Wijayawardene
et al. 2017a). The asexual morphs for the genus are
unknown. The species within Pernoneutypa are saprobes,
pathogens and are widely distributed in terrestrial habitats
(Lumbsch and Huhndorf 2010; Maharachchikumbura et al.
2015; Shang et al. 2017). Cultures and sequences for the
genus are available, but are lacking for the type.
Peroneutypa scoparia Carmarán & A.I. Romero (2006)
Facesoffungi number: FoF01998; Fig. 132
Holotype: MFLU 19-0623
Saprobic on decayed wood at a mangrove stand. Sexual
morph Ascomata 234–347 9 347–439 lm, solitary to
scattered, lacking stromatic tissues, deeply immersed, dark
brown to black, globose to subglobose, uni to multi-loculate, ostiolate, papillate. Ostioles central, apex composed of
brown outer amorphous layer and inner yellow cells of
textura porrecta and ostiolar canal filled with periphyses,
195–530 lm high, 330–720 lm wide. Peridium up to 25–
35 lm wide, thick, 3-layered, incorporating occasionally in
the host tissues. Paraphyses 1.5–2 lm wide, arising from
the base of perithecia, short, thin-walled cells, guttulate,
unbranched with apex blunt. Asci 19.5–23 9 3.2–5 lm
(
x = 21 9 4.2 lm, n = 20), 8-spored, unitunicate, clavate,
long pedicellate, apically rounded, with J-, subapical ring,
initially deliquescing. Ascospores 2.8–4.8 9 1.4–1.9 lm
(
x = 4.2 9 1.4 lm, n = 40), overlapping, allantoid, hyaline to light brown, aseptate, not constricted at the septum,
smooth-walled with one to few small guttules at the both
ends, straight to slightly curved. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA, reaching
10 cm diam. after 2 weeks at 20–25 C, medium dense,
irregular to regular, slightly raised, surface slightly rough,
with uneven edge, undulate margin, cottony, colony from
above: pale grey at the margin, greenish-grey at the centre;
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b Fig. 127 Phylogram of maximum likelihood analysis based on ITS,
TUB2 and TEF1-a sequences for Pestalotiopsis species. Related
sequences were obtained from Maharachchikumbura et al.
(2012, 2014). Neopestalotiopsis saprophyta (MFLUCC 12-0282) is
used as the outgroup taxon. Sixty five strains are included in the
combined analyses which comprise 1545 characters (565 characters
for ITS, 472 characters for TUB, 508 characters for TEF1-a) after
alignment. Single gene analyses are carried out and the topology of
each tree had clade stability. Tree topology of the maximum
likelihood analysis is similar to the maximum parsimony analysis.
The best RaxML tree with a final likelihood value of - 10917.490545
is presented. The matrix had 716 distinct alignment patterns, with
12.01% undetermined characters or gaps. Estimated base frequencies
were as follows: A = 0.234122, C = 0.292180, G = 0.213547,
T = 0.260152; substitution rates AC = 1.238816, AG = 3.435200,
AT = 1.254551, CG = 1.048645, CT = 4.518698, GT = 1.000000;
gamma distribution shape parameter a = 0.803705. Maximum parsimony analysis of 945 constant characters and 384 informative
characters resulted in two equally most parsimonious tree of 1643
steps (CI = 0.525, RI = 0.685, RC = 0.360, HI = 0.475). Bootstrap
support values for ML (first set) and MP equal to or greater than 50%
are given above the nodes. The ex-type strains are in bold and black.
The newly generated sequence is indicated in bold and blue
from below: yellow to brown; mycelium greenish-grey, not
produced pigmentation on PDA media.
Material examined: THAILAND, Phetchaburi Province,
Cha-am District, from a dieback diseased marine wood, 11
January 2018, Vin. Kumar, KRB010 (MFLU 19-0623),
living culture (MFLUCC 18-1111).
GenBank numbers: ITS: MK603519, TUB2:
MK101307.
Notes: The species Peroneutypa scoparia (MFLUCC
18-1111) is similar to the type of P. scoparia (MFLUCC
11-0478) in Diatrypaceae, based on molecular data and
morphology. It is characterized by immersed ascomata
with a cylindrical or subconical, periphysate ostiole, apically rounded, clavate asci and allantoid, aseptate, and
hyaline to light brown ascospores. Peroneutypa aeriensis
has unique ascomatal characters with multiple, short necks
Phylogenetically P. scoparia (MFLUCC 18-1111) shares
the same clade with P. scoparia (MFLUCC 11-0478) with
a close affinity. Our isolate grouped within the Peroneutypa clade with low statistical support. This species is
associated to the dieback symptoms. Here we add a TUB2
sequence with our entry. This is the first Peroneutypa
species reported from a mangrove habitat (Jones et al.
2015, 2019).
Xylariales Nannf., genera incertae sedis
Notes: Xylariales have been revised by several recent
studies researchers (Tang et al. 2009; Maharachchikumbura et al. 2015, 2016; Senanayake et al. 2015; Wendt et al.
2018; Daranagama et al. 2018; Voglmayr et al. 2018;
Wijayawardene et al. 2018a; Doilom et al. 2018).
Maharachchikumbura et al. (2016) accepted 24 genera,
181
while Wijayawardene et al. (2018a) listed 44 genera in
Xylariales, genera incertae sedis.
Fasciatispora K.D. Hyde
The most recent account of this genus are by Daranagama et al. (2018) who we follow here.
Fasciatispora arengae Konta & K.D. Hyde
Facesoffungi number: FoF05113; Fig. 133
Culture characteristics: Ascospores germinated on
MEA within 24 h and germ tube was produced from both
ends. Colony on MEA, at first whitish, felty, azonate, with
fluffy margins, hyphae, septate, branched, and smooth.
Material examined: THAILAND, Phang-nga Province,
on dead leaflet of Arenga pinnata (Arecaceae), 5 December
2014, S. Konta, PHR07 g (MFLU 15-0301), ex-type living
culture (MFLUCC 15-0326).
GenBank numbers: ITS: MK120275, MK120276,
MK120277, LSU: MK120300, MK120301, MK120302,
SSU: MK120304, MK120305, MK120306.
Notes: Doilom et al. (2018) introduced Fasciatispora
arengae from southern Thailand based on morphology.
Phylogenetic analyses indicated that F. arengae clustered
with other species in the ‘Fasciatispora’ clade; F. nypae
and F. calami (99 MP/1.00 BYPP, Fig. 134). Accordingly,
Hyde (1991), Wijayawardene et al. (2018a) and Doilom
et al. (2018) assigned Fasciatispora to Amphisphaeriales
genera incertae sedis, which was confirmed by Daranagama et al. (2018). Subsequently, several studies
transferred Fasciatispora to Xylariaceae (Alias et al. 1994;
Hyde 1995a; Hidayat et al. 2007; Hyde and Wong 1999;
Kang et al. 1999; Liu et al. 2015; Hyde et al. 2017a). In this
study, phylogenetic analysis of combined four gene regions
(TUB2, ITS, LSU, and RPB2) showed that Fasciatispora
did not form a clade close to Amphisphaeriaceae as was
mentioned in Daranagama et al. (2018), it formed a separate between the Graphostromataceae and ‘Hypoxylon’
clades (Fig. 134). Fasciatispora is, therefore, best accommodated in Xylariales genera incertae sedis.
Basidiomycota R.T. Moore
Agaricomycetes Doweld
Agaricomycetidae Parmasto
Agaricales Underw.
Cortinariaceae R. Heim ex Pouzar
Notes: The limits of the family Cortinariaceae remain
unknown. Most species in this family are in the genus
Cortinarius, with a cosmopolitan distribution and over
2000 described species. Many genera formerly placed in
the Cortinariaceae, (e.g., Phaeocollybia, Hebeloma, Galerina) have been transferred to other families in Agaricales (Matheny et al. 2015). Furthermore, the sequestrate
genera, Thaxterogaster, Quadrispora, Protoglossum and
Hymenogaster pro parte as well as Cuphocybe, Rapacea
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Fig. 128 Castanediella meliponae (URM 91974, holotype). a Surface of nest of Melipona scutellaris. b (Left to right) Surface and reverse of
colony growth on PDA. c Conidiogenous cell and conidia. d Conidiophore and conidia. e Conidia. Scale bars: c–e = 10 lm
and species of Rozites, once thought to be genera within the
Cortinariaceae, are currently included in the genus Cortinarius (Peintner et al. 2001, 2002). The basidiocarps range
from agaricoid to sequestrate, and many have poorly to
well–developed veils. The basidiospores are typically
ornamented and cinnamon brown in deposit.
Cortinarius (Pers.) Gray
Notes: Cortinarius species are important ectomycorrhizal fungi and are associated with different trees and
shrubs, belonging to the families Fagaceae, Salicaceae,
Caesalpiniaceae, Cistaceae, Dipterocarpaceae, Myrtaceae,
Rhamnaceae, Rosaceae and Pinaceae, as well as some
herbaceous plants in the Cyperaceae and Polygonaceae
(Frøslev et al. 2006). The basidiocarps range from agaricoid to sequestrate, and many have poorly to well-developed veils. The basidiospores are typically ornamented and
cinnamon brown in deposit. Hundreds of species are still
undescribed and below we present two new species
belonging to Cortinarius subgenus Telamonia, sect. Safranopedes (Bidaud, Moënne-Locc. & Reumaux) Liimat.,
Kytöv. & Niskanen from Europe.
Cortinarius minusculus Liimat. & Niskanen, sp. nov.
Index Fungorum number: IF555637; Facesoffungi
number: FoF05970; Fig. 135a
123
Etymology: Name based on a Latin adjective minusculus, meaning rather small.
Holotype: K. Liimatainen & T. Niskanen 12-032 (H)
Pileus 0.7–1.5 cm wide, at first conical to somewhat
convex, later planoconvex, often with an acute umbo, redbrown, paler at the margin, hygrophanous. Lamellae
medium spaced, adnexed to emarginated, brown. Stipe
2.5–5 cm long, 0.15–0.3 cm thick, cylindrical, at first very
pale greyish brown, covered with silky, white fibrils, later
darker, especially at the lower of the stipe. Context: in
pileus red brown, in stipe yellow brown, dark brown at the
base when older. Universal veil white, forming often distinct complete and incomplete on the stipe. Odour in
lamellae indistinct. Exsiccatae: pileus brown, stipe greyish
brown. Basidiospores (7.5–)8–9(–9.5) 9 4.7–5.3 lm,
av. = 8.5 9 4.9 lm, Q = 1.63–1.8, Qav. = 1.74, ellipsoid
to obovoid-ellipsoid, moderately verrucose, more strongly
at apex, somewhat dextrinoid. Lamellar trama hyphae
encrusted.
ITS sequence (GenBank MK211177, holotype) distinct
from other members of C. sect. Safranopedes. With a sister
group relationship to C. scotoides and deviating from it by
10 substitutions and indels.
Ecology and distribution: In hemiboreal and temperate
mixed forests with Betula, Corylus, Populus and Pyrola
rotundifolia on mull soil. Known from northern Europe.
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183
Fig. 129 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data for Castanediella
species and several closely related genera. Related sequences are
taken from Hernández-Restrepo et al. (2017) and Wanasinghe et al.
(2018a). Seventeen strains are included in the combined analyses
which comprise 1037 characters (557 characters for ITS, 480
characters for LSU) after alignment. Beltraniella endiandrae (CBS
137976) (Amphisphaeriaceae, Amphisphaeriales), Beltrania pseudorhombica (CBS 138003) (Beltraniaceae, Sordariales) and Beltraniopsis neolitseae (CBS 137974) (Incertae sedis) are used as the
outgroup taxa. Single gene analyses are carried out and the topology
of each tree had clade stability. Tree topology of the maximum
likelihood analysis is similar to the Bayesian analysis. The best
RaxML tree with a final likelihood value of - 3681.798352 is
presented. The matrix had 245 distinct alignment patterns, with 6.62%
undetermined characters or gaps. Estimated base frequencies were as
follows: A = 0.284691, C = 0.205176, G = 0.254615, T = 0.255518;
substitution rates AC = 0.678292, AG = 2.785917, AT = 1.405179,
CG = 0.578526, CT = 3.212102, GT = 1.000000; gamma distribution shape parameter a = 3.725904. Bootstrap values for maximum
likelihood (ML) equal to or greater than 75 are placed above the
branches respectively. The newly generated sequence is indicated in
bold and blue
Material examined: Finland, Uusimaa, Vantaa, Tammisto, Tammisto’s Nature Reserve Area, herb-rich mixed
forest with Quercus, Corylus, Betula, Pinus and Populus,
17 September 2012, K. Liimatainen & T. Niskanen 12-032
(H, holotype; K, isotype).
GenBank numbers: ITS: MK211177 (loc. cit. 12-031
(H)), MK211178.
Notes: Cortinarius minusculus is a small and slender
medium brown Telamonia with distinct white veil, indistinct odour, and ellipsoid to obovoid-ellipsoid spores. It
grows on mull soil with several different deciduous trees
and Pyrola. The species formed a well-supported monophyletic group within C. sect. Safranopedes in our phylogenetic analysis.
Cortinarius subscotoides Niskanen & Liimat. sp. nov.
Index Fungorum number: IF555638; Facesoffungi
number: FoF05971; Fig. 135b
Etymology: The name refers to the affinity to C.
scotoides.
Holotype: K. Liimatainen & T. Niskanen 12-010 (H)
Pileus 2–4 cm wide, at first conical, later planoconvex,
with an acute umbo, dark brown, paler at the margin,
hygrophanous. Lamellae medium spaced, adnexed to
emarginated, at first pale greyish brown, later brown. Stipe
2.5–5 cm long, 0.5–0.9 cm thick, cylindrical, at first silky
whitish fibrillose, soon brown, especially at the base of the
stipe. Context: in pileus dark brown, in stipe brown, darker
brown at the base. Universal veil white, rather thin,
forming a complete girdle on the middle part of the stipe.
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Fig. 130 Diatrypella delonicis (MFLU 16-1016, holotype). a Herbarium material. b Appearance of ascomata on host substrate. c Longitudinal section through stroma with ascomata. d Vertical section
through ascomata. e Close up of the peridium. f Paraphyses. g–i Asci.
j Ascospores. k Germinating ascospore. l, m Colony on PDA (l from
above view, m from below view). Scale bars: b = 1 mm,
d = 500 lm, e–i, k = 20 lm, j = 10 lm
Odour in lamellae indistinct. Exsiccatae: pileus dark
brown, stipe greyish brown. Basidiospores 7–8 9 4.8–
5.3 lm, x = 7.5 9 5.1 lm, Q = 1.4–1.6, x = 1.47, lacrymoid to shortly obovoid-ellipsoid, strongly verrucose,
somewhat dextrinoid. Lamellar trama hyphae encrusted.
ITS sequence (GenBank MK211175, holotype) distinct
from other members of C. sect. Safranopedes. With a close
relationship to C. scotoides and deviating from it by five
substitutions and indels.
Ecology and distribution: In hemiboreal and temperate
mixed forests with Corylus on mull soil. Known from
northern Europe.
Material examined: Finland, Varsinais-Suomi, Lohja,
Virkkala, Pähkinäniemi, Corylus forest with some Betula,
Populus and Picea on calcareous ground, 16 September
2012, K. Liimatainen & T. Niskanen 12-010 (H, holotype;
K, isotype).
GenBank numbers: ITS: MK211175 (loc. cit. K. Liimatainen & T. Niskanen 12-015 (H)), MK211176.
Notes: Cortinarius subscotoides has a dark brown
pileus, short but stout stipe, indistinct smell, and lacrymoid
to shortly obovoid-ellipsoid, strongly verrucose spores. It
grows on mull, calcareous soil at least with Corylus. In our
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phylogenetic analysis the species formed a well-supported
clade within C. sect. Safranopedes (Fig. 136).
Psathyrellaceae Vilgalys et al.
Notes: Psathyrellaceae was proposed by Redhead et al.
(2001) based on molecular phylogenetic studies. Initially,
the family comprised the genera Psathyrella (Fr.) Quél.,
Lacrymaria Pat., Coprinellus P. Karst., Coprinopsis P.
Karst., and Parasola Redhead et al. The latter three genera
were segregated from the polyphyletic genus Coprinus
Pers. s. lat. in the same paper. Only a few species (Coprinus comatus (O.F. Müll.) Pers. and related taxa)
remained in the Coprinus sensu stricto, and the genus was
transferred to Agaricaceae Chevall. Subsequently, several
smaller genera were added to Psathyrellaceae. According
to Kirk et al. (2008), it comprises 12 genera.
Coprinopsis P. Karst.
Notes: Members of the genus Coprinopsis live as
saprotrophs on various substrata such as wood, soil, dung,
herbal remnants or burnt ground. Its basidiomes are
agaricoid, brittle, and often short lived and/or deliquescent,
with blackish to black spore print (Vesterholt 2012). The
genus contains around 200 species globally (Kirk et al.
2008). The phylogenetic tree is shown in Fig. 137.
Coprinopsis psammophila Mešić & Tkalčec, sp. nov.
Index Fungorum number: IF555781; Facesoffungi
number: FoF05972; Fig. 138
Etymology: The species is named after its substrate,
sandy soil in a desert shrubland.
Holotype: CNF 1/6401
Pileus 10–25 mm broad at maturity, subglobose to
ellipsoid at first, later obtusely or truncately conical, often
with inflexed margin before expanding, finally subapplanate and radially splitting, partially deliquescent, surface finely and shallowly plicate-sulcate except at the
centre, pale grey-brownish to light brown at first, later grey
to grey-brown, finally black(ish), entirely covered with a
delicate, poorly developed, whitish to cream coloured,
silky veil. Lamellae free, rather crowded, up to 2.5 mm
broad, soon becoming black, partially deliquescent. Stipe
20–70 9 1.5–3.5 mm, cylindrical, mostly with a broadened (bulbous to subcylindrical) base (up to 6 mm wide),
with rather long, completely distinctly narrower or gradually narrowing pseudorhiza, central, widely hollow, whitish, dry, minutely fibrillose-floccose, sometimes striate;
basal part always buried in the sand, but often also its lower
half, or even most of it. Context very thin. Odour and taste
not recorded. Spore print black.
Basidiospores [300/3/1] (8.5–)8.8–9.8–10.9(–11.2) 9
5.8–6.4–7.2(–7.5) lm (in KOH 2.5%), averages of different basidiomes 9.5–10 9 6.3–6.6 lm, Q = 1.31–1.52–
1.71, av. Q = 1.47–1.56, ellipsoid or suboblong
185
(subcylindrical), more rarely (sub) ovoid, not flattened,
with rounded to slightly acute base and very obtuse apex,
smooth, slightly to moderately thick-walled (up to 0.6 lm),
with wide (2–3.2 lm in KOH, 1.4–2.5 lm in H2O) and
central germ-pore, brown-black in KOH, dark red-brown in
H2O, opaque, non-amyloid and non-dextrinoid. Basidia
13–28 9 6–10 lm, clavate, 4-spored, thin-walled, hyaline,
surrounded by (3–)4–6(–7) hymenophysalides (pseudoparaphyses). Cheilocystidia not observed. Pleurocystidia
conical or fusiform, up to ca. 100 lm long and 18–45 lm
wide, thin-walled, hyaline, rather abundant (but scattered)
to scarce. Pileipellis a cutis, composed of repent, hyaline,
thin-walled, 2–20 lm wide hyphae, with narrowest hyphae
on the surface. Veil hyphae 1–9 lm broad, occasionally
branched, not or only slightly constricted at the septa, thinwalled to slightly thick-walled (up to 0.5 lm), sparsely
diverticulate (excrescences pretty wide with rounded tip),
hyaline, mostly smooth, sometimes finely to coarsely
encrusted. Clamp connections absent.
Habit and habitat: In a scattered group (approximately
30 basidiomes at a radius of ca. 10 m), terrestrial, on a
sandy soil among sparse herbaceous plants and low bushes
in an arid, lowland, Mediterranean desert shrubland.
Distribution: known only from the type locality in
Libya, Sirte District.
Material examined: LIBYA, Sirte District, by Qasr Abu
Hadi town, 13.9 km S-SE from Sirte, 31050 5500 N,
16390 5400 E, 46 m a.s.l., on sandy soil in a desert shrubland,
14 November 2008, leg. M. Čerkez (CNF 1/6401,
holotype).
GenBank numbers: ITS: MK491274, LSU: MK492278.
Notes: Morphologically and ecologically, Coprinopsis
psammophila is characterised by occasionally branched
and sparsely diverticulate veil hyphae (sect. Alachuani
(Singer) D.J. Schafer), mostly ellipsoid or suboblong
(subcylindrical), smooth basidiospores with very obtuse
apex and wide germ-pore, absence of clamp connections,
presence of pseudorhiza, often bulbous stipe base, and
terrestrial growth on sandy soil. This set of characters
clearly differentiate C. psammophila from all other species
in the genus. Among all known species in the section
Alachuani, only C. alcobae (A. Ortega) Valade also lives
on sandy soil, but has no pseudorhiza, and differs in other
morphological characters. Furthermore, only the poorly
known Coprinus dryophilus Pat. has a rooting stipe, but
lives on wood and also differs in other morphological
characters. A megablast search of NCBI’s GenBank
nucleotide database, using the ITS sequence from our
holotype collection of C. psammophila, showed no closely
related species. The closest hit was C. verticillata (SchulzWedd.) Redhead et al. (GenBank Number MH861734,
Identities = 653/725(90%), Gaps = 23/725(3%)). In phylogenetic tree of Coprinopsis species based on ITS
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186
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Fungal Diversity (2019) 96:1–242
b Fig. 131 Phylogram generated from maximum likelihood analysis
based on combined ITS and TUB2 sequence data of selected taxa.
Related sequences were obtained from GenBank. Ninety-five strains
are included in the analyses, which comprise 971 characters including
gaps. Single gene analyses were carried out and compared with each
species, to compare the topology of the tree and clade stability. Tree
was rooted with Kretzschmaria deusta (CBS 826.72) and Xylaria
hypoxylon (CBS 122620). Tree topology of the ML analysis was
similar to BI. The best scoring RAxML tree with a final likelihood
value of - 9028.552721 is presented. The matrix had 404 distinct
alignment patterns, with 11.19% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.232808,
C = 0.250740, G = 0.235523, T = 0.280930; substitution rates AC =
1.126828, AG = 3.047992, AT = 1.477104, CG = 0.932402, CT =
3.736033, GT = 1.000000; gamma distribution shape parameter
a = 0.466736. The maximum parsimonious dataset consisted of
constant 316, 546 parsimony-informative and 109 parsimony-uninformative characters. The parsimony analysis of the data matrix
resulted in the maximum of two equally most parsimonious trees with
a length of 2920 steps (CI = 0.407, RI = 0.751, RC = 0.306,
HI = 0.593) in the first tree Maximum likelihood bootstrap (ML)
values [ 65%) and Bayesian posterior probabilities (PP) [ 0.90%
and are given above the nodes. The scale bar indicates 0.2 changes.
The ex-type strains are in bold and black. The newly generated
sequences are indicated in bold and blue
sequences (Fig. 139) C. psammophila is nested near two
other species of sect. Alachuani, C. strossmayeri (Schulzer)
Redhead et al. and C. picacea (Bull.) Redhead et al.
Coprinopsis villosa L. Nagy, Desjardin, Vágvölgyi &
Papp, in Nagy et al., Mycologia 105(1): 120 (2013)
Index Fungorum number: IF487717; Facesoffungi
number: FoF05973; Fig. 140
Pileus up to 20 mm high and 7 mm broad when still
unexpanded, narrowly ellipsoid at first, then narrowly
conical (never expanded in our collection), plicate-sulcate
except at the centre, white at first, then becoming brownish-grey to grey, covered with medium abundant, rather
loosely attached, floccose veil, reddish brown all over or
light brown at the wider central zone and whitish around.
Lamellae free, crowded, white at first, black at maturity,
with whitish edge. Stipe up to 32 9 3 mm, cylindrical or
somewhat tapering towards the base, central, widely hollow, dry, tomentulose, white. Odour and taste not
observed. Spore print black.
Basidiospores
[100/2/1]
8.6–10.1–11.4 9 6.2–7–
7.8 lm (in KOH 2.5%), averages of different basidiomes
9.6–10.7 9 6.7–7.3 lm,
Q = 1.30–1.45–1.57,
av.
Q = 1.42–1.47, ellipsoid to ovoid in frontal view, often
(sub) amygdaloid in side view, not flattened, with rounded
to slightly conical base and rounded apex, smooth, thickwalled (1–1.2 lm), with 1.2–1.8 lm wide, central germpore, medium rusty brown in H2O, medium brown in KOH,
semi-transparent, non-amyloid and non-dextrinoid. Basidia
17–33 9 8–11 lm, clavate or spatulate with rounded or
subcylindrical upper part, 4-spored, thin-walled, hyaline,
187
surrounded by 3–5(–6) hymenophysalides (pseudoparaphyses). Cheilocystidia up to ca. 40 lm in diameter, mostly
subglobose or ellipsoid, thin-walled, hyaline, very delicate
and easily collapsing in our material. Pleurocystidia
30–65 9 20–45 lm, ovoid, conical, oblong (subcylindrical) or ellipsoid, thin-walled, hyaline, rather abundant but
scattered. Veil on pileus composed of two types of elements: (1) ellipsoid to elongated cells, often inflated and/or
constricted at septa, arranged in chains forming tufts, up to
90 lm wide, gradually narrowing towards the pointed tuft
tip (to only a few lm), not branched or diverticulate,
subhyaline to yellow–brown (with intracellular pigment),
thin-walled to moderately thick-walled (up to 1 lm), and
(2) narrow, 1–6 lm wide, branched, sparsely diverticulate,
hyaline, thin-walled to moderately thick-walled (up to
0.8 lm) hyphae laying on the broader ones. Pileipellis a
cutis, composed of repent, ellipsoid to elongated, mostly
inflated, hyaline, thin-walled to moderately thick-walled
(up to 1 lm), 4–50 lm wide hyphae. Clamp connections
present and rather abundant in all tissues.
Material examined: CROATIA, Zagreb County,
800 m W-NW from Kostanjevac Podvrški (near Samobor),
45490 5100 N, 15350 1700 E, 250 m a.s.l., 13 October 2007,
leg. M. Čerkez (CNF 1/4862, new country record).
Habit and habitat: On horse dung (Nagy et al. 2013).
Our collection was composed of five immature basidiomes
(spores matured after sampling) growing from a heap of
horse dung mixed with straw.
Distribution: Hitherto known from Croatia, Germany,
Hungary, Spain, Sweden and USA (Hawaii) (Nagy et al.
2013; Ruiz & Cerdán 2016).
GenBank numbers: ITS: MK491273, LSU: MK492277.
Notes: ITS sequence obtained from our material of
Coprinopsis villosa is identical to the sequence from the
type material (GenBank number HQ847031). C. villosa is
characterised by fimicolous habitat (horse dung), brown
veil on the pileus (at least near the centre), smooth and
medium-sized basidiospores, veil composed of two types
of elements (broader, unbranched, non-diverticulate cells
in chains together with a narrower, branched and sparsely
diverticulate cells), and the presence of clamp connections.
According to the dominant type of velar elements, it
belongs to the section Lanatuli (Fr.) D.J. Schafer. The only
other known Coprinopsis species that has a veil on the
pileus composed of sausage like elements together with a
branched and diverticulate elements is C. candidolanata
(Doveri & Uljé) Keirle et al. However, it has a pure white
veil, densely diverticulate narrower veil hyphae, no clamp
connections, and it occurs on deer and sheep dung. In the
protologue of C. villosa, Nagy et al. (2013) wrote that they
did not see clamp connections. However, in Croatian,
Spanish (Ruiz & Cerdán 2016) and Hawaiian material
(Keirle et al. 2004; mentioned in the protologue under
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Fig. 132 Peroneutypa scoparia (MFLU 19-0623, new record). a–c Appearance of ascomata on host. d Vertical section through ascoma. e, f
Ostiole, section through neck with periphyses. h, i Asci. j–m Ascospores. Scale bars: d = 100 lm, e, f = 50 lm, g–m = 5 lm
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‘‘Other specimens examined’’) clamp connections are
present. A length/width ratio (Q) of basidiospores is
another difference worth mentioning between the original
description and the description of Croatian material.
Basidiospores reported in original description are more
elongated (Q = 1.47–1.57–1.64) than those measured in
Croatian material (Q = 1.30–1.45–1.57). C. villosa is
reported here as new to Croatian mycobiota. An overview
of coprinoid taxa recorded for Croatia until 2000 is given
by Mešić & Tkalčec (2003).
Cantharellales Gäum.
Notes: The order Cantharellales represents most likely
the oldest lineage among homobasidiomycetes and equally
the oldest lineage in which the ectomycorrhizal mode of
nutrition has been acquired. However, this lineage also
harbours few lichen-forming, several lichenocolous as well
as many saprotrophic and parasitic species (Hibbett et al.
2014). Also referred to as the cantharelloid clade, this
lineage typically contains species producing basidia with a
variable number of spores depending on the genus. Two
sterigmata are found in many Clavulina and Membranomyces, for example, as well as in some rare tropical
African Cantharellus, but a variable number from four to
eight sterigmata per basidium in a single species is the
situation that predominates in Cantharellus, Craterellus,
Botryobasidium and Sistotrema.
Botryobasidiaceae Julich.
Notes: It was first introduced as subfamily Botryobasidioideae of the Corticiaceae in 1958, and was validly
published by Parmasto (1968). Later on, Jülich (1982)
raised the status of subfamily to the rank of family, as the
Botryobasidiaceae. Corticioid genera Botryobasidium,
Botryodontia, Botryohypochnus (considered a synonym of
Botryobasidium), Candelabrochaete, Suillosporium, and
Waitea, were described based mainly on similarities in the
their basidiocarp micromorphology. Botryobasidium has
smooth spores and is usually 6–8 sterigmata, while
Botryophypochnus produces spiny spores and contains
4-sterigmate basidia. Langer (1994) based on molecular
data analysis suggested for synonymization and proved
Botryobasidium as paraphyletic. Almost all species produce thin and delicate basidiomata with distinct wide
hyphae. Many species are associated with asexual morph
genera like Haplotrichum, Acladium, etc. The proposed
taxon has somewhat similar in morphological features of
Acladium, but molecular data strongly supports to be distinct from all other known genera of Botryobasidiaceae.
Therefore, this taxon is being proposed as new genus.
189
Neoacladium P.N. Singh & S.K. Singh, gen. nov.
Index Fungorum number: IF556212; Facesoffungi
number: FoF5687
Etymology: Name refers to its morphological similarity
to the genus Acladium.
Type species: Neoacladium indicum P.N. Singh & S.K.
Singh
Saprobic on dead bark of tree of terrestrial habitats.
Asexual morph. Colonies effuse, cottony or velvety, rhizoidal, light yellow,powdery, reverse greyish orange.
Mycelium mostly semi-immersed or immersed, single or
interwoven, light olivaceous. Stroma none. Chlamydospores produced from lateral hyphae, bulbous and
inflated, simple to branched, constricted near septa, pigmented, trident shaped, subhyaline to light olivaceous.
Setae and hyphopodia absent. Conidiophores semimacronematous, mononematous, fasciculate, dichotomously loosely interwoven, branched, straight or flexuous,
subhyaline to light olivaceous, smooth, septate. Conidiogenous cells integrated to terminal sometimes discrete,
determinate, acropleurogenous, polyblastic, inflated or
sometimes cylindrical, persistent occasionally sessile,
denticulate, denticles numerous. Conidia solitary or catenate, dry, variously shaped, sub-globose- globose, clavateobclavate, lenticular, pyriform to ampulliform, smoothwalled, pigmented, sub-hyaline to light olivaceous. Sexual
morph Undetermined.
Notes: Phylogenetic analysis of ITS and LSU sequence
data indicates that Neoacladium is a distinct genus in
Botryobasidiaceae, which forms a clade sister to Botryobasidium. Neocladium is close to Acladium, but differs
from all other allied genera in having subhyaline to light
olivaceous variously shaped conidia, viz. globose to subglobose, clavate, obclavate, lenticular, ampulliform and
pyriform, catenate conidia, dentate and phialidic condiogenous cells and presence of abundant trident like pigmented chlamydospores.
Neoacladium indicum P.N. Singh & S.K. Singh, sp. nov.
Index Fungorum number: IF556211; Facesoffungi
number: FoF5688; Figs. 139, 141
Etymology: Specific epithet ‘‘indicum’’ refers to the
country of origin.
Holotype: AMH 10054
Saprobic on dead bark of Leucaena leucocephala L.
Asexual morph Colonies uniformly spread on dead bark,
yellow, velvety. Conidiophores semi-macronematous,
mononematous, fasciculate, dichotomously branched,
loosely interwoven, straight to flexuous, subhyaline to light
olivaceous, smooth-walled, septate, 337–752 9 9–13 lm.
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190
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b Fig. 133 Fasciatispora arengae (MFLU 15-0301, new record). a, b
Appearance of ascomata on host substrate. c Close up of ascomata.
d Section of ascoma. e Peridium. f Paraphyses. g–j Asci. k J ? reaction with Melzer’s reagen in apical ring. l Germinated ascospore.
m Colony on culture. n–s Ascospores. Scale bars: a, b = 1000 lm,
c = 200 lm, d = 100 lm, g–j = 50 lm, e, f = 20 lm, l, n–
s = 10 lm, k = 5 lm
Fig. 134 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU, RPB2 and TUB2 sequence data for
Fasciatispora species and several closely related genera in Xylariales.
Related sequences are taken from Voglmayr et al. (2018), Wendt
et al. (2018) and Daranagama et al. (2018). One hundred and fiftyseven strains are included in the combined analyses which comprise
7548 characters (1288 characters for ITS, 2397 characters for LSU,
1246 characters for RPB2, 2617 characters for TUB2) after
alignment. Microdochiaceae is used as the outgroup taxon. Single
gene analyses are carried out and the topology of each tree had clade
stability. Tree topology of the maximum likelihood analysis is similar
to the Bayesian analysis. The best RaxML tree with a final likelihood
191
Conidiogenous cells integrated to terminal, sometimes
discrete, determinate, acropleurogenous, polyblastic, inflated, sometimes cylindrical, persistent, occasionally sessile,
denticulate, denticles numerous, 1–2 celled, 0–1 septate,
subhyaline to light olivaceous, 7.48–62.92 lm (
x=
20.52 9 6.32 lm, n = 30). Conidia solitary or produced in
short chains, variously shaped, globose to sub-globose,
valued of - 156397.840016 is presented. The matrix had 4604
distinct alignment patterns, with 59.36% undetermined characters or
gaps. Estimated base frequencies were as follows: A = 0.239282,
C = 0.272288, G = 0.248516, T = 0.239913; substitution rates AC =
1.231573, AG = 3.991205, AT = 1.258243, CG = 0.936670, CT =
5.480741, GT = 1.000000; gamma distribution shape parameter
a = 0.408051. Bootstrap values for maximum likelihood (ML) equal
to or greater than 50, Bayesian posterior probabilities (BYPP) equal to
or greater than 0.9 are placed above and below the branches
respectively. Our strains generated sequences are indicated in blue.
Branches with 100% MLBS and 1.00 BYPP are shown as blue nodes
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192
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Fig. 135 Basidiomata and basidiospores (in MLZ). of a Cortinarius minusculus (K. Liimatainen & T. Niskanen 12-032, H). b Cortinarius
subscotoides (K. Liimatainen & T. Niskanen 12-015, H). Photographs Kare Liimatainen. Scale bar: a, b (basidiospores) = 10 lm
clavate, obclavate, pyriform, lenticular, sometimes broadly
ellipsoidal to ampulliform, pigmented, smooth-walled,
base truncate, subhyaline to light olivaceous, 10.5–
25.88 9 9.36–16.5 lm (
x = 15.75 9 13.55 lm, n = 30).
Sexual morph Undetermined
Culture characteristics: Conidia germinating on malt
extract agar (MEA). Colonies light yellow (4A4), reaching
4.2 cm diam. in 10 days at 25 C, rhizoidal with irregular
margin, surface powdery, reverse greyish orange (5B5).
Hyphae septate, unbranched to branched, pigmented,
smooth and thin walled, subhyaline to light olivaceous,
5–5.62 lm wide.
Material examined: INDIA, Maharashtra, Pune District,
on dead bark of Leucaena leucocephala (Fabaceae), 30
July 2017, P.N. Singh; (AMH 10054, holotype), ex-type
living culture (NFCCI 4480).
GenBank numbers: ITS: MK391496, LSU: MK391493.
Notes: The proposed taxon is compared with the asexual
genus Acladium in having subhyaline to light olivaceous
ampulliform to pyriform conidia produced from tubular to
inflated and conidiogenus cells which are persistent, and
occasionally sessile. Chlamydospores in the proposed
taxon produced laterally and form trident like structure,
which are pigmented. Most of these features are absent in
Fig. 136 Phylogram resulting from the RaXML (Stamatakis 2014)
analysis of ITS regions. Bootstrap values greater than 50% are
indicated above branches. The names in blue represent the new
species of Cortinarius and the specimens in boldface the type
specimens of the species. The tree is rooted with C. section Bovini.
The ex-type strains are in bold. The newly generated sequences are
indicated in blue
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Fig. 137 Maximum likelihood phylogenetic tree of Coprinopsis
species based on an ITS1–5.8S–ITS2 DNA sequence alignment and
calculated with PhyML software (Guindon et al. 2010). Related
sequences are taken from Nagy et al. (2011, 2012, 2013), Raut et al.
(2011), Osmundson et al. (2013), Raut et al. (2015), Tibpromma et al.
(2017), Phookamsak et al. (2019) and Vu et al. (2019). Forty-three
strains are included in the gene sequence analyses with total of 737
characters after alignment. The tree is rooted with Coprinellus
xanthothrix (SZMC-NL-3417) and C. domesticus (SZMC-NL-1292).
The best PhyML tree with a final likelihood value of -3591.26595 is
193
presented. Estimated base frequencies were as follows: A = 0.23036,
C = 0. 0.22632, G = 0.21420, T = 0.32912; substitution rates AC =
1.48357, AG = 3.68147, AT = 1.56733, CG = 0.64802, CT =
4.50774, GT = 1.00000; gamma distribution shape parameter
a = 0.570; proportion of invariant is 0.348. The ex-type strains are
indicated in bold. The newly generated sequences are indicated in
blue. Maximum likelihood bootstrap values greater than 70% are
indicated at the nodes. The bar indicates the number of nucleotide
substitutions per site
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194
Fungal Diversity (2019) 96:1–242
Fig. 138 Coprinopsis psammophila (CNF 1/6401, holotype). a, b Basidiomes. c–i Basidiospores. j–l Pleurocystidia. m Veil on the pileus (phase
contrast). Scale bars: a = 10 mm, b = 5 mm, c–i = 3 lm, j–m = 10 lm
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195
Fig. 139 Neoacladium indicum (AMH 10054, holotype). a Yellow
velvety fungal colonies spreaded on outer bark surface. b Stereoscopic
surface view of fungal colonies from substratum. c Colony morphology (above view). d Dichotomously branched conidiophores. e Enlarged microscopic view of dichotomously branched conidiophores
with intercalary, lateral and terminal dentate conidiogenous cells and
conidia. f A detached branch of conidiophore bearing conidiogenous
cells and conidia. g Catenate conidia. h–i Trident shaped and inflated
chlamydospores. Scale bars: d–i = 20 lm
Acladium. However, the conidial shape, length–width are
very much variable in N. indicum, which separate it from
other reported taxa in this group. Proposed taxon is also
distinct from other genera in this group, like Burgoa,
Minimedusa, Ceratobasidium, (Rogers 1935; Weresub and
LeClair 1971) in having non-bulbuliferous vegetative
parenchymatous propagules. During this study the sexual
stage Botryobasidium of Acladium was not encountered.
The proposed N. indicum (AMH 10054 holotype), is
different from other Botryobasidium species based on the
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196
Fungal Diversity (2019) 96:1–242
Fig. 140 Coprinopsis villosa (CNF 1/4862, new geographical record). a Basidiomes. b Basidiospores. c Veil on the pileus. d Veil on the pileus
(phase contrast). e Cheilocystidia (phase contrast). f–h Pleurocystidia. Scale bars: a = 3 mm, c = 100 lm, e–h = 20 lm, b, d = 10 lm
sequence analyses. On megablast analysis, ITS sequence of
N. indicum showed 80% (524/651) identity and 53 gaps
(8%) with B. subcoronatum, 84% (343/410) identity and 27
gaps (6%) with B. simile (GEL2348), and 81% (521/645)
identity 54 gaps (8%) with B. intertextum UC2022959. The
phylogenetic analysis clearly establishes Neoacladium as a
123
novel genus, and N. indicum as the type species with strong
bootstrap values (Figs. 142, 143).
Hydnaceae Chevall.
= Clavulinaceae Donk, Beih. Nova Hedwigia 1(4): 407
(1970)
Fungal Diversity (2019) 96:1–242
= Cantharellaceae J. Schröt., in Cohn, Krypt.-Fl. Schlesien (Breslau) 3.1 (25–32): 413 (1888) [1889]
= Sistotremataceae Jülich, Bibl. Mycol. 85: 390 (1982)
= Pterygellaceae Jülich, Bibl. Mycol. 85: 386 (1982),
= Heteroacanthellaceae P. Roberts, Mycologist 12(4):
147 (1998)
= Repetobasidiaceae Jülich, Bibl. Mycol. 85: 388 (1982)
Notes: Index Fungorum (http://www.indexfungorum.
org) and MycoBank (www.mycobank.org) are still considering Hydnaceae in a different, much narrower sense,
whereas recent phylogenetic studies (Moncalvo et al. 2006;
Hibbett et al. 2014; Lawrey et al. 2016) have demonstrated
that Hydnum L., the type genus of Hydnaceae, belongs in
the core lineage of the cantharelloid clade. Hence, the older
family name ‘‘Hydnaceae’’ has nomenclatural priority over
other family names that could be considered to name the
family that groups members of this core clade. The morphological character that seems to support this lineage is
the stichic nature of basidia (Pine et al. 1999), although this
is only based on observations for very few species. The
family is principally comprised of edible species (all
Craterellus, Cantharellus and Hydnum). Type genus:
Hydnum L., Species Plantarum: 1178 (1753)
Cantharellus Adans.
Notes: Cantharellus is an ectomycorrhizal genus of edible
mushrooms and is reputed for its high culinary and commercial value. The genus has a worldwide distribution and is
one of the prominent fungal genera in Africa (De Kesel et al.
2016) and Madagascar (Buyck et al. 2015), with several
species being sold along roads and in local markets in large
quantities. The genus is infamous for the difficulty of species
delimitation and identification (Olariaga et al. 2016). The
recent use of DNA sequences for the delimitation of species
has more than doubled the number of worldwide accepted
species over the past ten years, with numerous species continuing to be described from the African continent (e.g.
Buyck et al. 2017; Das et al. 2018a). Sequencing of recent
collections from Africa and Madagascar has also allowed for
the epitypification of many of the earlier species (e.g. Buyck
and Hofstetter 2018; Buyck et al. 2018).
Cantharellus goossensiae (Beeli) Heinem., Bull. Jard. Bot.
Etat Brux. 28(4): 406 (1958).
MycoBank number: MBT385407; Index Fungorum
number: IF556561; Figs. 144, 145
Basionym: Hygrophorus goossensiae Beeli, Bull. Société Royale Botanique Belge 61: 99 (1928).
Iconography: HEINEMANN (1958, Fig. 42; 1959, pl.
XXVII, Fig. 3).
Original diagnosis: « Pileo carnoso crasso, convexe,
glabro levi, atro-violacea, 5–6 cm lato; stipite crasso solida
dein cavo, sulcato, ochraceo, 3–4 9 2 cm; lamellis crassis
decurrentibus venoso connexis, ochraceis; sporis
197
ellipsoideis, hyalinis, levis, 8–9 9 4–5 lm; carne
ochraceopallida contactu aeris rubescente. »
Original description (freely translated from French)‘‘Pileus thick and fleshy, unevenly convex, smooth, glabrous, dark violaceous, 5–6 cm diam.; stipe thick, solid then
hollowing, ochraceous, 3–4 9 2 cm; gills thick, decurrent,
interveined, yellowish ochre; spores ellipsoid, hyaline,
smooth, 8–9 9 4–5 lm; flesh becoming pinkish when cut
or exposed.’’
Epitype description, Figs. 144, 145
Basidiomata fleshy and firm, dispersed or solitary. Pileus
82 mm diam., with strongly inrolled margin, at surface dull,
tomentose to fibrillose, yellowish brown with olive tinges,
slightly paler in the centre (4–5DE series), toward the margin
rather dull yellowish to grayish brown, musterd brown
(5E6–4, 5F4–5). Hymenophore composed of slightly higher
radial gill folds compared to the strongly developed interstitial venation, nearly off-white to pale cream when young,
then developing ochraceous tints. Stipe thick and massive,
30 9 11-26 mm, surface splitting in horizontally arranged
patches or rims in the upper portion, smooth elsewhere,
slightly paler than pileus and developing dark grayish brown
to orange brown tinges, not hollowing. Context off-white to
pale cream, moderately yellowing, finally turning ferruginous when cut, 6-7 mm thick above the upper stipe-hymenophore transition. Smell faint, but typical and fruity.
Taste mild. Spore print insufficient but very pale.
Spores ellipsoid to narrowly ellipsoid, (5.8)6.2–6.57–
7.0(7.3) 9 (3.8)4.0–4.38–4.7(5.0) lm, Q = (1.3)1.4–1.51–
1.6(1.7), smooth, hyaline. Basidia narrowly clavulate, measuring mostly 40–55(63) 9 7–8(9) lm, predominantly 4–5spored. Subhymenium filamentous. Cystidia not observed.
Pileipellis composed of long, multi-celled hyphal extremities, aggregating together in trichoids or tufts, composed of
subcylindrical, thin-walled cells; the terminal cell variable in
length, mostly 30–70 lm long, ca 10–20 lm wide and most
often wider than the subterminal cells, often slightly constricted in the middle portion, sometimes even clavate,
broadly obtuse at the tip, rarely subcylindrical or somewhat
narrowing upward. Clamp connections present everywhere.
Epitype: CENTRAL AFRICAN REPUBLIC. DzangaSangha Forest Reserve, near Bayanga, close to Bai-Hakou
base camp, N2.859934 – E16.467492, under monospecific
Gilbertiodendron dewevrei forest, on bare sandy soil, 19
May 2016, 1648/Buyck 16.064 (PC 0125008), epitype is
designated here !; Democratic Republic of the Congo.
Central African forest district, near Selenge-Lukolela,
isolated on the soil of the Gilbertiodendron dewevrei forest,
August 1925, M. Goossens-Fontana 372 (BR), holotype.
Additional material examined and sequenced: CENTRAL AFRICAN REPUBLIC. Dzanga-Sangha Forest
Reserve, near Bayanga, close to Bai-Hakou base camp,
N2.859934
–
E16.467492,
under
monospecific
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198
Fungal Diversity (2019) 96:1–242
Fig. 141 Neoacladium indicum (AMH 10054, holotype).
a Branched chlamydospores. b Hyphae and different types of conidia.
c Numerous globose to sub-globose conidia. d, e Catenate conidia.
f Lenticular conidium. g Pyriform conidium. h, i Clavate conidia. j, k
Obclavate conidia with protuberant hila. Scale bars: a–k = 20 lm
Gilbertiodendron dewevrei forest, on bare sandy soil, 16
May 2016, 1644/, Buyck 16.032 (PC 0124999); id.,
1647/Buyck 16.063 (PC 0125002), 18 May 2016,
1655/Buyck 16.080 (PC 0125003); id., 1660/Buyck 16.087
(PC0125001); id., 1662/Buyck 16.091 (PC0125004); id.,
19 May 2016, 1664/Buyck 16.096 (PC0125000); id.,
1665/Buyck 16.097 (PC0125007); id., 26 May 2016,
1679/Buyck 16.127 (PC0125005); id., 24 May 2016,
Buyck 16.114 (PC0125006).
Notes: This species, which was first described as a Hygrophorus (Beeli 1928), seems locally quite common in the
Gilbertiodendron dewevrei rain forest. The original description was based on a watercolor sketching a single, very
juvenile fruiting body. The latter, therefore, is hardly representative for what this species might look like when fully
mature, particularly when considering the impressive variability of the general field habit repeatedly reported already for
other chanterelles (Buyck et al. 2016b, 2016c; Das et al.
2018b; Olariaga et al. 2015). Moreover, the original description was very succinct, both for macro- and microscopic
features, and sketches a species characterized by a convex,
irregularly lobed and dark violet young cap sitting on top of an
ochraceous yellow, short and fleshy stipe and having a similarly colored, strongly veined-anastomosing hymenophore.
Yet, the feature that has most frequently been emphasized as characteristic for this species (Eyssartier 2001;
123
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199
Fig. 142 Molecular phylogenetic analysis of ITS gene region of
Neoacladium indicum (AMH 10054, holotype) by Maximum likelihood method. Sixteen strains are included in the sequence analyses
which comprise 668 positions after alignment. Auricularia auricularjudae CBS 7 (JF440710) (Auriculariaceae, Auriculariales) is used as
the outgroup taxon. Bootstrap values for maximum likelihood (ML)
greater than 50 are placed along the branches, respectively. The
evolutionary history was inferred by using the Maximum likelihood
method based on the Jukes-Cantor model (Jukes and Cantor 1969).
The tree with the highest log likelihood (- 4900.5990) is shown.
Evolutionary analyses were conducted in MEGA7 (Kumar et al.
2016). The ex-type strains are in bold and black. The newly generated
sequence is indicated in bold and blue
Buyck et al. 2012; De Kesel et al. 2016) is the mention - in
the original description - of the unusual pinkish reddening
of its context when exposed, as opposed to the yellowing
context of most other Central African chanterelles with
similar colors. As already previously discussed (Buyck
et al. 2012), we think that this pinkish red context is
probably a secondary and perhaps unimportant or even illinterpreted character. Very similar examples were recently
discussed for two North American species (Buyck et al.
2016b), including C. septentrionalis A.H.Sm., for which
the type description equally mentions a characteristic
pinkish lilac tinge in the context.
Fortunately, however, the microscopic features of C.
goossensiae allow to distinguish it rather easily from other
tropical African chanterelles as it offers the unique combination of comparatively small, ellipsoid spores and a
pileipellis composed of thin-walled, clamped hyphal
extremities terminating frequently in rather short but quite
wide (11–13 lm diam.), obtusely rounded terminal cells
(Heinemann 1958; Buyck et al. 2012) as here indeed
illustrated again for the epitype (Fig. 147c). The spores of
the holotype are clearly smaller than mentioned in the
original description, viz. 5.5–6.72–8 9 3–3.97–5 lm,
Q = 1.37–1.71– 2.0 (see Eyssartier (2001)). The small
spore size is here confirmed by our collections and is a
useful feature to distinguish this species from C. cyanoxanthus (see Buyck et al. (2012)).
During a recent field trip in the Central African rain
forest, not so far from where this species was originally
described, Buyck collected many specimens sharing identical microscopic features with the holotype of C.
goossensiae. One of these included a juvenile specimen
that corresponded exactly to the published watercolor of
the holotype (see Fig. 145a). As a result, we have no doubt
about the correctness of our identification as C. goossensiae and we can finally epitypify this enigmatic species and
assess its phylogenetic affinities and morphological variability using molecular sequencing.
The epitype description is based on the epitype only, but
we have obtained (near-) identical sequences from all of
our collections, (seven sequences have been used to represent the species in our phylogenetic analysis, Fig. 146).
These sequences confirm the impressive field variability of
this species, both in size and general color. Variations in
size concern not only the stipe, which can exceptionally be
more slender and up to 6 cm long, but also the pileus which
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200
Fungal Diversity (2019) 96:1–242
Fig. 143 Molecular Phylogenetic analysis of LSU of Neoacladium
indicum (AMH 10054, holotype) by Maximum likelihood method.
Twenty seven strains are included in the sequence analyses which
comprise 2176 positions after alignment. Auricularia auricular-judae
CBS 7 (JF440710) (Auriculariaceae, Auriculariales) is used as the
outgroup taxon. Bootstrap values for maximum likelihood (ML)
greater than 50 are placed along the branches respectively. The
evolutionary history was inferred by using the Maximum likelihood
method based on the Kimura 2-parameter model (Kimura 1980). The
tree with the highest log likelihood (- 3161.4740) is shown.
Evolutionary analyses were conducted in MEGA7 (Kumar et al.
2016). The ex-type strains are in bold and black. The newly generated
sequence is indicated in bold and blue
reached 15 cm diam. in one collection). Moreover, C.
goossensiae is another example of a species that changes
drastically in general appearance between very young and
more adult stages, exactly as for the closely related and
recently neotypified African C. miniatescens (Buyck et al.
2016a). Very young stages of the latter species are entirely
velutinous and dark wine red, while the adult stages
develop into nearly smooth, bright yellow basidiomata (for
illustrations, see Buyck et al. (2016a)). This is exactly what
happens in C. goossensiae where very young stages are
dark violet and tomentose, but then tend to become smooth
and much paler with age, usually turning brownish to
yellow or even pale cream in one collection. The explanation of this metamorphosis lies in both cases with the
dense fibrillose to tomentose layer that entirely covers the
young cap of both species. This surface layer is of a very
different color compared to the color of the underlying cap
surface which will be gradually exposed as it expands.
Secondly, depending on the intensity of the yellowing of
the context (in which we could not detect any pinkish
tinges), the overall color of the mature pileus varies greatly,
from pale cream over yellow to reddish brown.
For somebody acquainted with European chanterelles,
some specimens of the African C. goossensiae are reminiscent of C. amethysteus in the field because of a similar,
robust stature and identical, violet-purplish tinges on the
pileus surface (see Olariaga et al. (2016)). Notwithstanding
the fact that both species share the omnipresence of clamp
connections in their tissues, they belong in different subgenera (Fig. 146). Indeed, C. goossensiae appears to
belong in the exclusively African subg. Pseudocantharellus. The species is strongly reminiscent of C. miniatescens,
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201
Fig. 144 Cantharellus goossensiae. a Young specimen (Buyck
16.087) showing the typical dark violet pileus and short, fleshy,
yellowish stipe as illustrated (Heinemann 1959) for the holotype.
b Mature rusty brown discolored form (Buyck 16.080). c Mature
yellow discolored form (Buyck 16.096). d Detail of the hairy pileus
covering in young specimens (Buyck 16.087). e Epitype (Buyck
16.064). f typical, amethysteus-like form (Buyck 10.063). g detail of a
mature specimen (Buyck 16.127) with a strongly veined hymenophore. Photos B. Buyck. Scale bars: a–g = 1 cm
in particular because both share a strongly veined-anastomosing hymenophore that is off-white when young.
However, C. goossensiae forms much more robust, fleshy
fruiting bodies that lack the long and slender stipe and the
very thin pileus typical of the latter species. Fully grown
specimens of C. goossensiae are also more variable in color
compared to C. miniatescens as the latter never develops
the brown to olive tinges observed in several collections of
the former (see Fig. 144b, c, e). Some collections of C.
goossensiae developed a typical orange tinge in its upper
part of the stipe, something that appeared to be a good
feature for the correct identification of specimens that lost
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Fig. 145 Cantharellus
goossensiae (epitype).
Microscopic features. a Spores.
b Basidia and basidiola.
c Hyphal extremities of the
pileus surface. Drawings B.
Buyck. Scale bar = 10 lm, but
only 5 lm for spores
their violet pileus color. In addition, C. goossensiae generally lacks the dispersed remains of the red fibrillose
tomentum that are visible near the cap centre or near the
very cap margin of mature or older C. miniatescens (see for
illustrations Buyck et al. (2016a)). The latter species also
has narrower spores and more slender hyphal extremities
composed of more elongated cells at the pileus surface.
123
Cantharellus brunneopallidus Buyck, Randrianjohany &
V. Hofst., sp. nov.
MycoBank number: MB829484; Facesoffungi number:
FoF05997; Figs. 147, 148a–c
Etymology: Referring to the brownish color when
young, fading to yellowish cream, isabelline or dirty
whitish with age.
Holotype: MADAGASCAR. Tampolo, S 17.28724 –
E49.40868, growing on sandy soil between grass and
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amongst dead leaves under Intsia, 4 July 2011, Buyck,
Hofstetter & Randrianjohany leg., Buyck 1053/BB11.105
(PC0085584, holotype).
Fruiting bodies rather small, less than 30 mm high, in
groups of several to many dispersed to aggregated individuals. Pileus less than 30 mm diam., regular in outline,
more rarely lobed or wavy–undulate, first convex, then
becoming progressively more depressed in the centre,
finally often infundibuliform; margin strongly inrolled,
plane or slightly wavy, the marginal zone of the cap
remaining often downturned in age; surface finely fibrillose, slimy when wet, when young often unevenly coloured
in patches of yellowish brown to reddish brown or even
dark brown (5CD6–7,7DEF6–8), later sometimes locally
retaining these colors, elsewhere gradually paler, discoloring to pale cream or isabelline between pale brownish
squamae. Hymenophore decurrent, rather low and mostly
ca 1 mm high, quite abruptly delimited from the sterile
stipe surface, composed of crowded (25–35/cm), frequently
forking and thin gill folds, whitish and with concolorous,
even edges, without anastomosing–veined pattern in
between gills or on gill sides. Stipe shorter than the cap
diam., 9–15(19) 9 3–5(7) mm, subcylindrical or slightly
narrowing downward, slimy when wet, particularly in
lower half also covered with similar, fibrillose–erected to
appressed, pale brownish squamulae as the pileus surface,
off–white, but soon yellowing with age below, sometimes
brownish in lower half when strongly squamulose, not
hollowing. Context white, rather firm, relatively thick
(2–3 mm just outside the pileus centre), strongly bruising
chrome yellow when handled on the whole surface of the
fruiting bodies or in lower tissues from insect damage, but
not entirely becoming yellow. Taste mild. Smell weak.
Spore print not obtained.
Spores ellipsoid, (6.2)6.4–6.68–6.9(7.0) 9 (3.9)4.2–
4.47–4.8(5.0)
lm,
Q = (1.36)1.41–1.50–1.59(1.70),
smooth, hyaline. Basidia small and short, mostly
35–42 9 6–7 lm, usually six–spored. Cystidia none.
Subhymenium more pseudoparenchymatic than filamentous. Pileipellis composed of thin-walled to refringent,
intertwining, septate and mostly slender generative hyphae,
(4)6–9(12) lm diam., with many free endings and occasional ramifications; terminal cell mostly (30)40–65 lm
long, rounded obtuse or slightly tapering at the apex, thinwalled to refringent, but very few clearly thick-walled,
mostly quite regular in outline, sometimes locally undulating or more abruptly inflated. Clamp connections absent
from all tissues.
Additional material examined: MADAGASCAR. Tampolo, S 17.28724 – E49.40868, growing on sandy soil
between grass and amongst dead leaves under Intsia, 6 July
2018, 1054/Buyck 11.116 (PC0085585, paratypus).
203
Notes: The description is based on the holotype. The
paratype has similar features, including a near identical
spore size of (6.0)6.3–6.66–7.0(7.3) 9 (3.9)4.1–4.49–
4.7(4.8) lm, Q = (1.35)1.42–1.52–1.63(1.70). This species
is strongly reminiscent of the tropical African C. densifolius (sensu lato) because of the equally crowded gill folds
and overall color, and it is thus not surprising that it was
originally mistaken for that species when we collected it in
the field. Nevertheless, the very strong yellowing when
bruised is more typical of the Malagasy C. albidolutescens
Buyck, Eyssart. & V. Hofst., but the latter has less crowded
gill folds (see Buyck et al. (2015)). Our phylogeny now
shows that it is clearly an independent species. As we never
have found C. brunneopallidus in other habitats or with
other host trees during the many years of collecting in
Madagascar, it is possible that it might be a strict associate
on the island of the ectomycorrhizal tree genus Intsia
Thouars (Fabaceae), a very small tree genus with an indopacific distribution (Asamoah et al. 2012), but more sites
with Intsia need to be studied in Madagascar to confirm
this. Intsia bijuga (Colebr.) Kuntze, the host species on our
collecting site, is known to have survived in a few
remaining remnants of low elevation humid forests along
Madagascar’s east coast. This tree was always heavily
exploited for its high quality wood. It is listed for many
years already on the IUCN Red List of Threatened Species
and is believed to risk worldwide extinction within the next
20 years without appropriate action to stop illegal logging
(see Intsia bijuga page on Wikipedia for more details).
Cantharellus griseotinctus Buyck, Randrianjohany & V.
Hofst. sp. nov.
MycoBank number: MB829483; Facesoffungi number:
FoF05998; Figs. 148d–f, 149
Etymology: griseo- and –tinctus, referring to the greyish
tinges of the pileus surface.
Holotype: PC0085579
Diagnosis: differs from C. albolutescens in the less
crowded and better developed, less forked gill folds with
distinct rib-like anastomoses on the gill sides, as well as in
the much shorter terminal cells of the hyphal extremities in
the pileipellis.
Fruiting bodies medium-sized, growing single or in
small groups. Pileus relatively fleshy and firm, up to
45 mm diam., first convex, than plano–convex or becoming slightly depressed in the centre, surface a pale brown to
brownish gray (6–7DE4–7) or pale gray to grayish cream
toward the margin, from centre to margin gradually more
areolate–fissured, pileus centre distinctly warty–scurfy,
fading to hardly visible squamae to almost smooth toward
the margin, rarely developing strongly strigose–squamose
rims in concentrical arrangement. Hymenophore decurrent,
composed of well–developed, relatively thick gill folds, up
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b Fig. 146 Most likely phylogram (-ln = 12014.27555) inferred from
combined analyses of nucLSU and TEF1-a for 106 cantharelloid taxa,
using Craterellus tubaeformis as the outgroup taxon. The combined
gene sequence analyses comprise 1840 characters after exclusion of
ambiguously aligned regions (1211 characters for nucLSU, 629
characters for TEF1-a). Phylogenetic analyses were all conducted in
PhyML (Guindon and Gascuel, 2003) under GTR evolutionary model
with the number of substitutions categories, the proportion of
invariable sites and the gamma shape parameters estimated during
the searches. No significant conflict was detected between nucLSU
and TEF1-a based on comparison of bootstrap values recovered from
bootstrap analyses of individual gene (200 bootstrap replicates with
conflict assumed when two different relationships, one being monophyletic and the other being non-monophyletic, for the same set of
taxa were both supported with significant bootstrap values equal or
greater than 70%; Mason-Gamer and Kellog 1996). Searches for the
most likely tree included three independent runs. Model parameters
for the most likely tree were as follows: proportion of invariable
sites = 0.412; gamma shape parameter = 0.409; A = 0.26337,
C = 0.22919, G = 0.27754, T = 0.22990 for base frequencies; A–
C = 1.09799, A–G = 5.55043, A–T = 1.56560, C–G = 1.10930, C–
T = 10.70297, G–T = 1.00000 for substitution rates. The newly
generated sequences are indicated in bold and the taxa discussed in
bold blue. GenBank accessions for nucLSU/TEF1-a follow taxon
names. Branches that received significant support (equal or greater
than 70%) based on 500 BS replicates are in bold and BS values
indicated along the branches
to 4 mm high, unequal and shorter gills of variable length
present, radially undulate –wavy due to rib–like veins
running down the gill sides, off–white to pale cream with
even smooth edges, furcations frequent only near the pileus
margin. Stipe stout, shorter than the pileus diam., less than
20 mm long, 4–7 mm diam., surface white, minutely warty
– fibrillose, not hollowing. Context relatively firm, white,
but near the surface strongly bruising yellow when handled. Taste mild. Smell weak. Spore print not obtained.
Spores rather small, (6.0)6.3–6.64–7.0(7.5) 9 (3.7)
4.0–4.43–4.8(5.2) lm, Q = (1.30)1.39–1.51–1.62(1.72),
ellipsoid, smooth, with a small apiculus. Basidia clavulate,
small to almost medium–sized, 35–50(60) 9 7–8 lm,
mostly six–spored; sterigmata robust. Cystidia none. Subhymenium more pseudoparenchymatic than filamentous.
Pileipellis composed of ramifying, intermingled, strongly
thick–walled hyphal extremities, that are densely septate
(with [ 10 consecutive thick–walled cells) but easily
fragmenting when making preparations, sparsely branching, measuring mostly (4)6–8(10) lm diam.; cells becoming gradually shorter toward the apex, with the terminal
cell rarely longer than 30 lm, tapering and more or less
conical, or also sometimes more ellipsoid and inflated.
Clamp connections absent from all tissues.
Materials examined: MADAGASCAR. East Coast,
Analalava, 7 km West of Mahovelona (Foulpointe),
1742S-4927E, in low altitude humid forest, in small
205
groups among herbs on muddy, stony, steep slope, Buyck
1048/BB 11.054 (PC0085579, holotype !).
Notes: This species strongly resembles C. albidolutescens, (see Buyck et al. (2015)) but it differs from it
because of the less crowded and better developed, less
forked gill folds with distinct rib-like anastomoses on the
gill sides. The more spaced gills also immediately exclude
C. densifolius and look-alikes as a possible identification. It
was collected on the outer limit of one of the few remnants
of low altitude humid forests on the east coast. This forest
remnant near Analalava, not far from Foulpointe, is known
for its high biodiversity and conservation value, which is
well documented in particular for the many rare palm
species that are growing there (Rakotoarinivo et al. 2010).
Hymenochaetales Oberw.
Notes: The order was set up by Frey et al. (1977), based
on Hymenochaetaceae Donk, the most species in the order
have xanthochroic reaction. But recent molecular analyses
showed that more families without xanthochroic reaction
were included in the order (Miettinen and Larsson 2011).
Hymenochaetaceae Donk, Bull. bot. Gdns Buitenz.
Notes: The family Hymenochaetaceae was introduced
by Donk (1948), based on the type genus Hymenochaete
Lév. It is the most important family in Hymenochaetales,
because it has the major species in the order, many species
in the family are medicinal fungi and some are forest
pathogens (Dai et al. 2007, 2009). The xanthochroic
reaction is the most important characteristic of the family.
Currently around 20 genera and 500 species are accepted in
the family (Larsen and Cobb-Poulle 1990; Léger 1998;
Parmasto 2005; Kirk et al. 2008; Dai 2010; Cui et al. 2011;
He and Dai 2012; Ji et al. 2017, 2018; Zhou et al.
2016a, b, c).
Fomitiporia Murrill, N. Amer. Fl.
Notes: Fomitiporia was included in Phellinus Quél.
sensu lato previously, but is characterized by subglobose to
globose, hyaline, thick-walled, strongly dextrinoid and
cyanophilous basidiospores, and accepted as an independent genus (Fiasson & Niemelä 1984; Dai 2010; Amalfi &
Decock 2013; Chen & Cui 2017).
Fomitiporia carpinea X.H. Ji, X.M. Tian & Y.C. Dai, sp.
nov.
Index Fungorum number: IF555462; Facesoffungi
number: FoF04971; Fig. 150
Etymology: Referring to the species growing on
Carpinus.
Holotype: BJFC 025552.
Basidiocarp perennial, resupinate, inseparable, without
odour or taste when fresh, consistency woody hard, light in
weight when dry, up to 10 cm long, 4 cm wide and 7 mm
thick at centre, becoming more or less cushion-shaped with
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206
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Fig. 147 Cantharellus
brunneopallidus (PC0085584,
holotype). Microscopic
features. a Spores. b basidia and
basidiola. c Hyphal extremities
at pileus surface. Drawings B.
Buyck. Scale bar = 10 lm, but
only 5 for spores
age; margin receding, brown, up to 2 mm wide. Pore surface dark brown to bay when fresh, becoming yellowish
brown to umber up on drying, distinctly shining; pores
circular, 4–6 per mm; dissepiments thin and entire.
Subiculum brown, up to 2 mm thick. Tubes concolorous
with pore surface, stratified, up to 5 mm long. Hyphal
system dimitic in all parts; generative hyphae simple septate, skeletal hyphae negative in Melzer’s reagent, acyanophilous in Cotton Blue; tissue darkening but otherwise
unchanged in KOH. Subicular generative hyphae
123
infrequent, hyaline to pale yellowish brown, thin- to
slightly thick-walled, occasionally branched, simple septate, 3–4 lm in diam.; skeletal hyphae yellowish brown,
thick-walled, with a wide lumen, unbranched, regularly
arranged, 3–5 lm in diam. Tramal generative hyphae
hyaline to pale yellow brown, thin- to slightly thick-walled,
occasionally branched, frequently simple septate,
2.5–3.5 lm in diam.; skeletal hyphae yellowish brown,
thick-walled, with a narrow lumen, rarely branched,
interwoven, 3–4.5 lm in diam. Hymenial setae absent;
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207
Fig. 148 Cantharellus brunneopallidus (PC0085584, holotype).
a Freshly collected specimens. b Young specimen showing the dark
pileus. c holotype overview. d–f Cantharellus griseotinctus
(PC0085579, holotype). d showing color change after one day with
respect to the freshly collected specimens depicted in e–f (photos B.
Buyck) Scale bars: 1 cm
cystidioles present, ventricose, hyaline, thin-walled,
10–18 9 3–5 lm; basidia short clavate to capitate, with a
basal
simple
septum
and
four
sterigmata,
12–16 9 8.5–11 lm; basidioles in shape similar to basidia,
but slightly smaller, big irregular or rhombic crystals present
among trama. Basidiospores globose, hyaline, thick-walled,
smooth, dextrinoid in Melzer’s reagent, cyanophilous in
Cotton Blue, (6.1–)6.5–7.8(–8) 9 (5.5–)6–7.5(–7.8) lm,
L = 7.06 lm, W = 6.7 lm, Q = 1.05 (n = 30/1).
Material examined: CHINA, Gansu Province, Pingliang,
Kongtongshan Forest Park, on stump of Carpinus, 2 Sept
2017, Dai 18023 (BJFC 025552, holotype).
GenBank numbers: ITS: MH930812, LSU: MH930810.
Notes: Fomitiporia carpinea is similar to F. punctata (P.
Karst.) Murrill in sharing perennial, resupinate, cushion-
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Fig. 149 Cantharellus
griseotinctus (PC0085579,
holotype). Microscopic
features. a Spores. b basidia and
basidiola. c Hyphal extremities
at pileus surface. Drawings B.
Buyck. Scale bars = 10 lm, but
only 5 for spores
shaped basidiocarps, the approximately the same size
basidiospores, and occurring in temperate forests (Dai
2010, 2012). However, the latter species has smaller pores
(6–8 per mm), interwoven hyphae in subiculum, lacks
cystidioles, while F. carpinea has bigger pore (4–6 per
mm), hyphae regularly arranged in subiculum, the presence
of cystidioles. Phylogentically F. carpinea formed a distinct terminal lineage (Fig. 152), and is distant from F.
punctata.
123
Fomitiporia lagerstroemiae X.H. Ji, X.M. Tian & Y.C.
Dai, sp. nov.
Index Fungorum number: IF555461; Facesoffungi
number: FoF04972; Fig. 151
Etymology: Referring to the species growing on
Lagerstroemiae.
Holotype: BJFC 025858
Basidiocarp perennial, resupinate, inseparable, without
odour or taste when fresh, consistency woody hard, light in
weight when dry, up to 5 cm long, 3 cm wide and 8 mm
Fungal Diversity (2019) 96:1–242
209
Fig. 150 Microscopic structures of Fomitiporia carpinea (BJFC 025552, holotype). a Basidiospores. b Basidia. c basidioles. d Cystidioles.
e Hyphae from trama. f Hyphae from subiculum
thick at centre, becoming more or less cushion-shaped with
age; margin receding, dark brown, up to 2 mm wide. Pore
surface snuff brown when dry, distinctly shining; pores
circular, 7–9 per mm; dissepiments thin and entire.
Subiculum brown, very narrow to almost lacking. Tubes
concolorous with pore surface, stratified, up to 8 mm long.
123
210
Fungal Diversity (2019) 96:1–242
Fig. 151 Microscopic structures
of Fomitiporia lagerstroemiae
(BJFC 025858, holotype).
a Basidiospores. b Basidia.
c Basidioles. d Hymenial setae.
e Hyphae from subiculum
Hyphal system dimitic in all parts; generative hyphae
simple septate, skeletal hyphae negative in Melzer’s
reagent, acyanophilous in Cotton Blue; tissue darkening
but otherwise unchanged in KOH. Subicular generative
hyphae infrequent, hyaline to pale yellowish brown, thinto slightly thick-walled, occasionally branched, simple
septate, 2.5–4 lm in diam.; skeletal hyphae yellowish
brown, thick-walled, with a wide lumen, unbranched,
loosely interwoven, 3–4.5 lm in diam.Tramal generative
123
hyphae hyaline to pale yellow brown, thin- to slightly
thick-walled, occasionally branched, frequently simple
septate, 2.5–3 lm in diam.; skeletal hyphae yellowish
brown, thick-walled, with a narrow to medium size lumen,
rarely branched, interwoven, 3–4 lm in diam. Hymenial
setae frequent, ventricose, thick-walled, dark brown,
15–22 9 5–7 lm; cystidioles absent, basidia barrelshaped, with a basal simple septum and four sterigmata,
9.5–11.5 9 6.5–9 lm; basidioles in shape similar to
Fungal Diversity (2019) 96:1–242
211
basidia, but slightly smaller. Basidiospores subglobose,
hyaline, thick-walled, smooth, dextrinoid in Melzer’s
reagent, cyanophilous in Cotton Blue, (4.8–)5–6(–6.2) 9
(4–)4.5–5.5(–6)
lm,
L = 5.86 lm,
W = 5.13 lm,
Q = 1.14 (n = 30/1).
Material examined: VIETNAM, Dong Nai Province,
Dinh Quan District, Thac Mai Preservation Park, on stump
of Lagerstroemia, 14 Oct 2017, Dai 18335 (BJFC 025858,
holotype).
GenBank numbers: ITS: MH930812, LSU: MH930810.
Notes: Fomitiporia lagerstroemiae is similar to Fomitiporia hainaniana B.K. Cui & Hong Chen in sharing
perennial, resupinate, basidiocarps, the approximately the
same size pores, the presence of hymenial setae, interwoven tramal hyphae and occurring in tropical forests (Chen
and Cui 2017). However, Fomitiporia hainaniana has
smaller basidiospores (4–5 9 3.8–4.4 lm), and grows on
angiosperm wood rather than Lagerstroemia (Chen and Cui
2017). The phylogentic analysis of ITS and LSU sequence
(Fig. 152) shows that Fomitiporia lagerstroemiae is closely related to Fomitiporia bannaensis Y.C. Dai, and both
species has perennial, resupinate, basidiocarps, the
approximately the same size pores and hymenial setae.
However,
Fomitiporia
bannaensis
has
smaller
Fig. 152 Phylogenetic position of Fomitiporia carpinea (holotype)
and Fomitiporia lagerstroemiae (holotype) inferred from the ITS and
LSU sequences. Bootstrap support values for ML and MP greater than
50% and Bayesian posterior probabilities greater than 0.95 are given
near nodes respectively. The tree is rooted with Phellinus uncisetus
(MUCL 46231) and P. uncisetus (MUCL 47061). The new isolates
are in bold and black
123
212
basidiospores
(4.2–5.2 9 3.8–4.9 lm,
L = 4.68 lm,
W = 4.24 lm), the presence of cystidioles (Dai 2010).
Polyporales Gäum.
Notes: Polyporales is a large group of Agaricomycetes
with about 1800 described species (Kirk et al. 2008; Justo
et al. 2017). This order includes species with a wide variety
of basidiomata (pileate, stipitate, resupinate or multiple
flabelliform lobes) and hymenophore (poroid, hydnoid,
lamellate, merulioid or smooth) shapes (Binder et al. 2013;
Hibbett et al. 2014). Most of the species are saprotrophic
wood-decay fungi, while few are plant pathogens. Polyporales has been the focus of many phylogenetic studies
which have confirmed it as a strongly supported monophyletic clade of Agaricomycetes (Binder et al. 2013;
Hibbett et al. 2014; Justo et al. 2017).
Polyporaceae Fr. ex Corda
Notes: Polyporaceae (Polyporales) was proposed by
Fries (1838) with Polyporus P. Micheli as the type genus.
The family currently includes about 45 genera (Justo et al.
2017). Phylogenetic analyses have shown that Polyporaceae belongs to the ‘‘core polyporoid’’ within Polyporales
(Binder et al. 2013). Species in the family have poroid,
irregular or lamellate, occasionally corticioid hymenophore. The hyphal system is rarely monomitic, mostly
dimitic or trimitic, and cystidia are generally lacking. The
species are saprobes and cause white rot (Cannon and Kirk
2007; Justo et al. 2017).
Grammothele Berk. & M.A. Curtis
Notes: Grammothele, typified by G. lineata Berk. &
M.A. Curtis (Berkeley and Curtis 1869), is a genus with
about 22 species (www.indexfungorum.org), with thin and
Fungal Diversity (2019) 96:1–242
strongly adherent basidiomata, and shallow pores (Ryvarden 2015). The hyphal system is dimitic to trimitic, with
clamped generative hyphae, skeletal hyphae thick-walled
to solid, hyaline, darkening with age, dextrinoid or not.
Dendrohyphidia and others sterile elements are absent or
present and the basidiospores are ellipsoid to cylindrical,
thin-walled, smooth and non-amyloid. This tropical genus
occurs on mono- and dicotyledons (Karasinsk 2015;
Ryvarden 2015; Wu et al. 2016).
Grammothele aurantiaca A.M.S. Soares, sp. nov.
Index Fungorum number: IF555479; Facesoffungi
number: FoF05974; Fig. 153
Etymology: aurantiacus L. = orange, referring to the
colour of the basidiomata.
Holotype: MG 232015
Basidiomata annual, resupinate, strongly adnate, hard
and brittle when dried, up to 8 cm long, 4.5 cm wide,
1 mm thick; pore surface salmon to orange (5C5),
becoming dark orange with the age (5B8); margin narrow
to almost lacking, rust orange (5C4). Pores shallow, almost
invisible to the naked eye, round to angular, 6–8 per mm,
dissepiments thin, tubes concolorous, up to 1 mm deep.
Basidiospores scarce, subglobose, hyaline, thin-walled,
with tiny or without apiculus, smooth, IKI-, 4–5 lm wide
(L = 4.66 lm, W = 4.33 lm, Q = 1.07, n = 10/2). Basidia
not seen. Hyphal system dimitic, generative hyphae thickwalled, up to 3 lm wide, without clamps, IKI-; skeletal
hyphae thick-walled 4–5 lm, tightly interwoven, cherry
red in KOH, IKI-. Dendrohyphidia absent. Cystidia and
others sterile elements absents.
Material examined: BRAZIL, Amapá: Serra do Navio,
Serra dos Veados, on dead wood, 00 550 39,9600 N and 51
Fig. 153 Grammothele aurantiaca (MG 232015, holotype). a Basidioma. b Pores. c Hyphae from the sub-hymenium in KOH d basidiospores.
Photos: A.M.S. Soares; Drawing: R.L. Alvarenga. Scale bars: a = 1 cm, b, c = 10 lm, d = 5 lm
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590 20,7900 W, October 2014, W. Xavier, WX 2014-115,
(MG 232015, holotype; isotype in URM).
Additional specimens examined: Grammothele aurantiaca – BRAZIL, Amapá: Serra do Navio, Serra dos Veados,
on dead wood, February 2015, W. Xavier, WX 2015-88
(MG 226148). Porogramme lateritia – COSTA RICA,
Alajuela: Bijagua, Parque Nacional Volcan A renal La
Fortuna de San Carlos, July 2001, L. Ryvarden, 43 (O
F18637). DOMINICAN REPUBLIC: November 1892,
Elliot (O F903299). FRANCE, Guadeloupe: Camp Jacob,
sur un tronc d’un Synplocus maritimiersii, Duss 592 (O
F505994, holotype); Basse terre: October 1976, A. David,
2189 LY-AD (O F505992). VENEZUELA, Arugua: dead
hardwood, February 2006, L. Ryvarden (O F506199).
GenBank numbers: LSU: MH844886, ITS: MH842137.
Notes: Grammothele aurantiaca is characterized by the
salmon to dark orange basidiomata, dimitic hyphal system,
generative hyphae simple-septate, and skeletal hyphae
cherry red in KOH. Porogramme lateritia (Pat.) Pat. has
similar colour, however the pores are smaller [10–15(–20
per mm)], the hyphal system is monomitic with clamps and
the basidiomata become greyish to reddish when dried
(Lowe 1964; Ryvarden 1983). Besides, in the phylogenetic
tree (Fig. 156), G. aurantiaca clustered in the Grammothele clade with good support (1.00/89%) and is distantly
213
related to the Porogramme clade, with P. albocincta
(Cooke & Massee) Gibertoni (1.00/95%) (Fig. 156) as the
type species of the genus.
Grammothele micropora A.M.S. Soares & W.K.S. Xavier
sp. nov.
Index Fungorum number: IF555480; Facesoffungi
number: FoF05975; Fig. 154
Etymology: mikrós Gr. = small ? porus L. = pore,
referring to the tiny pores.
Holotype: MG 232012
Basidiomata annual, resupinate, effuse, strongly adnate,
hard and brittle when fresh and when dried, up to 15 cm
long, 8 cm wide, 1 mm thick; pore surface black when
fresh, dark-bluish when dried (1F1); margin narrow, concolorous with the pore surface becoming slightly whitish
with the age. Pores shallow, invisible to the naked eye,
irregular, round, sometimes angular to hexagonal, 25–30
per mm, dissepiments thin, tubes concolorous, up to 1 mm
deep. Basidiospores subglobose, hyaline, thick-walled,
smooth, with tiny or without apiculus, IKI-, (3.0–) 3.5–4(–
4.5) 9 3–3.5 (–4) lm (L = 3.81 lm, W = 3.14 lm,
Q = 1.21,
n = 15/1).
Basidia
narrowly
clavate,
7–10 9 3–5 lm, with four sterigmata, 2.5–3 lm long.
Hyphal system dimitic, generative hyphae, thin-walled,
Fig. 154 Grammothele micropora (MG 232012, holotype). a Basidiomata. b Pores. c Hyphae from the sub-hymenium. d1 Cystidioles. d2
Basidia. e Basidiospores. Photos: A.M.S. Soares; Drawings: R.L. Alvarenga. Scale bars: a = 1 cm, b, c = 10 lm, d, e = 5 lm
123
214
hyaline, clamped, 2–3 lm wide, IKI-; skeletal hyphae
dominating, thick-walled, brown to black in KOH, 5–7 lm
wide IKI-. Dendrohyphidia absent. Cystidia absent, but
fusoid cystidioles present, 6–9 9 4–5 lm.
Material examined: BRAZIL, Amapá: Serra do Navio,
0 550 9.9500 N and 51 590 19.800 W, on dead wood, October
2014. W. Xavier, WX 2014-116 (MG 232012, holotype;
isotype in URM).
Additional specimens examined: BRAZIL, Amapá:
Porto Grande, Floresta Nacional do Amapá, July 2009, H.
Sotão, H2009-256A (MG 232558). BRAZIL, Amapá:
Porto Grande, Floresta Nacional do Amapá, September
2011, A.M.S. Soares, M146 (MG 232692).
GenBank numbers: ITS: MH842144, LSU: MH842144.
Notes: the dark bluish to black basidiomata, the small
pores and basidiospores, among the smallest in the genus,
are characteristic of this species. Initially, G. micropora
was identified as Porogramme albocincta (Cooke & Massee) Gibertoni because of its dark bluish grey basidiomata
and very small pores (8–20 per mm). Nevertheless, G. micropora has even smaller pores and dimitic hyphal system,
whereas P. albocincta is monomitic and its basidiospores
are longer [4–6 (6.5) 9 3–3.5 lm] (Ryvarden 1979).
Grammothele boliviana Karasiński has similar colour (grey
to bluish grey), however the basidiospores are broadly
ellipsoid to ovate and larger [8.4–10.8(–11.5) 9 6–7.2(–
7.5) lm]. Differently from the new species, G. boliviana
has hyphidia and dendrohyphidia and is reported on palms
Fungal Diversity (2019) 96:1–242
(Karasinsk 2015). Additionally, in the phylogenetic tree, G.
micropora clustered within the Grammothele clade (1.00/
76%) and grouped with low to moderate support with
G. brasiliensis Ryvarden (0.95/70%) (Fig. 156). The type
species of the G. brasiliensis is from Brazil’s southeast
(state of São Paulo) and it is characterized by the dark grey
basidiomata, pores 5–6 per mm and cylindrical basidiospores (5–6 9 2.5–3 lm) (Ryvarden 2015).
Grammothele brasiliensis Ryvarden [as ‘brasilensis’], Syn.
Fung. (Oslo) 33: 38 (2015)
Facesoffungi number: FoF05976; Fig. 155
Description: Ryvarden (2015).
Material examined: BRAZIL, Amapá: Serra do Navio,
on dead wood, October 2014, W. Xavier, WX2014-87
(MG232006), WX2014-132 (MG232007), WX2014-100
(MG232008), WX2014-28 (MG232009), WX2014-101
(MG232010); WX2014-148 (MG232011). Grammothele
subargentea – BRAZIL, Amapá: Serra do Navio, on dead
wood, October 2014, W. Xavier, WX2014-26
(MG232013), WX2014-155 (MG232014). Grammothele
lineata – BRAZIL, Amapá: Serra do Navio, on dead wood,
October 2014, W. Xavier, WX2014-208 (MG232015),
WX2014-40 (MG232016).
GenBank numbers: ITS: MH844864, MH844550,
MH844679, MH844866, MH844553, MH844563,
MH842150, LSU: MH844863, MH844887, MH844583,
MH844865, MH844553, MH842149.
Fig. 155 Grammothele brasiliensis (MG 232007, new record). a Basidioma. b Pores. c Encrusted hyphae from the sub-hymenium. d Basidia.
e Basidiospores. Photos: A.M.S. Soares; Drawings: R.L. Alvarenga. Scale bars: a = 2 cm, b = 0.2 mm, c = 10 lm, d, e = 5 lm
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Fungal Diversity (2019) 96:1–242
Notes: This species is characterized by the dark grey
basidiomata, round and entire pores (5–6 per mm), dimitic
hyphal system, clamped generative hyphae, skeletal
hyphae strongly agglutinated with numerous crystals and
cylindrical basidiospores (5–6 9 2.5-3 lm) (Ryvarden
2015). Dendrohyphidia were easily identified on the studied material, which were not found by Ryvarden (2015).
He, however, presumed their presence in the dissepiments.
This species may be confused with Tinctoporellus epimiltinus (Berk. & Br.) Ryvarden, which has similar colour
(bluish white and light beige), pores (angular to round, 7–9
per mm), and dimitic hyphal system. However, the ellipsoid to subglobose basidiospores [4–5 (5.5) 9 2.5–3 lm],
absence of dendrohyphidia and cystidioles separate it from
G. brasiliensis (Ryvarden 1979). Besides, G. brasiliensis
did not group in the Tinctoporellus clade (1.00/95%), but
formed a monophyletic lineage with strong support (1.00/
100%) within the Grammothele clade (Fig. 156). This is
the first time this species is sequenced, the second record of
this species for Brazil and the first to the Brazilian
Amazonia.
The colour of G. subargentea (Speg.) Rajchenb.
resembles G. brasiliensis, but the former has larger pores
and longer basidiospores [(2–3 per mm; cylindrical (6–)
6.5–8.5 9 (–2.5) 3–4 lm)]. The type species of G. subargentea is from Paraguay, and the morphological characters
of the specimens analyzed in this study are very similar to
the original description (Ryvarden 2015). Although there is
no sequence of the type locality, our phylogenetic analyses
confirm this species in the Grammothele clade (0.91/69%).
Another species registered in Brazil, G. lineata Berk. &
M.A. Curtis, has also white to greyish basidiomata, later
pinkish, and might be mistaken for G. brasiliensis; however, the former has trimitic hyphal system and a partly
hydnoid surface with dots of numerous dark bundles of
skeletal hyphae (Ryvarden 2015). In addition, the specimens collected in the Brazilian Amazonia sistered with G.
lineata from China and Malaysia, and Grammothele sp.
and G. denticulata Y.C. Dai & L.W. Zhou (1.00/99%) from
China, in the Grammothele clade. Grammothele lineata
was originally collected in Cuba and there is no sequence
from the type locality. The new sequences generated from
Brazilian material may represent G. lineata due the
closeness to the type locality, since both are in the
Neotropics. Thus, the Asian collections should be reevaluated.
Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C.
David
Notes: The concept of Russulales was gradually developed in morphological analyses for several groups of
macro-fungi. Singer and Smith (1960) have noticed the
close morphological relationship between agaricoid and
215
gasteroid homobasidiomycetes taxa. The amyloid basidiospore ornamentations and gloeoplerous hyphae system
reminded the Dutch mycologist Marinus Anton Donk of
some families (e.g. Hericiaceae) may be members of an
unknown order (Donk 1971). Oberwinkler (1977) regarded
this order as Russulales which include agaricoid, discoid,
fused-reflexed, gasteroid, hydnoid, and resupinate taxa.
The order Russulales was officially acknowledged in the
Dictionary of Fungi (Kirk et al. 2001; Miller et al. 2006).
The independence of this order has been supported by
some recent phylogenetic analyses (Hibbett et al. 2007;
Zhao et al. 2017).
Russulaceae Lotsy
Notes: The family Russulaceae was established by
Dutch botanist Johannes Paulus Lotsy to accommodate
species with granular flesh, thick gills, spiny spores, and
milky hyphae and rounded cells (sphaerocytes). This
family includes gasteroid and agaricoid genera, Lactarius
Pers., Lactifluus (Pers.) Roussel, Multifurca Buyck & V.
Hofst. and Russula Pers., together with the crust-like genera Boidinia Stalpers & Hjortstam, Gloeopeniophorella
Rick, and Pseudoxenasma K.H. Larss. & Hjortstam (Lotsy
1907; Larsson and Larsson 2003; Miller et al. 2006;
Larsson 2007; Buyck et al. 2008, 2010; Verbeken and
Nuytinck 2013). The family comprises ectomycorrhizal
(Tedersoo and Nara 2010), arbutoid mycorrhizal (Smith
and Read 2008), monotropoid mycorrhizal (Bidartondo
2005), orchid mycorrhizal (Dearnaley 2007), and saprotrophic, wood-degrading species (Larsson and Larsson
2003).
Russula Pers.
Notes: The genus Russula was introduced by Persoon
(1796) with the type species R. emetica (Schaeff.) Pers.
This genus is characterized by fairly large basidiocarps,
brightly colored pileus, white to dark yellow basidiospore
print, attached lamellae, absence of volva and milky latex,
amyloid basidiospore ornamentations, and numerous
spherocysts in context (Sarnari 1998). Over 800 species are
accommodated in this genus (Li 2014; Li et al.
2015, 2018c). Most recent new Russula species were
described based on specimens from Asia (Li et al.
2016, 2018a, b; Sang et al. 2016; Das et al.
2017a, b, 2018b; Hyde et al. 2016, 2017b; Jiang et al. 2017;
Zhang et al. 2017). In this paper, we introduce a new
species in the genus Russula which was collected in
coniferous and broad-leaved intermixed forests from
Guangxi of South China.
Russula prasina G.J. Li & R.L. Zhao, sp. nov.
Index Fungorum number: IF570591; Facesoffungi
number: FoF05086; Fig. 157
123
216
Fig. 156 Maximum likelihood consensus phylogenetic tree of the
concatenated ITS1, 5.8S, ITS2 and partial 28S rDNA sequences.
Bootstrap supporting values (1000 replicates) and posterior probabilities (PP) from Bayesian analysis to each node are shown from left to
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Fungal Diversity (2019) 96:1–242
right. Only bootstrap values above 50% and BYPP above 0.75 are
provided. The newly generated sequences are indicated in bold and
black. The tree was rooted with Funalia trogii (XSD37)
Fungal Diversity (2019) 96:1–242
217
Fig. 157 Russula prasina (HMAS 281232, holotype). a, b Basidiocarps. c Basidiospores. d Basidia. e Subpellis. f Epipellis. Scale bars: a,
b = 5 cm, f = 20 lm, d, e = 10 lm
Etymology: Refers to the grass green tinged pileus of
this species.
Holotype: HMAS 281232
Basidiocarps small to large sized. Pileus 43–130 mm in
diam., first hemispheric, then plane to depressed at centre
when mature, slightly striate 25–30 mm towards the edge
when mature, not cracked, slightly viscid when wet, dull,
peeling 1/5–1/4 from the edge, grass green tinge of Winter
Green (XVIII330 i), Motmot Green (XX430 i) to Courge
Green (XVII250 i) at centre, pale yellowish green tinge of
Oil Yellow (V27 k), Javel Green (V27i) to Calliste Green
(VI31i) towards margin. Lamellae adnate, 3–5 mm in
height, 8–13 pieces per centimeter at edge, rarely forked
near stipe, often interveined, pale ocher tinge of Ochraceous-Buff (XV150 b), unchanging when bruised, lamellulae absent. Stipes central to subcentral, 4.1–7.7 9
2.4–3.6 cm, subcylindrical to cylindrical, surface dry,
rugulose longitudinally, dull, without annulus, slightly
attenuate downward, White (LIII) at upper part, ochraceous
yellow tinge of Light Ochraceous Salmon (XV130 b) to
Salmon Buff (XIV110 d) towards the base, stuffed at first,
becoming hollow when old. Context up to 3–5 mm at
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218
Fungal Diversity (2019) 96:1–242
Fig. 158 Phylogram generated from Bayesian analysis of ITS
sequence data of Russula subgenus Heterophyllidia. Related
sequences were obtained from Miller and Buyck (2002), Abarenkov
et al. (2010), Li et al. (2013), Guo et al. (2014), Dutta et al. (2015),
Zhao et al. (2015) and Zhang et al. (2017). Russula aurea and R.
paludosa are used as the outgroup taxa. Bootstrap (BS) support values
for ML equal to or greater than 65% and Bayesian posterior
probability (BYPP) values equal to or greater than 0.90 are given
above the nodes respectively as (BS/BYPP). Phylogenetic topology of
the ML analysis was similar to that of the BI. The type specimens are
in bold and black. The newly generated sequences are in blue
centre of pileus, White (LIII) first, unchanging when
bruised, fragile, no distinct odor; taste mild. Spore print
Ocher (Romagnesi III b–c).
Basidiospores [300/3/3] (5.9–) 6.2–7.3 (–7.6) 9 (5–)
5.4–6.5
(–7)
lm,
Q = 1.01–1.21
(–1.25)
(Q = 1.12 ± 0.07), hyaline, globose to subglobose, rarely
broadly ellipsoid; ornamentation composed of amyloid
obtuse warts and short crests that are mostly interconnected
as short crests and ridges, forming a complete network,
warts 0.3–0.6 lm high; suprahilar area nonamyloid to
rarely weakly amyloid. Basidia 35–53 9 8–11 lm, projecting 10–20 lm beyond hymenium, four-spored, sterigmata 4–6 lm long, hyaline, unchanging in KOH,
subclavate to clavate, rarely cylindrical. Pleurocystidia and
cheilocystidia not observed. Subhymenium cellular layer
15–30 lm thick composed of voluminous cells 10–20 lm
in diam., hyaline, rarely pale yellowish in KOH. Pileipellis
is composed of epipellis and subpellis; epipellis pseudoparenchymatous, 100–250 lm thick, composed of thinwalled, unbranched and inflated cells 10–20 lm wide,
narrowing to 5–7 lm towards the terminal cells;
pileocystidia absent; subpellis slightly gelified, a cutis
150–250 lm thick, composed of parallel to interweaved,
rarely branched and septate, hyaline hyphae 2–4 lm wide.
Stipitipellis cutis, upper layer composed of parallel filamentous hyphae 3–6 lm in diam., inner layer hyaline filamentous hyphae interweaved with sphaerocytes
15–25 lm in diam., some filamentous hyphae pale yellow
in KOH; caulocystidia not observed. Trama composed of
sphaerocytes 15–40 lm in diam. and scattered by filamentous and cystidioid hyphae. Clamp connections and
laticiferous hyphae absent in all tissues.
Material examined: CHINA, Guangxi Zhuang Autonomous Region, Baise City, Leye County, Huangjingdong
National Forest Park, N 24740 E 106320 , elv. 378 m., 6
August 2017, Hui-Jun Wang GX20170580 (HMAS
281232, holotype); ibid. elv. 464 m., GX20170846
(HMAS 279805); ibid. elv. 395 m., GX20170937 (HMAS
279806).
Habit and habitat: Scattered or single in coniferous and
broad-leaved intermixed forests (dominated by e.g. Pinus
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219
Fig. 159 Rhizophydium koreanum (CNUFC-17CPW1-1, holotype).
a Colony on PmTG agar. b Developing zoosporangium with branched
rhizoids. c Resting spores. d Developing thallus with two rhizoidal
axes (white arrows). e Uniflagellate zoospores (yellow arrows). Scale
bars: b–e = 20 lm
crassicorticea, P. yunnanensis var. tenuifolia, Quercus
acutissima, and Q. variabilis) at 300–500 m altitude.
Distribution: CHINA (Guangxi Zhuang Autonomous
Region). Season. August.
GenBank numbers: ITS: MH454351 (HMAS 281232),
MH454352 (HMAS 279805), MH454353 (HMAS
279806).
Notes: Russula prasina is a member of subgenus
Heterophyllidia, section Virescentinae because it has
greenish tinged pileus, small basidiospores, spherocysts in
epipellis, and absence of pileocystidiain pileipellis (Sarnari
1998). Unlike the furfuraceous and areolate pileus surfaces
of the other Virescentinae members, that of R. prasina is
relatively smooth. The hyphal cells in subpellis and
epipellis of R. prasina are reminiscent of those of R. parvovirescens Buyck, D. Mitch. & Parrent, in contrast,
R. parvovirescens has pileocystidia in pileipellis and
hymenium with pleurocystidia and cheilocystidia (Buyck
et al. 2006). This new species may be confused with some
green-capped Russula in China and adjacent region, e.g. R.
atroaeruginea G.J. Li, Q. Zhao & H.A. Wen, R.
dinghuensis J.B. Zhang & L.H. Qiu, R. nigrovirens Q.
Zhao, Y.K. Li & J.F. Liang, R. sikkimensis K. Das, Atri &
Buyck, R. virescens (Schaeff.) Fr., and R. viridirubrolimbata J.Z. Ying. These species can be distinguished from R.
prasina by the following morphological characteristics: R.
atroaeruginea differs in its radially yellowish-striped
pileus, larger basidiospores (6.3–) 6.8–8.1 (–9.0) 9 (5.9–)
6.1–7.4 (–7.8) lm, and habitat of in alpine subtropical
conifer forests dominated by Picea spp. (Li et al. 2013); R.
dinghuensis has pileus surface cracking into small patches
with rusty tinge, scattered lamellulae, white basidiospore
sprint, and trichoderm epipellis (Zhang et al. 2017); R.
nigrovirens can be differentiated by its longer basidia
45–75 9 9–14 lm, basidiospore ornamentation not forming a reticulum, and habitat of in alpine subtropical conifer
forests dominated by Picea sp., Rhododendron sp., Sorbus
sp. and Abies sp. (Zhao et al. 2015); R. sikkimensis has
basidiospore ornamentation up to 0.9 lm, cutis to trichoderm pileipellis and habitat of in alpine subtropical conifer
forests dominated by Abies spp. (Das et al. 2013); R. virescens differs in its white to pale cream spore print,
123
220
Fungal Diversity (2019) 96:1–242
Fig. 160 Phylogenetic tree of Rhizophydium koreanum (CNUFC17CPW1-1) and (CNUFC-17CPW1-2) and related species based on
Maximum likelihood analysis of 28S. Sequence of Allomyces
arbusculus was used as outgroup taxon. Numbers at the nodes
indicate the bootstrap values (C 50%) from 1000 replications. The
bar indicates the number of substitutions per position. The newly
generated sequences are indicated in blue and ex-type strain in bold
elliptical to subglobose basidiospores, and frequently
elongated and tapered terminal cells in pileipellis (Sarnari
1998, Li 2014); R. viridirubrolimbata can be differentiated
by its red tinge in pileus, and acrid context (Ying 1983).
Phylogenetic analysis showed that our strains formed a
distinct clade from known Russula species (Fig. 158). Thus
Russula prasina is introduced as a new species based on
morphological and phylogenetic analyses.
Rhizophydium Schenk
Notes: The genus Rhizophydium is the oldest genus
within the order Rhizophydiales, proposed by Schenk
(1858) and validated by Rabenhorst (1868). The species
belonging to this genus are characterized by the formation
of sphaerical zoosporangia with multiple, discharge pores,
and branched rhizoids arising from the sporangium (Barr
1968; Letcher et al. 2006). Members of the genus occur in
aquatic systems, primarily as parasites of algae, and from
pollen, keratin, and soil as saprotrophs (Barr 1968; Longcore 1996; Letcher et al. 2004, 2006). In recent years,
ultrastructural and molecular analyses have been used as a
tool to delimit species in the genus, resulting in significant
change in the taxonomy (Letcher et al. 2006; Seto et al.
2017). Many Rhizophydium species have been transferred
to new genera (Letcher et al. 2006, 2015).
During a study of Rhizophydiales in water samples in
Gwangju, Korea, a new species of Rhizophydium was
Chytridiomycota Arx
Rhizophydiomycetes Tedersoo, Koljalg et al.
Rhizophydiales Letcher
Notes: Currently, 17 families and 25 genera are placed
in order Rhizophydiales based on molecular phylogeny and
zoospore ultrastructure (Letcher et al. 2015; Van den
Wyngaert et al. 2017; Seto and Degawa 2018;
Wijayawardene et al. 2018b).
123
Fungal Diversity (2019) 96:1–242
isolated and is described here, based on morphological
characteristics and phylogenetic analyses.
Rhizophydium koreanum Hyang B. Lee, S.J. Jeon, T.T.
T. Nguyen, sp. nov.
Index Fungorum number: IF554571; Facesoffungi
number: FoF05792; Fig. 159
Etymology: koreanum, referring to the country from
which the species was recorded.
Holotype: CNUFC-17CPW1-1
On PmTG (peptonized milk, tryptone and glucose) agar:
Zoosporangium sphaerical, with many closely spaced and
highly branched rhizoids, measured (100–)107.5–120.5(–
131.5) lm diam. Rhizoidal system arising from a single
point at the base of the sporangium. Zoospores abundantly
produced, sphaerical, measured (3.0–)3.5(–4.0) lm diam.,
with a flagellum of (20–)23.5(–30) lm long. Resting spore
sphaerical, measured (10–)12.5(–13.5) lm diam.
Culture characteristics: Colonies reaching 8.5 mm
diam. on PmTG at 25 C in 7 days, cream. Optimal growth
was observed around 20 C.
Material examined: REPUBLIC OF KOREA, from
pond water collected at Chonnam National University
Arboretum, Gwangju, Korea, 20 October 2017 (CNUFC17CPW1-1, preserved as glycerol stock at - 80 C in the
Chonnam National University Fungal Collection; isotype
in Culture Collection of Nakdonggang National Institute of
Biological Resources [NNIBR], Sangju, Gyeongbuk
Province).
Notes: Based on phylogenetic analyses and morphological comparison, our isolate belongs to Rhizophydium.
Rhizophydium koreanum formed a distinct clade from other
species in the phylogenetic tree (Fig. 160). It differs from
the closely related species R. globosum and R. brooksianum
by forming larger sporangia.
GenBank numbers: LSU: MH298649, MH298650.
Acknowledgements Kevin D. Hyde thanks the Foreign Experts Bureau
of Yunnan Province, Foreign Talents Program (2018; Grant No.
YNZ2018002), Thailand Research grants entitled Biodiversity, phylogeny and role of fungal endophytes on above parts of Rhizophora
apiculata and Nypa fruticans (Grant No. RSA5980068), the future of
specialist fungi in a changing climate: baseline data for generalist and
specialist fungi associated with ants, Rhododendron species and Dracaena species (Grant No. DBG6080013), Impact of climate change on
fungal diversity and biogeography in the Greater Mekong Subregion
(Grant No. RDG6130001). K.D. Hyde also thanks Chiang Mai
University for the award of visiting Professor. We also thank to the
director Jun-Bo Yang and Plant Germplasm and Genomics Center in
Germplasm Bank of Wild Species, Kunming Institute of Botany for the
molecular laboratory support. Dr. Shaun Pennycook is thanked for his
essential nomenclatural review. Samantha C. Karunarathna thanks
CAS President’s International Fellowship Initiative (PIFI) for funding
his postdoctoral research (Grant No. 2018PC0006) and the National
Science Foundation of China (NSFC, project code 31750110478).
Department of Plant Medicine, National Chiayi University (NCYU) is
thanked to provide facilities for DNA molecular experiment. Rajesh
221
Jeewon thanks Mae Fah Luang University for the award of a Visiting
Scholar. Walter Rossi and Marco Leonardi wish to thank the following
entomologists who supplied them with the flies bearing Labouobeniales
and/or identified the host insects: R. Andrade (Portugal), G. Cocks
(Australia), J. Deeming (Great Britain), Martin J. Ebejer (Great Britain), Marina Krivosheina (Russia), Iain MacGowan (Great Britain),
Thomas Pape (Denmark), Alfio Raspi (Italy), N. Vikhrev (Russia), M.
von Tschirnhaus (Germany). H.B. Lee was supported by the Graduate
Program for the Undiscovered Taxa of Korea, and in part by the Project
on Survey and Discovery of Indigenous Fungal Species of Korea funded
by NIBR and Project on Discovery of Fungi from Freshwater and
Collection of Fungarium funded by NNIBR of the Ministry of Environment (MOE), and in part carried out with the support of Cooperative
Research Program for Agriculture Science and Technology Development (PJ013744), Rural Development Administration, Republic of
Korea. This work was in part supported by the BK21 plus program
through the National Research Foundation (NRF) funded by the Ministry of Education of Korea. Jian-Kui Liu thanks the National Natural
Science Foundation of China (NSFC 31600032). Marcela Cáceres
thanks the CNPq (Conselho Nacional de Desenvolvimento Cientı́fico e
Tecnológico) for a research grant (309058/2015-5), funding for collecting trips (401186/2014-8), and a collaborative project with RL as
Special Visiting Professor (314570/2014-4). Funding for phylogenetic
work on Graphidaceae was provided by a grant from the National
Science Foundation (NSF) to The Field Museum: DEB-1025861
‘‘ATM – Assembling a taxonomic monograph: The lichen family
Graphidaceae’’; PI Thorsten Lumbsch, CoPI Robert Lücking. Armin
Mangold is thanked for providing two photographs of the type material
of Thelotrema annulatum. Vanessa P. Abreu, André W. C. Rosado and
Olinto L. Pereira thank the CAPES, CNPq, and FAPEMIG for financial
support and ICMBio/FLONA-Paraopeba for providing facilities and
permits for the exploration surveys of the mycodiversity in their protected areas. We would like to thank the Instituto Chico Mendes de
Conservação da Biodiversidade (ICMBio) of the Parque Montanhas do
Tumucumaque and the Instituto Brasileiro de Meio Ambiente
(IBAMA) for support during the field trips; and Msc. RLM Alvarenga
for the figures. We thank the Graduate Program for the Biodiversity and
Biotechnology Network of the Legal Amazon (UFPA-MPEG, Brazil);
the Museu Paraense Emı́lio Goeldi (MPEG), the Universidade do
Estado do Amapa (UEAP) and the Universidade Federal de Pernambuco (UFPE) for the logistical support of its laboratories and herbaria.
Further, we acknowledge the Conselho Nacional de Desenvolvimento
Cientı́fico Programa de Capacitação for the scholarship to AMSS
(Programa de Capacitação Institucional 303073/2018-7); CNPq (Sisbiota 563342/2010-2, PROTAX 562106/2010-3), and FACEPE (APQ
0788-2.03/12) for funding this research. T.G.L. Oliveira, C.M. SouzaMotta, J.D.P. Bezerra, and O.M.C. Magalhães are grateful to the Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq),
Coordenação de Aperfeiçoamento de Pessoal de Nı́vel Superior
(CAPES) and Fundação de Amparo à Ciência e Tecnologia de Pernambuco (FACEPE) for financial support and/or scholarships; to Dr.
Jarcilene S. Almeida Cortez, Dr. Alexandre R. Machado and Greicilene
M. R. Albuquerque for their help during the execution of this study, and
also to the Instituto Fazenda Tamanduá (Tamanduá Farm) for support
during the collections of plant material. PN Singh and Sanjay K Singh
thanks to Director Agharkar Research Institute, Pune, India for providing facilities. The technical help in preparing photo-plates by Mr.
S.B. Gaikwad is acknowledged. Martina Réblová acknowledges support by a long-term research development project No. RVO 67985939
of the Czech Academy of Sciences, Institute of Botany. José Ewerton F.
dos Santos and Renan N. Barbosa acknowledgment scholarship and
financial support from Conselho Nacional de Pesquisa e Desenvolvimento Cientifico (CNPq). We would like to acknowledge the URM
Herbarium and URM Culture Collection staff and Professor Dr. Roger
Fagner R. de Melo for taxonomy assistance. Guo-Jie Li and Rui-Lin
Zhao would like to thank National Natural Science Foundation of China
123
222
(Project IDs GJL: 31500013, RLZ: 31470152 and 31360014) for
financial support. The material of Plectocarpon on S. tricosa was studied
for the Galapagos Lichen Inventory, a joint project of the Charles
Darwin Foundation (CDF) and the Galapagos National Park (DPNG),
part of a national biodiversity assessment ‘‘Biodiversidad Genética del
Ecuador’’ led by the Instituto Nacional de Biodiversidad del Ecuador
(INABIO). The authors are indebted to our colleagues and collaborators
at these institutions, especially Daniel Lara Solı́s, Galo Quedaza and
Victor Carrión (DPNG), Arturo Izurieta and Maria-José Barragan
Paladines (CDF), and Diego Inclán and Rosa Batallas (INABIO) for
research permits and logistical support. Monika Dayarathne would like
to thank Thailand Research Fund (TRF) Grant No. MRG6080089
entitled; Taxonomy and phylogeny of foliar fungi from Mangrove and
to Dr. Putarak Chomnunti. Saranyaphat Boonmee would like to thank
the Thailand Research Fund (No. TRG6180001), the National Research
Council of Thailand (No. 61215320023) and Plant Genetic Conservation Project under the Royal Initiation of Her Royal Highness Princess
Maha Chakri Sirindhorn-Mae Fah Luang University. Armin Mešić and
Zdenko Tkalčec are grateful to Croatian Science Foundation for their
financial support under the project HRZZ-IP-2018-01-1736 (ForFungiDNA) and thank to Milan Čerkez for collecting specimens of
coprinoid taxa. Chaynard Phukhamsakda would like to thank the Royal
Golden Jubilee PhD Program under Thailand Research Fund (RGJ) for
a personal grant to C. Phukhamsakda (The scholarship no. PHD/0020/
2557 to study towards a PhD). Lei Cai acknowledges China-Thailand
Joint Lab on Microbial Biotechnology (Most KY201701011) for
financial support. Rungtiwa Phookamsak thanks CAS President’s
International Fellowship Initiative (PIFI) for young staff (Grant No.
2019FYC0003), the Research Fund from China Postdoctoral Science
Foundation (Grant No. Y71B283261), the Yunnan Provincial Department of Human Resources and Social Security (Grant No.
Y836181261), and National Science Foundation of China (NSFC)
project code 31850410489 for financial support. Putarak Chomnunti
would like to thank the National Research Council of Thailand (Grant
No. 256108A3070006) for financial support. Ting-Chi Wen and
Yuanpin Xiao would like to thank the National Natural Science
Foundation of China (No. 31760014) and the Science and Technology
Foundation of Guizhou Province (No. [2016]2863). This research work
was partially supported by Chiang Mai University. TB Gibertoni, AMS
Soares, HL Plautz-Jr, HMP Sotão and WKS Xavier would like to thank
the Instituto Chico Mendes de Conservação da Biodiversidade of the
Parque Nacional Montanhas do Tumucumaque and the Instituto Brasileiro de Meio Ambiente for support during the field trips; the Graduate
Program for the Biodiversity and Biotechnology Network of the Legal
Amazon (UFPA-MPEG), the Museu Paraense Emı́lio Goeldi (MPEG),
the Universidade do Estado do Amapa and the Universidade Federal de
Pernambuco for the logistical support of their laboratories and herbaria;
CNPq for the scholarship of AMSS (Programa de Capacitação Institucional 303073/2018-7); and CNPq (Sisbiota 563342/2010-2, PROTAX 562106/2010-3) and FACEPE (APQ 0788-2.03/12) for funding
this research; and Msc. RLM Alvarenga for the figures. B. Buyck thanks
the ATM of the Paris’ Museum and ‘‘l’Institut Ecologie et Environnement’’ (CNRS-INEE) for funding the field trip with Shelly Masi to
Africa; Shelly is thanked for all the practical help and sharing her
experience. Terence Fuh and the staff of the Primate Habituation Programme of the Dzanga-Ndoki National Park of the ‘‘Réserve spéciale de
Foret Dense de Dzanga-Sangha’’ at Bayanga, as well as all staff, Aka
guides and visitors of the Bai hakou field station for logistical support,
field assistance and the very enjoyable company during our stay. This
research was made possible through research permit 034/MENESR/
DIRCAB/DGESRSTI/DRSTSPI/SSSTI/16 from the ‘‘Ministère de
l’éducation nationale, de l’enseignement supérieur et de la recherche
scientifique’’ of the Central African Republic. B. Buyck & V. Hofstetter
are grateful to the staff members of CNRE in Antananarivo for logistic
support and field assistance; while field work in Madagascar was
financed in part by the National Geographic Society (grants 6365-98,
123
Fungal Diversity (2019) 96:1–242
7921-05) and in more recent years by the ATM-project ‘‘Past and
present biodiversity’’ of the Muséum national d’histoire naturelle (Dirs.
Ph. Janvier and S. Peigné). R Jeewon thanks University of Mauritius for
research support. Napalai Chaiwan would like to thank the Thailand
Research Fund (PHD60K0147). The material of Plectocarpon on S.
tricosa was studied for the Galapagos Lichen Inventory, a joint project
of the Charles Darwin Foundation (CDF) and the Galapagos National
Park (DPNG), part of a national biodiversity assessment ‘‘Biodiversidad Genética del Ecuador’’ led by the Instituto Nacional de Biodiversidad del Ecuador (INABIO). The authors are indebted to our
colleagues and collaborators at these institutions, especially Danny
Rueda, Daniel Lara Solı́s, Galo Quedaza and Victor Carrión (DPNG),
Arturo Izurieta and Maria-José Barragan Paladines (CDF), and Diego
Inclán and Rosa Batallas (INABIO) for research permits and logistical
support. This publication is contribution number 2248 of the Charles
Darwin Foundation for the Galapagos Islands. Danushka S. Tennakoon
would like to thank Lakmali Dissanayake and Binu Samarakoon for
their support. D.N. Wanasinghe would like to thank CAS President’s
International Fellowship Initiative (PIFI) for funding his postdoctoral
research (Number 2019PC0008). Peter E. Mortimer and D.N. Wanasinghe thank the National Science Foundation of China and the Chinese
Academy of Sciences for financial support under the following grants:
41761144055, 41771063 and Y4ZK111B01. Yusufjon Gafforov thanks
CAS President’s International Fellowship Initiative (Grant No.
2018VBB0021) and German Academic Exchange Service Fellowship
(Grant No. 57314018) and Ministry of innovative development of the
Republic of Uzbekistan (Projects No. P3-2014-0830174425 and PP20170921183) for funding his research projects. Qiu-Ju Shang would
like to thank Dr. Julio Mena-Portales, the senior researcher of the
Division of Mycology and president of the Scientific Council, Institute
of Ecology and Systematic, Havana, Cuba, for his kind support on
manuscript writing. PhD students from Mae Fah Laung and Chiang Mai
Universities thank the Mushroom Research Foundation for research
financial support and PhD Fellowships. Mingkwan Doilom would like
to thank the 5th batch of Postdoctoral Orientation Training Personnel in
Yunnan Province (Grant No. Y934283261) and the 64th batch of China
Postdoctoral Science Foundation (Grant No. Y913082271). Ningguo
Liu, Zuo-Yi Liu, Hong-Yan Su, A. Akulov and Huang Zhang are deeply
thanked for their kind support on manuscript writing. Jianchu Xu thanks
Key Research Program of Frontier Sciences ‘‘Response of Asian
mountain ecosystems to global change’’, CAS (Grant No. QYZDYSSW-SMC014). D.N. Wanasinghe thanks the 64th batch of China
Postdoctoral Science Foundation (Grant No. Y913083271). Alan JL
Phillips acknowledges the support from UID/MULTI/04046/2019
Research Unit grant from FCT, Portugal to BioISI.
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Affiliations
Kevin D. Hyde1,2,3,4,5,6,71 • Danushka S. Tennakoon1,2,3,6,7 • Rajesh Jeewon8 • D. Jayarama Bhat9,10 •
Sajeewa S. N. Maharachchikumbura11 • Walter Rossi12 • Marco Leonardi12 • Hyang Burm Lee13 •
Hye Yeon Mun14 • Jos Houbraken15 • Thuong T. T. Nguyen13 • Sun Jeong Jeon13 • Jens Christian Frisvad16 •
Dhanushka N. Wanasinghe1,3,4,71 • Robert Lücking17 • André Aptroot18 • Marcela E. S. Cáceres19 •
Samantha C. Karunarathna1,4,5,71 • Sinang Hongsanan3,20 • Rungtiwa Phookamsak1,3,4,5,71 • Nimali I. de Silva1,3,5 •
Kasun M. Thambugala21 • Ruvishika S. Jayawardena3 • Indunil C. Senanayake3,20 • Saranyaphat Boonmee3 •
Jie Chen22 • Zong-Long Luo23 • Chayanard Phukhamsakda2,3 • Olinto L. Pereira24 • Vanessa P. Abreu25 •
André Wilson Campos Rosado24 • Buyck Bart26 • Emile Randrianjohany27 • Valérie Hofstetter28 •
Tatiana B. Gibertoni29 • Adriene Mayra da Silva Soares30 • Helio Longoni Plautz Jr.31 • Helen Maria Pontes Sotão30 •
William Kalhy Silva Xavier32 • Jadson Diogo Pereira Bezerra33 • Thays Gabrielle Lins de Oliveira33 •
Cristina Maria de Souza-Motta33 • Oliane Maria Correia Magalhães33 • Digvijayini Bundhun3,34 •
Dulanjalee Harishchandra2,3,35 • Ishara S. Manawasinghe2,3,35 • Wei Dong3,6,34,36 • Sheng-Nan Zhang3,34 •
Dan-Feng Bao3,23,34 • Milan C. Samarakoon3,5,37 • Dhandevi Pem2,3,6,20 • Anuruddha Karunarathna1,3,7,34 •
Chuan-Gen Lin2,3,6 • Jing Yang2,3,6,37 • Rekhani H. Perera2,3,6,37 • Vinit Kumar3,34 • Shi-Ke Huang1,2,3,6 •
Monika C. Dayarathne1,2,3,6 • Anusha H. Ekanayaka1,2,3 • Subashini C. Jayasiri1,3 • Yuanpin Xiao2,3,6,38 •
Sirinapa Konta1,2,3,6 • Tuula Niskanen39 • Kare Liimatainen39 • Yu-Cheng Dai40 • Xiao-Hong Ji40 •
Xue-Mei Tian41 • Armin Mešić42 • Sanjay K. Singh43 • Kunthida Phutthacharoen2,3,6 • Lei Cai4 •
Touny Sorvongxay3 • Vinodhini Thiyagaraja1,3,6,34 • Chada Norphanphoun2,3,6,7,38 • Napalai Chaiwan1,2,3,6 •
Yong-Zhong Lu3,6,38 • Hong-Bo Jiang1,2,3,6 • Jin-Feng Zhang3,37 • Pranami D. Abeywickrama2,3,35 •
Janith V. S. Aluthmuhandiram2,3,35 • Rashika S. Brahmanage2,3,35 • Ming Zeng1,2,3,6 • Thilini Chethana2,3,35 •
Deping Wei1,3,34 • Martina Réblová45 • Jacques Fournier46 • Jana Nekvindová47 • Renan do Nascimento Barbosa48 •
José Ewerton Felinto dos Santos33 • Neiva Tinti de Oliveira33 • Guo-Jie Li44 • Damien Ertz49,50 •
Qiu-Ju Shang2,3,37 • Alan J. L. Phillips51 • Chang-Hsin Kuo7 • Erio Camporesi52,53,54 • Timur S. Bulgakov55 •
Saisamorn Lumyong3,5,68,69 • E. B. Gareth Jones3,56 • Putarak Chomnunti2,3 • Eleni Gentekaki2,3 •
Frank Bungartz57,58,59 • Xiang-Yu Zeng3,38 • Sally Fryar60 • Zdenko Tkalčec42 • Junmin Liang44 •
Guangshuo Li44,61 • Ting-Chi Wen38,62 • Paras Nath Singh43 • Yusufjon Gafforov63,64,70 • Itthayakorn Promputtha5,72 •
Erandi Yasanthika2,3 • Ishani D. Goonasekara1,2,3 • Rui-Lin Zhao44 • Qi Zhao1 • Paul M. Kirk65 •
Jian-Kui Liu37,66 • JiYe Yan35 • Peter E. Mortimer1,71 • Jianchu Xu1,4,71 • Mingkwan Doilom1,3,4,5,67,71
1
Key Laboratory for Plant Diversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of
Science, Kunming 650201, Yunnan, People’s Republic of
China
10
No. 128/1-J, Azad Housing Society,
Curca, Goa Velha 403108, India
11
Department of Crop Sciences, College of Agricultural and
Marine Sciences, Sultan Qaboos University, PO Box 8,
123 Al Khoud, Oman
2
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
12
3
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
Sect. Environmental Sciences, Dept. MeSVA, University of
L’Aquila, 67100 Coppito, AQ, Italy
13
4
East and Central Asia Regional Office, World Agroforestry
Centre (ICRAF), Kunming 650201, Yunnan, People’s
Republic of China
Division of Food Technology, Biotechnology &
Agrochemistry, College of Agriculture & Life Sciences,
Chonnam National University, Gwangju 61186, Korea
14
5
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
Fungal Resources Research Division, NNIBR,
Sangju-si 37242, Korea
15
6
Mushroom Research Foundation, 128 M.3 Ban Pa Deng T. Pa
Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8,
3584 CT Utrecht, The Netherlands
16
7
Department of Plant Medicine, National Chiayi University,
300 Syuefu Road, Chiayi City 60004, Taiwan, People’s
Republic of China
Department of Biotechnology and Biomedicine, DTUBioengineering, Technical University of Denmark,
2800 Kongens Lyngby, Denmark
17
8
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit, Mauritius
Botanischer Garten und Botanisches Museum Berlin, Freie
Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin,
Germany
9
Formerly Department of Botany, Goa University, Goa, India
123
Fungal Diversity (2019) 96:1–242
18
Laboratório de Botânica/Liquenologia, Instituto de
Biociências, Universidade Federal de Mato Grosso do Sul,
Avenida Costa e Silva s/n, Bairro Universitário,
Campo Grande, Matto Grosso do Sul CEP 79070-900, Brazil
19
Departamento de Biociências, Universidade Federal de
Sergipe, Itabaiana, Sergipe CEP: 49500-000, Brazil
20
21
22
23
24
25
26
27
28
36
Yunnan Institute of Food Safety, Kunming University of
Science and Technology, Kunming 650500, People’s
Republic of China
37
Guizhou Key Laboratory of Agricultural Biotechnology,
Guizhou Academy of Agricultural Sciences,
Guiyang 550006, Guizhou, People’s Republic of China
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Sciences and Oceanography, Shenzhen
University, Shenzhen 518060, People’s Republic of China
38
Engineering Research Center of Southwest BioPharmaceutical Resources, Ministry of Education, Guizhou
University, Guiyang 550025, People’s Republic of China
Industrial Science and Management (International Program),
Faculty of Science and Technology, Thammasat University
(Rangsit Center), Klong Luang, Rangsit, Pathumthani 12121,
Thailand
39
Jodrell Laboratory, Royal Botanic Gardens, Kew,
Surrey TW9 3AB, UK
40
Institute of Microbiology, Beijing Forestry University,
PO Box 61, Beijing 100083, People’s Republic of China
41
Shandong Provincial Key Laboratory of Applied Mycology,
Qingdao Agricultural University, Qingdao 266109, People’s
Republic of China
42
Rud̄er Bošković Institute, Bijenička cesta 54, 10000 Zagreb,
Croatia
43
National Fungal Culture Collection of India (NFCCI),
Biodiversity and Palaeobiology Group, MACS Agharkar
Research Institute, G.G. Agarkar Road, Pune 411 004, India
44
State Key Laboratory of Mycology, Institute of
Microbiology, Chinese Academy of Sciences,
Beijing 100101, People’s Republic of China
45
Institute of Botany, Czech Academy of Sciences,
252 43 Průhonice, Czech Republic
46
Las Muros, 09420 Rimont, France
47
Institute for Clinical Biochemistry and Diagnostics,
University Hospital Hradec Králové, 500 05 Hradec Králové,
Czech Republic
48
Centro de Biociências, Departamento de Micologia,
Universidade Federal de Pernambuco, Avenida da
Engenharia, S/N 50740-600 – Cidade Universitária, Recife,
Pernambuco, Brazil
49
Department Research, Meise Botanic Garden, Nieuwelaan
38, 1860 Meise, Belgium
50
Fédération Wallonie-Bruxelles, Direction Générale de
l’Enseignement non obligatoire et de la Recherche
scientifique, Rue A. Lavallée 1, 1080 Brussels, Belgium
51
Faculdade de Ciências, Biosystems and Integrative Sciences
Institute (BioISI), Universidade de Lisboa, Campo Grande,
1749-016 Lisbon, Portugal
52
A.M.B, Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314,
Brescia, Italy
53
A.M.B. Gruppo, Micologico Forlivese ‘‘Antonio
Cicognani’’, Via Roma 18, Forlı́, Italy
54
Società per gli Studi Naturalistici della Romagna,
C.P. 143, Bagnacavallo, RA, Italy
55
Russian Research Institute of Floriculture and Subtropical
Crops, Yana Fabritsiusa Street 2/28, Sochi, Krasnodar region,
Russia 354002
56
Department of Botany and Microbiology, College of Science,
King Saud University, P.O Box 2455, Riyadh 11451,
Kingdom of Saudi Arabia
Instituto de Ecologı́a, A. C., Apartado Postal 63,
91000 Xalapa, Veracruz, Mexico
College of Agriculture and Biological Science, Dali
University, Dali 671003, Yunnan, People’s Republic of
China
Departamento Fitopatologia, Universidade Federal de
Viçosa, Viçosa, Minas Gerais, Brazil
Universidade Federal de Viçosa, Departamento de
Microbiologia, Viçosa, Minas Gerais, Brazil
Institut de Systématique, Evolution, Biodiversité (ISYEB –
UMR 7205), Muséum national d’Histoire naturelle, Sorbonne
Université, CNRS, CP 39, 12 Rue Buffon, 75005 Paris,
France
Centre National de Recherche sur l’Environnement (CNRE),
Lab. de Microbiologie de l’Environnement (LME),
BP 1739, Antananarivo, Madagascar
Department of Plant Protection, Agroscope ChanginsWädenswil Research Station ACW, Rte De Duiller,
1260 Nyon 1, Switzerland
29
Departamento de Micologia, Universidade Federal de
Pernambuco, Avenida da Engenharia, S/N,
Recife 50740-600, Brazil
30
Museu Paraense Emı́lio Goeldi, Coord. Botânica, Av.
Perimetral 1901, Terra Firme, C.P. 399, Belém,
PA 66040-170, Brazil
31
32
33
34
35
241
Instituto de Ciências Biológicas, Laboratório de
Microbiologia, Av. Augusto Corrêa, Belém, Pará 66075-110,
Brazil
Coordenação de Engenharia Ambiental, Universidade do
Estado do Amapá, Av. Presidente Vargas,
Macapá 68900-070, Brazil
Programa de Pós-Graduação em Biologia de Fungos,
Departamento de Micologia Prof. Chaves Batista, Centro de
Biociências, Universidade Federal de Pernambuco, Recife,
PE 50670-901, Brazil
Department of Entomology and Plant Pathology, Faculty of
Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
Beijing Key Laboratory of Environment Friendly
Management on Fruit Diseases and Pests in North China,
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences,
Beijing 100097, People’s Republic of China
123
242
Fungal Diversity (2019) 96:1–242
57
Biodiversity Integration Knowledge Center, Arizona State
University, PO Box 874108, Tempe, AZ 85287-4108, USA
65
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS,
UK
58
Charles Darwin Foundation for the Galapagos Islands,
Puerto Ayora, Ecuador
66
59
Instituto Nacional de Biodiversidad, Quito, Ecuador
School of Life Science and Technology, University of
Electronic Science and Technology of China,
Chengdu 611731, People’s Republic of China
60
67
College of Science and Engineering, Flinders University,
GPO Box 2100, Adelaide, SA 5001, Australia
Institute of Animal Science, State Key Laboratory of Animal
Nutrition, Chinese Academy of Agricultural Sciences,
Beijing 100193, People’s Republic of China
61
College of Life Sciences, Hebei University,
Baoding 071002, Hebei Province, People’s Republic of
China
68
Center of Excellence in Microbial Diversity and Sustainable
Utilization, Chiang Mai University, Chiang Mai 50200,
Thailand
69
Academy of Science, The Royal Society of Thailand,
Bangkok 10300, Thailand
70
Institute of Applied Ecology, Chinese Academy of Sciences,
Shenyang 110016, People’s Republic of China
62
State Key Laboratory Breeding Base of Green Pesticide and
Agricultural Bioengineering, Key Laboratory of Green
Pesticide and Agricultural Bioengineering, Ministry of
Education, Guizhou University, Guiyang 550025, People’s
Republic of China
63
Laboratory of Mycology, Institute of Botany, Academy of
Sciences of the Republic of Uzbekistan, 32 Durmon Yuli
Street, Tashkent, Uzbekistan 100125
71
Centre for Mountain Futures (CMF), Kunming Institute of
Botany, Kunming 650201, Yunnan, People’s Republic of
China
64
Department of Ecology, University of Kassel, HeinrichPlettStrasse, 40, 34132 Kassel, Germany
72
Center of Excellence in Bioresources for Agriculture,
Industry and Medicine, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
123