Fungal Diversity
https://doi.org/10.1007/s13225-019-00421-w
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Fungal diversity notes 929–1035: taxonomic and phylogenetic
contributions on genera and species of fungi
Rungtiwa Phookamsak1,2,3,4,6 • Kevin D. Hyde1,3,4,5 • Rajesh Jeewon7 • D. Jayarama Bhat8 •
E. B. Gareth Jones17,48 • Sajeewa S. N. Maharachchikumbura9 • Olivier Raspé10,11 • Samantha C. Karunarathna1,2,4 •
Dhanushka N. Wanasinghe1,2,3,4 • Sinang Hongsanan3,12 • Mingkwan Doilom1,2,3,4,6 • Danushka S. Tennakoon1,3,5,13 •
Alexandre R. Machado14 • André L. Firmino15 • Aniket Ghosh16 • Anuruddha Karunarathna1,3,13,17 •
Armin Mešić18 • Arun Kumar Dutta19 • Benjarong Thongbai20 • Bandarupalli Devadatha21 •
Chada Norphanphoun3,5,13,22 • Chanokned Senwanna3,5,17 • Deping Wei1,3,5,17 • Dhandevi Pem3,5,12 •
Frank Kwekucher Ackah23 • Gen-Nuo Wang24 • Hong-Bo Jiang1,3,5 • Hugo Madrid25 • Hyang Burm Lee26 •
Ishani D. Goonasekara1,3,5 • Ishara S. Manawasinghe3,27 • Ivana Kušan18 • Josep Cano28 • Josepa Gené28 •
Junfu Li1,3 • Kanad Das29 • Krishnendu Acharya19 • K. N. Anil Raj30 • K. P. Deepna Latha30 •
K. W. Thilini Chethana3,27 • Mao-Qiang He31 • Margarita Dueñas32 • Margita Jadan18 • Marı́a P. Martı́n32 •
Milan C. Samarakoon3,6,33 • Monika C. Dayarathne1,3,5 • Mubashar Raza31,34 • Myung Soo Park35 •
M. Teresa Telleria32 • Napalai Chaiwan1,3,5 • Neven Matočec18 • Nimali I. de Silva1,3,5,6 • Olinto L. Pereira36 •
Paras Nath Singh37 • Patinjareveettil Manimohan30 • Priyanka Uniyal16 • Qiu-Ju Shang3,33 •
Rajendra P. Bhatt16 • Rekhani H. Perera3,5,33 • Renato Lúcio Mendes Alvarenga38 • Sandra Nogal-Prata32 •
Sanjay K. Singh37 • Santhiti Vadthanarat6 • Seung-Yoon Oh35 • Shi-Ke Huang1,3,5,22 • Shiwali Rana37 •
Sirinapa Konta1,3,5 • Soumitra Paloi19 • Subashini C. Jayasiri1,3,5 • Sun Jeong Jeon26 • Tahir Mehmood16 •
Tatiana Baptista Gibertoni38 • Thuong T. T. Nguyen26 • Upendra Singh16 • Vinodhini Thiyagaraja1,3,5,17 •
V. Venkateswara Sarma21 • Wei Dong3,5,17,39 • Xian-Dong Yu39 • Yong-Zhong Lu3,5,22 • Young Woon Lim35 •
Yun Chen40 • Zdenko Tkalčec18 • Zhi-Feng Zhang31,34 • Zong-Long Luo3,5,41 • Dinushani A. Daranagama42 •
Kasun M. Thambugala52 • Saowaluck Tibpromma1,2,3,4 • Erio Camporesi43,44,45 • Timur S. Bulgakov46 •
Asha J. Dissanayake3 • Indunil C. Senanayake3,12 • Dong Qin Dai47 • Li-Zhou Tang47 • Sehroon Khan1,4 •
Huang Zhang39 • Itthayakorn Promputtha6,50 • Lei Cai31 • Putarak Chomnunti3,49 • Rui-Lin Zhao31 •
Saisamorn Lumyong6,51 • Saranyaphat Boonmee3 • Ting-Chi Wen22 • Peter E. Mortimer1 •
Jianchu Xu2,4
Received: 13 December 2018 / Accepted: 26 February 2019
Ó The Author(s) 2019
Abstract
This article is the ninth in the series of Fungal Diversity Notes, where 107 taxa distributed in three phyla, nine classes, 31
orders and 57 families are described and illustrated. Taxa described in the present study include 12 new genera, 74 new
species, three new combinations, two reference specimens, a re-circumscription of the epitype, and 15 records of sexualasexual morph connections, new hosts and new geographical distributions. Twelve new genera comprise Brunneofusispora, Brunneomurispora, Liua, Lonicericola, Neoeutypella, Paratrimmatostroma, Parazalerion, Proliferophorum, Pseudoastrosphaeriellopsis, Septomelanconiella, Velebitea and Vicosamyces. Seventy-four new species are Agaricus
memnonius, A. langensis, Aleurodiscus patagonicus, Amanita flavoalba, A. subtropicana, Amphisphaeria mangrovei,
Baorangia major, Bartalinia kunmingensis, Brunneofusispora sinensis, Brunneomurispora lonicerae, Capronia camelliaeyunnanensis, Clavulina thindii, Coniochaeta simbalensis, Conlarium thailandense, Coprinus trigonosporus, Liua muriformis, Cyphellophora filicis, Cytospora ulmicola, Dacrymyces invisibilis, Dictyocheirospora metroxylonis, Distoseptispora thysanolaenae, Emericellopsis koreana, Galiicola baoshanensis, Hygrocybe lucida, Hypoxylon teeravasati,
& Jianchu Xu
j.c.xu@cgiar.org
Extended author information available on the last page of the article
123
Fungal Diversity
Hyweljonesia indica, Keissleriella caraganae, Lactarius olivaceopallidus, Lactifluus midnapurensis, Lembosia brigadeirensis, Leptosphaeria urticae, Lonicericola hyaloseptispora, Lophiotrema mucilaginosis, Marasmiellus bicoloripes,
Marasmius indojasminodorus, Micropeltis phetchaburiensis, Mucor orantomantidis, Murilentithecium lonicerae,
Neobambusicola brunnea, Neoeutypella baoshanensis, Neoroussoella heveae, Neosetophoma lonicerae, Ophiobolus
malleolus, Parabambusicola thysanolaenae, Paratrimmatostroma kunmingensis, Parazalerion indica, Penicillium dokdoense, Peroneutypa mangrovei, Phaeosphaeria cycadis, Phanerochaete australosanguinea, Plectosphaerella kunmingensis, Plenodomus artemisiae, P. lijiangensis, Proliferophorum thailandicum, Pseudoastrosphaeriellopsis kaveriana,
Pseudohelicomyces menglunicus, Pseudoplagiostoma mangiferae, Robillarda mangiferae, Roussoella elaeicola, Russula
choptae, R. uttarakhandia, Septomelanconiella thailandica, Spencermartinsia acericola, Sphaerellopsis isthmospora,
Thozetella lithocarpi, Trechispora echinospora, Tremellochaete atlantica, Trichoderma koreanum, T. pinicola, T. rugulosum, Velebitea chrysotexta, Vicosamyces venturisporus, Wojnowiciella kunmingensis and Zopfiella indica. Three new
combinations are Baorangia rufomaculata, Lanmaoa pallidorosea and Wojnowiciella rosicola. The reference specimens of
Canalisporium kenyense and Tamsiniella labiosa are designated. The epitype of Sarcopeziza sicula is re-circumscribed
based on cyto- and histochemical analyses. The sexual-asexual morph connection of Plenodomus sinensis is reported from
ferns and Cirsium for the first time. In addition, the new host records and country records are Amanita altipes, A.
melleialba, Amarenomyces dactylidis, Chaetosphaeria panamensis, Coniella vitis, Coprinopsis kubickae, Dothiorella
sarmentorum, Leptobacillium leptobactrum var. calidus, Muyocopron lithocarpi, Neoroussoella solani, Periconia cortaderiae, Phragmocamarosporium hederae, Sphaerellopsis paraphysata and Sphaeropsis eucalypticola.
Keywords 86 new taxa Agaricomycetes Ascomycota Basidiomycota Dacrymycetes Dothideomycetes
Eurotiomycetes Lecanoromycetes Leotiomycetes Mucoromycetes Mucoromycota Pezizomycetes
Sordariomycetes Taxonomy
Table of contents
Ascomycota R.H. Whittaker
Dothideomycetes O.E. Erikss. & Winka
Dothideomycetidae P.M. Kirk et al.
Capnodiales Woron.
Teratosphaeriaceae Crous & U. Braun
929. Hyweljonesia indica P.N. Singh & S.K. Singh, sp.
nov.
Pleosporomycetidae C.L. Schoch et al.
Pleosporales Luttr. ex M.E. Barr
Dictyosporiaceae Boonmee & K.D. Hyde
930. Dictyocheirospora metroxylonis Konta & K.D. Hyde,
sp. nov.
Didymosphaeriaceae Munk
931. Vicosamyces Firmino, A.R. Machado & O.L. Pereira,
gen. nov.
932. Vicosamyces venturisporus Firmino, A.R. Machado
& O.L. Pereira, sp. nov.
Lentitheciaceae Yin. Zhang et al.
933. Keissleriella caraganae Chaiwan, Phookamsak,
Wanas. & K.D. Hyde, sp. nov.
934. Murilentithecium lonicerae Phookamsak, Chaiwan,
Wanas. & K.D. Hyde, sp. nov.
123
935. Phragmocamarosporium hederae Wijayaw., R.K.
Schumach. & K.D. Hyde, Index Fungorum 370: 1 (2018),
new host record
Leptosphaeriaceae M.E. Barr
936. Leptosphaeria urticae D. Pem, E.B.G. Jones & K.D.
Hyde, sp. nov.
937. Plenodomus artemisiae A. Karunarathna, Phookamsak & K.D. Hyde, sp. nov.
938. Plenodomus lijiangensis Phookamsak, A. Karunarathna & K.D. Hyde, sp. nov.
939. Plenodomus sinensis Tennakoon, Phookamsak &
K.D. Hyde, in Tennakoon et al., Phytotaxa 324(1): 76
(2017), new hosts and asexual morph records
940. Sphaerellopsis isthmospora A. Karunarathna,
Phookamsak & K.D. Hyde, sp. nov.
941. Sphaerellopsis paraphysata Crous & Alfenas, in
Trakunyingcharoen et al., IMA Fungus 5(2): 411 (2014),
new host record from Yunnan, China
Lophiotremataceae K. Hiray. & Kaz. Tanaka
942. Lophiotrema mucilaginosis M. Raza & L. Cai, sp.
nov.
Occultibambusaceae D.Q. Dai & K.D. Hyde
943. Brunneofusispora S.K. Huang & K.D. Hyde, gen.
nov.
944. Brunneofusispora sinensis S.K. Huang & K.D. Hyde,
sp. nov.
Fungal Diversity
Parabambusicolaceae Kaz. Tanaka & K. Hiray.
945. Lonicericola Phookamsak, Jayasiri & K.D. Hyde,
gen. nov.
946. Lonicericola hyaloseptispora Phookamsak, Jayasiri &
K.D. Hyde, sp. nov.
947. Parabambusicola thysanolaenae Goonas., Jayasiri,
Phookamsak & K.D. Hyde, sp. nov.
948. Paratrimmatostroma Jayasiri, Phookamsak, D.J. Bhat
& K.D. Hyde, gen. nov.
949.
Paratrimmatostroma
kunmingensis
Jayasiri,
Phookamsak, D.J. Bhat & K.D. Hyde, sp. nov.
Periconiaceae (Sacc.) Nann.
950. Periconia cortaderiae Thambug. & K.D. Hyde, in
Thambugala et al., Mycosphere 8(4): 734 (2017), new host
record from Yunnan, China
Phaeosphaeriaceae M.E. Barr
951. Amarenomyces dactylidis Mapook, Camporesi &
K.D. Hyde, in Hyde et al., Fungal Divers 87: 78 (2017),
new host record from Yunnan, China
952. Brunneomurispora Phookamsak, Konta, Wanas. &
K.D. Hyde, gen. nov.
953. Brunneomurispora lonicerae Konta, Phookamsak,
Wanas. & K.D. Hyde, sp. nov.
954. Galiicola baoshanensis Phookamsak, Wanas. & K.D.
Hyde, sp. nov.
955. Neosetophoma lonicerae Phookamsak, Wanas. &
K.D. Hyde, sp. nov.
956. Ophiobolus malleolus S.K. Huang, Bulgakov & K.D.
Hyde, sp. nov.
957. Phaeosphaeria cycadis Wanas., Phookamsak & K.D.
Hyde, sp. nov.
958. Wojnowiciella kunmingensis Phookamsak, Wanas. &
K.D. Hyde, sp. nov.
959. Wojnowiciella rosicola (W.J. Li et al.) Wanas.,
Phookamsak & K.D. Hyde, comb. nov.
Pseudoastrosphaeriellaceae Phookamsak & K.D. Hyde
960. Pseudoastrosphaeriellopsis Devadatha, Wanas., Jeewon & V.V. Sarma, gen. nov.
961. Pseudoastrosphaeriellopsis kaveriana Devadatha,
Wanas., Jeewon & V.V. Sarma, sp. nov.
Roussoellaceae J.K. Liu et al.
962. Neoroussoella heveae Senwanna, Phookamsak &
K.D. Hyde, sp. nov.
963. Neoroussoella leucaenae Jayasiri, E.B.G. Jones &
K.D. Hyde, Mycosphere 10(1): 1–186 (2019), new host
record from Yunnan, China
964. Roussoella elaeicola Konta & K.D. Hyde, sp. nov.
Sulcatisporaceae Kaz. Tanaka & K. Hiray.
965. Neobambusicola brunnea Y. Chen & Norphanphoun,
sp. nov.
Thyridariaceae Q. Tian & K.D. Hyde
966. Liua Phookamsak & K.D. Hyde, gen. nov.
967. Liua muriformis Phookamsak, H.B. Jiang & K.D.
Hyde, sp. nov.
Dothideomycetes, orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf.
968. Lembosia brigadeirensis Firmino, A.R. Machado &
O.L. Pereira, sp. nov.
Botryosphaeriales C.L. Schoch et al.
Botryosphaeriaceae Theiss. & P. Syd.
969. Dothiorella acericola Phookamsak, Tennakoon &
K.D. Hyde, sp. nov.
970. Dothiorella sarmentorum (Fr.) A.J.L. Phillips, A.
Alves & J. Luque, Mycologia 97(2): 522 (2005), new host
record from Russia
971. Sphaeropsis eucalypticola A.J.L. Phillips, in Phillips
et al., Stud Mycol 76: 158 (2013), new host record
Microthyriales G. Arnaud
Microthyriales, genera incertae sedis
972. Parazalerion Madrid, Gené & Cano, gen. nov.
973. Parazalerion indica Madrid, Gené, & Cano, sp. nov.
Muyocopronales Mapook et al.
Muyocopronaceae K.D. Hyde
974. Muyocopron lithocarpi Mapook, Boonmee & K.D.
Hyde, in Mapook et al., Phytotaxa 265(3): 235 (2016), new
host record from Yunnan, China
Tubeufiales Boonmee & K.D. Hyde
Tubeufiaceae M.E. Barr
975. Pseudohelicomyces menglunicus J.F. Li, Phookamsak & K.D. Hyde, sp. nov.
Eurotiomycetes O.E. Erikss. & Winka
Chaetothyriomycetidae Doweld
Chaetothyriales M.E. Barr
Cyphellophoraceae Réblová & Unter.
976. Cyphellophora filicis Hongsanan, Phookamsak &
K.D. Hyde, sp. nov.
Herpotrichiellaceae Munk
977. Capronia camelliae-yunnanensis M. Raza, Z.F.
Zhang & L. Cai, sp. nov.
Eurotiomycetidae Geiser & Lutzoni
Eurotiales G.W. Martin ex Benny & Kimbr.
Trichocomaceae E. Fisch.
978. Penicillium dokdoense Hyang B. Lee & T.T.T.
Nguyen, sp. nov.
Lecanoromycetes, O.E. Erikss. & Winka
123
Fungal Diversity
Lecanoromycetes, families incertae sedis
Micropeltidaceae Clem. & Shear
979. Micropeltis phetchaburiensis Dayarathne, Hongsanan
& K.D. Hyde, sp. nov.
Leotiomycetes O.E. Erikss. & Winka
Helotiales Nannf. ex Korf & Lizoň
Lachnaceae Raitv.
980. Velebitea I. Kušan, Matočec & Jadan, gen. nov.
981. Velebitea chrysotexta I. Kušan, Matočec & Jadan, sp.
nov.
Pezizomycetes O.E. Erikss. & Winka
Pezizales J. Schröt.
Pezizaceae Dumort.
982. Sarcopeziza sicula (Inzenga) Agnello, Loizides & P.
Alvarado, Ascomycete.org 10(4): 179 (2018), re-circumscription of the epitype
Sordariomycetes O.E. Erikss. & Winka
Diaporthomycetidae Senan. et al.
Atractosporales H. Zhang et al.
Conlariaceae H. Zhang et al.
983. Conlarium thailandense X.D. Yu, H. Zhang & K.D.
Hyde, sp. nov.
Cytosporaceae Fr.
984. Cytospora ulmicola Norphanphoun, Bulgakov, T.C.
Wen & K.D. Hyde, sp. nov.
Melanconiellaceae Senan. et al.
985. Septomelanconiella Samarak. & K.D. Hyde, gen.
nov.
986. Septomelanconiella thailandica Samarak. & K.D.
Hyde, sp. nov.
Dayarathne,
Schizoparmaceae Rossman
988. Coniella vitis Chethana, J.Y. Yan, X.H. Li & K.D.
Hyde, Pl Dis 101: 2129 (2017), new host record from
Russia
Diaporthomycetidae, families incertae sedis
Distoseptisporaceae K.D. Hyde & McKenzie
989. Distoseptispora thysanolaenae Goonas., Dayarathne,
Phookamsak & K.D. Hyde, sp. nov.
Diaporthomycetidae, genera incertae sedis
990. Proliferophorum G.N. Wang, H. Zhang & Senan.,
gen. nov.
123
Hypocreomycetidae O.E. Erikss. & Winka
Glomerellales Chadef. ex Réblová et al.
Plectosphaerellaceae W. Gams et al.
992. Plectosphaerella kunmingensis Phookamsak, J.F. Li
& K.D. Hyde, sp. nov.
Hypocreales Lindau
Cordycipitaceae Kreisel ex G.H. Sung et al.
993. Leptobacillium leptobactrum var. calidus (W. Gams)
Zare & W. Gams, Mycol Prog 15: 1003 (2016), new record
for India
Hypocreaceae De Not.
994. Trichoderma koreanum S-Y. Oh, M.S. Park & Y.W.
Lim, sp. nov.
995. Trichoderma pinicola S-Y. Oh, M.S. Park & Y.W.
Lim, sp. nov.
996. Trichoderma rugulosum M.S. Park, S-Y. Oh & Y.W.
Lim, sp. nov.
Hypocreales, genera incertae sedis
997. Emericellopsis koreana Hyang B. Lee, S.J. Jeon &
T.T.T. Nguyen, sp. nov.
Savoryellomycetidae Hongsanan et al.
Diaporthales Nannf.
Pseudoplagiostomataceae Cheew. et al.
987.
Pseudoplagiostoma
mangiferae
Phookamsak & K.D. Hyde, sp. nov.
991. Proliferophorum thailandicum G.N. Wang, H. Zhang
& Senan., sp. nov.
Savoryellales Boonyuen et al.
Savoryellaceae Jaklitsch & Réblová
998. Canalisporium kenyense Goh, W.H. Ho & K.D.
Hyde, Can J Bot 76: 148 (1998), reference specimen
Sordariomycetidae O.E. Erikss. & Winka
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae Réblová et al.
999. Chaetosphaeria panamensis Huhndorf & F.A. Fernández, Fungal Divers 19: 33 (2005), new host record
from Taiwan
1000. Thozetella lithocarpi R.H. Perera & K.D. Hyde, sp.
nov.
Coniochaetales Huhndorf et al.
Coniochaetaceae Malloch & Cain
1001. Coniochaeta simbalensis S. Rana & S.K. Singh, sp.
nov.
Phyllachorales M.E. Barr
Phyllachoraceae Theiss. & H. Syd.
1002. Tamsiniella labiosa S.W. Wong, K.D. Hyde, W.H.
Ho & S.J. Stanley, Can J Bot 76(2): 334 (1998), reference
specimen
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss.
Lasiosphaeriaceae Nannf.
Fungal Diversity
1003. Zopfiella indica Devadatha, Jeewon & V.V. Sarma,
sp. nov.
Xylariomycetidae O.E. Erikss. & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Amphisphaeriaceae G. Winter
1004. Amphisphaeria mangrovei Devadatha & V.V.
Sarma, sp. nov.
Sporocadaceae Corda
1005. Bartalinia kunmingensis Thiyag., Wanas.,
Phookamsak & K.D. Hyde, sp. nov.
1006. Robillarda mangiferae Thiyag., Wanas., Phookamsak & K.D. Hyde, sp. nov.
Xylariales Nannf.
Diatrypaceae Nitschke
1007. Neoeutypella M. Raza, Q.J. Shang, Phookamsak &
L. Cai, gen. nov.
1008. Neoeutypella baoshanensis M. Raza, Q.J. Shang,
Phookamsak & L. Cai, sp. nov.
1009. Peroneutypa mangrovei Devadatha & V.V. Sarma,
sp. nov.
Hypoxylaceae DC.
1010. Hypoxylon teeravasati Devadatha, V.V. Sarma &
E.B.G. Jones, sp. nov.
Basidiomycota R.T. Moore
Agaricomycetes Doweld
Agaricomycetidae Parmasto
Agaricales Underw.
Agaricaceae Chevall.
1011. Agaricus memnonius M.Q. He & R.L. Zhao, sp.
nov.
1012. Agaricus langensis M.Q. He & R.L. Zhao, sp. nov.
1013. Coprinus trigonosporus Tkalčec & Mešić, sp. nov.
Amanitaceae E.-J. Gilbert
1014. Amanita altipes Zhu L. Yang, M. Weiss & Oberw.,
Mycologia 96(3): 636 (2004), new record from Thailand
1015. Amanita flavoalba Mehmood & R.P. Bhatt, sp. nov.
1016. Amanita melleialba Zhu L. Yang, Qing Cai & Yang
Y. Cui, in Ariyawansa et al., Fungal Divers: https://doi.org/
10.1007/s13225-015-0346-5, [163] (2015), new record
from Thailand
1017. Amanita subtropicana Mehmood & R.P. Bhatt, sp.
nov.
Hygrophoraceae Lotsy
1018. Hygrocybe lucida K. Acharya & A.K. Dutta, sp. nov.
Marasmiaceae Roze ex Kühner
1019. Marasmius indojasminodorus A.K. Dutta, K.
Acharya & K. Das, sp. nov.
Omphalotaceae Bresinsky
1020. Marasmiellus bicoloripes K.P.D. Latha, K.N.A Raj
& Manim., sp. nov.
Psathyrellaceae Vilgalys et al.
1021. Coprinopsis kubickae (Pilát & Svrček) Redhead
et al., in Redhead et al., Taxon 50(1): 229 (2001), new
record for Croatia
Boletales E.-J. Gilbert
Boletaceae Chevall.
1022. Baorangia major Raspé & Vadthanarat, sp. nov.
1023. Baorangia rufomaculata (Both) Raspé & Vadthanarat, comb. nov.
1024. Lanmaoa pallidorosea (Both) Raspé & Vadthanarat,
comb. nov.
Cantharellales Gäum.
Clavulinaceae Donk
1025. Clavulina thindii U. Singh, sp. nov.
Polyporales Güum.
Phanerochaetaceae Jülich
1026. Phanerochaete australosanguinea Telleria, M.
Dueñas & M.P. Martı́n, sp. nov.
Russulales Kreisel ex P.M. Kirk et al.
Russulaceae Lotsy
1027. Lactarius olivaceopallidus Uniyal, sp. nov.
1028. Lactifluus midnapurensis S. Paloi & K. Acharya, sp.
nov.
1029. Russula choptae A. Ghosh & K. Das, sp. nov.
1030. Russula uttarakhandia A. Ghosh & K. Das, sp. nov.
Stereaceae Pilát
1031. Aleurodiscus patagonicus Nogal, Telleria, M. Dueñas & M.P. Martı́n, sp. nov.
Trechisporales K.H. Larss.
Hydnodontaceae Jülich
1032. Trechispora echinospora Telleria, M. Dueñas, I.
Melo & M.P. Martı́n, sp. nov.
Auriculariomycetidae Jülich
Auriculariales J. Schröt.
Auriculariaceae Fr. ex Lindau
1033. Tremellochaete atlantica Alvarenga, sp. nov.
Dacrymycetes Doweld
Dacrymycetales Henn.
Dacrymycetaceae J. Schröt.
1034. Dacrymyces invisibilis M. Dueñas, Telleria & M.P.
Martı́n, sp. nov.
Mucoromycota Doweld
Mucoromycetes Doweld
123
Fungal Diversity
Mucorales Fr.
Mucoraceae Dumort.
1035. Mucor orantomantidis Hyang B. Lee, P.M. Kirk &
T.T.T. Nguyen, sp. nov.
Introduction
Fungi are well-known as a large and diverse group of
microorganisms that play important functional roles from
agricultural, ecological and economic perspectives. They are
crucial to natural ecosystems as decomposers degrading dead
organic materials, accelerating rock weathering and response
to plant growth, nutrient cycling, as well as maintaining plant
diversity (Kendrick 2000; Finlay 2008; Zechmeister-Boltenstern et al. 2015; Drinkwater et al. 2017; Horwath 2017; Hyde
et al. 2018a, b; Willis 2018). They are heterotrophic and may
change their lifestyles from endophytic to pathogenic to
saprobic on plants or other organisms as well as other fungi
depending on the environmental circumstances (Hyde et al.
2007, 2018a; Promputtha et al. 2007, 2010; Slippers and
Wingfield 2007; Ghimire and Hyde 2008; Hyde and Soytong
2008; Gomes et al. 2013; Zhan et al. 2016; Ariyawansa et al.
2018; Haelewaters et al. 2018; Liyanage et al. 2018; Lofgren
et al. 2018; Wang et al. 2018; Sun et al. 2019). Sixteen phyla
are accepted in the Kingdom Fungi (Tedersoo et al. 2018;
Wijayawardene et al. 2018b).
Hawksworth (1991, 2001) estimated 1.5 million species of
fungi worldwide, with fewer than 5–10% having been
described. Hawksworth and Lücking (2017) attempted to
derive an updated estimate of global fungal diversity based on
scientific evidence such as the extrapolations of plant/fungus
ratios, including molecular and fieldwork data from the same
sites. They concluded that there is an estimated 2.2–3.8 million undescribed species with taxa awaiting discovery in
biodiversity hot spots, with only 120,000 species described
and accepted (Hawksworth and Lücking 2017).
In our ongoing research compiling notes on new fungal
taxa, reference specimens, new data, and other taxonomic
contributions, more than 900 species have been introduced,
re-circumscribed and illustrated worldwide based up on
morphological characteristics and phylogenetic analyses.
This is the ninth paper in the fungal diversity series with
more than 100 species contributions which were mainly
collected from China, some other Asian countries, as well
as other parts of the world.
Materials and methods
Materials and methods follow the previous fungal diversity
notes (Hyde et al. 2016; Tibpromma et al. 2017). Fresh and
dried specimens in this study were collected from Australia,
123
Brazil, Chile, China, Croatia, Equatorial Guinea, India, Italy,
Korea, New Zealand, Russia, Saudi Arabia, Taiwan, Thailand, UK and the USA. Media agar used to cultivated fungi is
shown in Table 1. The genes and primers used in this study
are shown in the Table 2. Phylogenetic analyses were performed based on Bayesian inference (BI), maximum likelihood (ML) and maximum parsimony (MP) (see Table 2).
Phylum Ascomycota R.H. Whittaker
We follow the latest treatments and updated accounts of
Ascomycota in Wijayawardene et al. (2017a, 2018a).
Class Dothideomycetes O.E. Erikss. & Winka
The Classification of families in Dothideomycetes follow
Hyde et al. (2013), Liu et al. (2017a) and Wijayawardene
et al. (2018a). The subclasses, orders and families of Dothideomycetes are listed in alphabetical order.
Subclass Dothideomycetidae P.M. Kirk
Capnodiales Woron.
Teratosphaeriaceae Crous & U. Braun
Teratosphaeriaceae was introduced by Crous et al.
(2007a) and is typified by Teratosphaeria Syd. & P. Syd.
The family was introduced to accommodate several
important leaf spot and extremotolerant species initially
included in the genera Teratosphaeria, Mycosphaerella
and related asexual morph genera. Recently, 59 genera
were listed in this family (Wijayawardene et al. 2018a).
The latest treatments of genera in Teratosphaeriaceae were
outlined in Quaedvlieg et al. (2014), Wäli et al. (2014) and
Hyde et al. (2017).
Hyweljonesia R.G. Shivas et al.
A monotypic genus, Hyweljonesia was introduced in
Teratosphaeriaceae by Shivas et al. (2016) to accommodate H. queenslandica R.G. Shivas et al. (as the type
Table 1 Abbreviations of media agar used for fungal cultivation in
this study
Name
Abbreviation
Cornmeal dextrose agar
CMD
Creatine sucrose agar
CREA
Czapek yeast autolysate agar
CYA
Malt extract agar
MEA
Oat agar
OA
Potato carrot agar
PCA
Potato dextrose agar
Sea water agar
PDA
SWA
Spezieller nährstoffarmer agar
SNA
Synthetic mucor agar
SMA
Water agar
WA
Yeast extract sucrose agar
YES
Fungal Diversity
Table 2 Genes sequenced, primers used and phylogenetic analyses performed in this study
Family
Genes/loci
Primers
Phylogenetic
analysis
References
Agaricaceae
LSU, TEF1-a and
ITS (Agaricus);
ITS (Coprinus)
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), EF1-983F/EF1-1567R
(TEF1-a) (Agaricus); ITS1/
ITS4 (ITS, Coprinus)
BI and ML
(Agaricus);
ML (Coprinus)
Amanitaceae
LSU and RPB2
LR0R/LR5(LSU), rpb2-Am6F/Am-7R (RPB2)
ML
Amphisphaeriaceae
LSU, SSU and ITS
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), NS1/NS4 (SSU)
BI, ML and MP
Vilgalys and Hester (1990), White et al.
(1990), O’Donnell (1993), Glass and
Donaldson (1995), Carbone and Kohn
(1999), Liu et al. (1999), Rehner
(2001), Samuels et al. (2002), Matheny
(2005), Hong et al. (2006), Cai et al.
(2014), Raspé et al. (2016)
Asterinaceae
LSU
LR0R/LR5 (LSU)
BI
Auriculariaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR5
(LSU)
BI and ML
Boletaceae
ATP6, RPB2 and
TEF1-a
ATP6-1M40F/ATP6-2 M
(ATP6), bRPB2-6F/bRPB27.1R (RPB2), EF1-983F/EF12218R (TEF1-a)
ML
Botryosphaeriaceae
ITS and TEF1-a
(Dothiorella); ITS,
TEF1-a and TUB2
(Sphaeropsis)
ITS5/ITS4 (ITS), EF1-728F/
EF1-986R (TEF1-a), BT2a/
BT2b (TUB2)
BI and ML
(Dothiorella);
BI, ML and
MP
(Sphaeropsis)
Chaetosphaeriaceae
LSU, ITS and TUB2
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), BT2a/BT2b (TUB2)
BI and ML
Clavulinaceae
ITS
ITS1/ITS4 (ITS)
ML
Coniochaetaceae
LSU and ITS
ITS5/ITS4 (ITS), LR0R/LR7
(LSU)
ML
Conlariaceae
LSU, SSU and ITS
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), NS1/NS4 (SSU)
BI and ML
Cordycipitaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR7
(LSU)
ML and MP
Cyphellophoraceae
ITS, LSU and SSU
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), NS1/NS4 (SSU)
BI and ML
Cytosporaceae
ITS, LSU, and ACT
ITS1/ITS4 (ITS), NL1/NL4
(LSU), ACT512F/ACT783R
(ACT)
ML
Dacrymycetaceae
ITS
ITS5/ITS4 (ITS)
BI, ML and MP
Diaporthomycetidae,
genera incertae sedis
(Proliferophorum)
LSU, SSU and ITS
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), NS1/NS4 (SSU)
BI and ML
Diatrypaceae
ITS and TUB2
ITS1/ITS4 (ITS), BT2a/BT2b
(TUB2)
BI, ML and MP
Dictyosporiaceae
ITS, LSU and TEF1a
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), EF1-983F/EF1-2218R
(TEF1-a)
BI, ML and MP
Didymosphaeriaceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR5
(LSU)
BI
Distoseptisporaceae
ITS, LSU and TEF1a
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), EF1-983F/EF1-2218R
(TEF1-a)
BI, ML and MP
Herpotrichiellaceae
ITS, LSU and SSU
ITS5/ITS4 (ITS), LR0R/LR5
(LSU), NS1/NS4 (SSU)
BI and ML
Hydnodontaceae
ITS
ITS5/ITS4 (ITS)
BI, ML and MP
Hygrophoraceae
ITS
ITS1/ITS4 (ITS)
ML
123
Fungal Diversity
Table 2 continued
Family
Genes/loci
Primers
Phylogenetic
analysis
Hypocreaceae
TEF1-a and RPB2
EF1-728F/TEF1rev (TEF1-a), fRPB2-5F/fRPB2-7cR (RPB2)
BI and ML
Hypocreales, genera
incertae sedis
(Emericellopsis)
ITS and TUB2
ITS1/ITS4 (ITS), BT2a/BT2b (TUB2)
ML
Hypoxylaceae
ITS, LSU, RPB2
and TUB2
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), fRPB2-5F/fRPB2-7cR
(RPB2), BT2a/BT2b (TUB2)
BI, ML and
MP
Lachnaceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR7 (LSU)
BI and ML
Lasiosphaeriaceae
LSU, TUB2, ITS
and RPB2
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), BT2a/BT2b (TUB2),
fRPB2-5F/fRPB2-7cR (RPB2)
BI, ML and
MP
Lentitheciaceae
LSU, SSU, ITS
and TEF1-a
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a)
BI and ML
Leptosphaeriaceae
LSU, SSU and
ITS
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU)
BI and ML
Lophiotremataceae
ITS, LSU, SSU,
TEF1-a and
RPB2
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a), fRPB2-5f/fRPB2-7cR (RPB2)
BI and ML
Marasmiaceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR3 (LSU)
BI and ML
Melanconiellaceae
ITS, LSU and
RPB2
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), fRPB2-5f/fRPB2-7cR
(RPB2)
BI, ML and
MP
Micropeltidaceae
LSU and SSU
LR0R/LR5 (LSU), NS1/NS4 (SSU)
ML and MP
Microthyriales, genera
incertae sedis
(Parazalerion)
LSU
LR0R/LR5 (LSU)
ML
Mucoraceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR5F (LSU)
ML
Muyocopronaceae
LSU and SSU
LR0R/LR5 (LSU), NS1/NS4 (SSU)
BI and ML
Occultibambusaceae
LSU, SSU, ITS,
RPB2 and
TEF1-a
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR7 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a), fRPB2-5f/fRPB2-7cR (RPB2)
BI, ML and
MP
ITS1/ITS4 (ITS), LR0R/LR7 (LSU)
BI and ML
Parabambusicolaceae
SSU, ITS, LSU
and TEF1-a
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a)
BI and ML
Periconiaceae
ITS, LSU and
TEF1-a
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), EF1-983F/EF1-2218R
(TEF1-a)
BI and ML
Pezizaceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR7 (LSU)
ML
Phaeosphaeriaceae
LSU, SSU, TEF1a and ITS
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a)
BI and ML
Phanerochaetaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR5 (LSU)
BI, ML and
MP
Phyllachoraceae
LSU, SSU, and
ITS
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU)
BI and ML
Plectosphaerellaceae
LSU, ITS and
TEF1-a
ITS1/ITS4 (ITS), LR0R/LR5 (LSU), EF1-983F/EF1-2218R
(TEF1-a)
BI and ML
Psathyrellaceae
ITS
ITS1/ITS4 (ITS)
ML
Pseudoastrosphaeriellaceae
LSU, SSU, ITS,
RPB2 and
TEF1-a
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a), fRPB2-5f/fRPB2-7cR (RPB2)
BI and ML
Pseudoplagiostomataceae
ITS, LSU, TUB2
and TEF1-a
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), BT2a/BT2b (TUB2),
EF1-983F/EF1-2218R (TEF1-a)
BI, ML and
MP
Roussoellaceae
LSU, SSU, TEF1a, ITS and RPB2
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU), EF1983F/EF1-2218R (TEF1-a), fRPB2-5f/fRPB2-7cR (RPB2)
BI and ML
Omphalotaceae
123
References
Fungal Diversity
Table 2 continued
Family
Genes/loci
Primers
Phylogenetic analysis
Russulaceae
ITS and LSU
ITS1/ITS4 (ITS), LR0R/LR3 (LSU)
ML (Lactarius); BI and
ML (Lactifluus), BI
(Russula)
Savoryellaceae
LSU and ITS
ITS5/ITS4 (ITS), LR0R/LR5 (LSU)
BI and ML
Schizoparmaceae
ITS, LSU, HIS3
and TEF1-a
ITS4/ITS5 (ITS), LR0R/LR5 (LSU); H3-1A/H3-1B
(HIS3), EF1-728F/EF1-986R (TEF1-a)
MP
Sporocadaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR5 (LSU)
BI and ML
Stereaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR5 (LSU)
BI, ML and MP
Sulcatisporaceae
LSU and ITS
ITS5/ITS4 (ITS), LR0R/LR5 (LSU)
ML
Teratosphaeriaceae
ITS and LSU
ITS5/ITS4 (ITS), LR0R/LR7 (LSU)
ML
Thyridariaceae
LSU, SSU,
TEF1-a, ITS
and RPB2
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), NS1/NS4 (SSU),
EF1-983F/EF1-2218R (TEF1-a), fRPB2-5f/fRPB2-7cR
(RPB2)
BI and ML
Trichocomaceae
TUB2 and CMD
BT2a/BT2b (TUB2), CMD5/CMD6 (CMD)
ML
Tubeufiaceae
ITS, LSU, TEF1a and RPB2
ITS5/ITS4 (ITS), LR0R/LR5 (LSU), EF1-983F/EF12218R (TEF1-a), fRPB2-5f/fRPB2-7cR (RPB2)
BI and ML
species) isolated from a cocoon of an unidentified
microlepidoptera parasitized by a chalcidoid wasp (Hymenoptera: Chalcoidea), collected from tropical forests of
northern Queensland, Australia. The genus is characterized
by white, septate, smooth-walled, hyaline to subhyaline
mycelial hyphae often form hyphal tufts from which
straight, unbranched, light brown, smooth-walled, and
septate conidiophores arise laterally. Subhyaline, cuneiform, smooth-walled conidia are produced on characteristic
integrated, pale brown and minutely verruculose conidiogenous cells forming apical whorls (1–5) of conidiogenous
cells with inconspicuous conidial scars (Shivas et al. 2016).
In this study, a new species, H. indica is introduced, which
was collected as a saprobe associated with leaves of Shorea
robusta Roth colonized by black moulds in India. Phylogenetic analysis from maximum likelihood based on a
combined LSU and ITS sequence dataset (Fig. 1) is provided to clarify its phylogenetic affinities within
Teratosphaeriaceae.
Hyweljonesia indica P.N. Singh & S.K. Singh, sp. nov.
MycoBank number: MB821804; Facesoffungi number:
FoF03526, Fig. 2
Etymology: The specific epithet ‘‘indica’’ refers to the
country of origin.
Holotype: AMH 9889
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Saprobic on leaves of Shorea robusta (Dipterocarpaceae) forests in terrestrial habitats. Sexual morph
Undetermined. Asexual morph Vegetative hyphae
smooth-walled, septate, subhyaline to light olivaceous, up
to 4 lm wide. Conidiophores arising from loose to
References
compact hyphal tufts, macronematous, lateral, unbranched
to rarely branched at base, 0–1-septate, straight, smoothwalled, light olivaceous, 4.5–24.5 lm long (
x = 9.6 lm,
n = 30); base flared, 3.5–8.5 lm wide (
x = 5.87 lm,
n = 30); apex narrow, cylindrical, 1.5–4.5 lm wide
(
x = 2.74 lm, n = 30). Conidiogenous cells terminal, 1(–
2), straight, smooth-walled, subhyaline to olivaceous,
cylindrical to clavate, scars inconspicuous, 5.5–12.8 9 2–
5 lm (
x = 10.3 9 3.28 lm, n = 30). Conidia acrogenous
to rarely acropleurogenous, produced in apical whorl of 1–
12 conidia, simple, aseptate, obovoid to pyriform, smoothwalled, hyaline, apex rounded, base truncate, hilum
refractive,
2.4–6.8 9 1.5–2.6 lm
(
x = 4.6 9 2 lm,
n = 30).
Culture characteristics: Colonies on PDA reaching
average 12.5 mm diam. in 12 days, after 2 weeks of
incubation at 25 °C, colonies were circular, margin regular,
smooth, and orange white (6A2). Later turning to grey
(2C1), mucoid, centre raised, umbonate, periphery white
(6A1), with abundant hyphal tufts, sulcate, up to
7500 9 132–220 lm. Reverse brownish orange (5C4),
margin smooth-walled, wrinkled.
Material examined: INDIA, Uttar Pradesh, Gorakhpur
District, on Shorea robusta (leaf infested with black mold),
5 May 2016, P.N. Singh, AMH 9889 (holotype), ex-type
living culture, NFCCI 4146 (National Fungal Culture
Collection of India-WDCM 932).
GenBank numbers: ITS = MF322773, LSU = MF322775.
Notes: Detail study of in vitro cultural characteristics
and morphology revealed a few morphological similarities
with Hyweljonesia queenslandica. However, H. indica is
distinct in having obovoid to pyriform conidia which are
significantly larger when compared to the cuneiform
123
Fungal Diversity
conidia of H. queenslandica (Fig. 2). Conidiogenous cells
of H. indica mostly arise singly from the conidiophores,
while they are produced in 1–5 whorls of H. queenslandica
(Shivas et al. 2016).
Sequence analysis of ITS and LSU positions Hyweljonesia indica in the genus Hyweljonesia closely related
to H. queenslandica with strong bootstrap support (100%
ML; Fig. 1). The BLASTn search of ITS sequence shows
95% similarity (468/491) with H. queenslandica (BRIP
61322b) and same similarity was recorded for LSU
sequence with 98% similarity (838/851). Thus following
the guidelines of Jeewon and Hyde (2016) this is a new
species. To our understanding this genus and species is
isolated and reported for the first time from India as a
saprobic black mold associated with leaves of Shorea
robusta.
Subclass Pleosporomycetidae C.L. Schoch et al.
Pleosporales Luttr. ex M.E. Barr
Fig. 1 Phylogram generated
from maximum likelihood
analysis based on the combined
ITS and LSU sequences of
representative species in
Teratosphaeriaceae. Bootstrap
support value for maximum
likelihood equal to or greater
than 50% are indicated at the
nodes. The novel species is
shown in blue. The ex-type
strains are indicated in bold.
The tree is rooted to Harknessia
ellipsoidea (CPC 13077)
123
Dictyosporiaceae Boonmee & K.D. Hyde
We follow the latest treatments and updated accounts of
Dictyosporiaceae in Boonmee et al. (2016), Wang et al.
(2016), Hyde et al. (2017), Tibpromma et al. (2018) and
Yang et al. (2018b). Recently, 12 genera were listed in this
family (Wijayawardene et al. 2018a).
Dictyocheirospora M.J. D’souza et al.
Dictyocheirospora was introduced by Boonmee et al.
(2016) with D. rotunda M.J. D’souza et al. as the type
species. Boonmee et al. (2016) included Dictyocheirospora
in the new family Dictyosporiaceae based on the fact that
Dictyocheirospora species have dark sporodochial colonies, and produce aeroaquatic cheiroid dictyospores. Many
species were subsequently accommodated in this genus
(Wang et al. 2016; Hyde et al. 2017; Tibpromma et al.
2018; Yang et al. 2018b) and 17 species are listed in Index
Fungorum (2019). In this study, Dictyocheirospora
metroxylonis Konta & K.D. Hyde, sp. nov. is introduced
from dead Metroxylon sagu (Arecaceae) in Thailand based
on morphological and multigene phylogenetic support.
Fungal Diversity
Fig. 2 Hyweljonesia indica (AMH 9889, holotype). a Lower surface
of Shorea robusta leaf showing patches of black moulds. b Colony
characteristics on PDA (front view). c Enlarged view of single colony
on PDA showing mycelial tufts. d Conidiophores bearing conidiogenous cells and whorls of conidia arising from tuft of mycelial
hyphae. e Tufts of white vegetative mycelial hyphae in stereoscopic
view. f Numerous conidiophores arising laterally from loose and
tufted mycelial hyphae. g Enlarged view of single conidiophore
bearing whorl of conidia. h Conidiophore branched at base. i Conidiophore bearing two conidiogenous cells and attached conidia.
j Obovoid to pyriform hyaline conidia with refractive conidial scars.
Scale bars d, f–j = 10 lm
Dictyocheirospora metroxylonis Konta & K.D. Hyde, sp.
nov.
Index Fungorum number: IF555290; Facesoffungi
number: FoF04833, Fig. 4
Etymology: Name reflects the host genus Metroxylon.
Holotype: MFLU 15-0028
Saprobic on dead Metroxylon sagu. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Sporodochia on natural substrate in small groups, punctiform,
100–200 lm diam. (
x = 130 lm, n = 10), velvety, greyish
123
Fungal Diversity
to dark brown. Mycelium immersed, composed of brown,
smooth, thin-walled, septate, branched hyphae. Conidiophores micronematous, pale brown, smooth, thin-walled.
Conidiogenous cells 3–8 9 3–5 lm (
x = 5.2 9 4.6 lm,
n = 10), holoblastic, integrated, terminal, determinate, pale
brown, smooth-walled. Conidia 45–69 9 15–29 lm
(
x = 61 9 20 lm, n = 20), solitary, monoblastic, acrogenous, cheiroid, pale brown, consisting of 4–6 rows of cells,
rows digitate, cylindrical, inwardly curved at the tip, arising from a basal cell, each arm composed of 9–14 cells,
distoseptate, constricted at thr septa, large guttule in each
central cell. Conidial arm 29–58 9 5–7 lm (
x = 47 9
6 lm, n = 10) (Fig. 3).
Culture characteristics: Conidia germinated on MEA
within 24 h and germ tubes produced from the basal cells
of the conidium. Colonies on MEA reaching 7–7.5 cm
diam. after 2 weeks, at 25–28 °C, initially white, becoming
grey-light brown, not sporulating on media.
Material examined: THAILAND, Krabi Province, on
dead Metroxylon sagu Rottb. (Arecaceae), 8 December
2014, S. Konta, KBR04d (MFLU 15-0028, holotype), extype living culture, MFLUCC 15-0282.
GenBank numbers: ITS = MH742321, LSU = MH742313,
SSU = MH742317, (MFLUCC 15-0282a); ITS = MH742322,
LSU = MH742314, SSU = MH742318, TEF1-a = MH764301
(MFLUCC 15-0282b); ITS = MH742323, LSU = MH742315,
SSU = MH742319, TEF1-a = MH764302 (MFLUCC
15-0282c); ITS = MH742324, LSU = MH742316, SSU =
MH742320, TEF1-a = MH764303 (MFLUCC 15-0282d).
Notes: Dictyocheirospora metroxylonis differs from
other Dictyocheirospora species by its conidial size, and
number of rows and cell numbers in each row. Phylogenetic analyses of a combined ITS, LSU, SSU and TEF1-a
sequence dataset (Fig. 3) show that D. metroxylonis forms
a distinct lineage, clustered with other Dictyocheirospora
species with moderate support in ML analysis (84% ML)
and high support in BI analysis (0.99 BYPP). Since Dictyocheirospora has been introduced in Dictyosporiaceae
(Dothideomycetes), many species were subsequently
introduced to this genus with morphological and phylogenetic evidence. Interestingly, D. metroxylonis strain
MFLUCC 150282d formed a clear zone against contaminated fungi on MEA during our experiment (Fig. 4, r).
Didymosphaeriaceae Munk
We follow the latest treatment and updated accounts of
Didymosphaeriaceae in Ariyawansa et al. (2014), Wanasinghe et al. (2018) and Tibpromma et al. (2018). There are
26 genera accepted in Didymosphaeriaceae (Wijayawardene et al. 2018a). Here we introduce a monotypic genus
Vicosamyces.
123
Vicosamyces Firmino, A.R. Machado & O.L. Pereira, gen.
nov.
MycoBank number: MB822577; Facesoffungi number:
FoF03786
Etymology: The generic epithet ‘‘Vicosamyces’’ refers to
the city ‘‘Viçosa’’, where the type was collected.
Biotrophic or necrotrophic associated with plant disease
on living leaves, forming a large, irregular, slightly raised,
rough, orange brown wound, with orange margin. Sexual
morph Ascomata immersed in orange brown wound tissue,
solitary, brown, globose to pyriform, ostiolate. Peridium
thin-walled, composed of dark brown, pseudoparenchymatous cells, of textura angularis to textura prismatica.
Hamathecium comprising numerous, cylindrical, filiform,
septate, unbranched, hyaline pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, cylindrical, subsessile to
short pedicellate, with furcate pedicel, apically rounded
with well-developed ocular chamber. Ascospores overlapping 1–2-seriate, brown, 2-celled, apiosporous, smaller at
the lower cell, subfusoid to clavate, or obovoid, narrower
towards the lower cell. Asexual morph Undetermined.
Type species: Vicosamyces venturisporus Firmino, A.R.
Machado & O.L. Pereira
Notes: Vicosamyces is introduced as a new genus based on
morphology and phylogenetic support (LSU and ITS
sequence dataset). Phylogenetic analysis of a combined
LSU and ITS sequence dataset (Fig. 5) shows the fungus
belongs to Didymosphaeriaceae, clustering with the genus
Austropleospora R.G. Shivas & L. Morin. Vicosamyces has
2-celled, apiospores, while, Austropleospora has muriform
ascospores (Morin et al. 2010; Thambugala et al. 2014;
Ariyawansa et al. 2015a). Both genera have been found as
biotrophic or necrotrophic pathogens associated with plant
disease on living leaves, or stems. However, these two
genera are associated with different symptoms on the host
tissue. Austropleospora forms subglobose ascomata, solitary or in groups, immersed in small, brown, raised necrotic
spots on Chrysanthemoides monilifera ssp. rotundata
(Asteraceae) (Morin et al. 2010; Thambugala et al. 2014).
Vicosamyces forms globose to pyriform ascomata, solitary,
immersed in large, orange-brown wound, with orange
margin on leaves of Eugenia sp. (Myrtaceae). In this study,
the phylogenetic relationship of Austropleospora and Vicosamyces was not well-resolved. Phylogenetic analysis
obtained from more informative genes will provide a better
phylogenetic relationship of these genera.
Vicosamyces venturisporus Firmino, A.R. Machado &
O.L. Pereira, sp. nov.
MycoBank number: MB822578; Facesoffungi number:
FoF03787, Fig. 6
Fungal Diversity
Fig. 3 Maximum likelihood majority rule consensus tree for the
analysed Dictyosporiaceae isolates based on a dataset of combined
ITS, LSU and TEF1-a sequence data. Bootstrap support values for
maximum likelihood (ML) and maximum parsimony (MP) greater
than 75% and Bayesian posterior probabilities greater than 0.95 are
indicated above the nodes as ML/MP/PP. Branches with 100% ML,
100% MP and 1.00 BYPP are shown as black circle at the nodes. The
tree is rooted with Periconia igniaria (CBS 379.86, CBS 845.96). The
new taxon is in red and ex-type strains are in black bold
Etymology: The specific epithet ‘‘venturisporus’’ refers
to the ascospores which are similar in shape to the ascospores of the genus Venturia.
Holotype: VIC 44320
Biotrophic or necrotrophic associated with plant disease
on living leaves, forming a large, irregular, slightly raised,
rough, orange brown wound, with orange margin. Sexual
morph Ascomata 240–340 9 250–310 lm, immersed in
orange brown wound, solitary, brown, globose to pyriform,
ostiolate. Peridium thin-walled, composed of dark brown,
pseudoparenchymatous cells, of textura angularis to textura prismatica. Hamathecium comprising 2–2.5 lm wide,
123
Fungal Diversity
Fig. 4 Dictyocheirospora metroxylonis (MFLU 15-0028, holotype).
a Sporodochia on the substrate. b–c Close up sporodochia on the
substrate. d–f Immature conidia. g–m Mature conidia. n–p
Germinating conidium. q Colony on MEA. r Colony on MEA with
clear zone against contaminated fungi. Scale bars a = 500 lm, b,
c = 100 lm, g–p = 20 lm, d–f = 10 lm
numerous, cylindrical, filiform, septate, unbranched, hyaline pseudoparaphyses. Asci 125–152.5 9 14–15 lm,
8-spored, bitunicate, fissitunicate, cylindrical, subsessile to
short pedicellate, with furcate pedicel, apically rounded
with well-developed ocular chamber. Ascospores
22.5–30 9 6–8 lm, overlapping 1–2-seriate, upper cell
brown with reddish tint, lower cell pale brown with a
reddish tint, 2-celled, apiosporous, smaller at the lower
cell, subfusoid to clavate, or obovoid, narrower towards the
lower cell, with rounded to acute ends, slightly constricted
at the septum, guttulate, smooth-walled. Asexual morph
Undetermined.
Material examined: BRAZIL, Minas Gerais, Viçosa,
Recanto das Cigarras, on leaves of Eugenia sp. (Myrtaceae), 10 September 2015, A.R. Machado (VIC 44320,
holotype).
GenBank
numbers:
MF802828 (CDA1494);
MF802829 (CDA1495);
MF802830 (CDA495).
123
ITS = MF802825,
ITS = MF802826,
ITS = MF802827,
LSU =
LSU =
LSU =
Lentitheciaceae Y. Zhang ter et al.
The family Lentitheciaceae was introduced by Zhang
et al. (2009a) with L. fluviatile (Aptroot & Van Ryck.) K.D.
Hyde as the type species. Thirteen genera are included in this
family (Wanasinghe et al. 2014a, 2018; Knapp et al. 2015;
Phookamsak et al. 2015a; Tanaka et al. 2015; Wijayawardene et al. 2015, 2018a; Dayarathne et al. 2018). We follow
the latest treatment and updated accounts of Lentitheciaceae
in Wanasinghe et al. (2014a), Wijayawardene et al. (2015),
Tibpromma et al. (2017) and Dayarathne et al. (2018). Based
on phylogenetic analysis of a combined LSU, SSU, ITS and
Fungal Diversity
Fig. 5 Bayesian inference tree obtained from the concatenated ITS
and LSU sequences including 83 taxa of representative genera in
Didymosphaeriaceae. Taxa of Pleosporaceae (Pleosporales) were
selected as the outgroup. Bayesian posterior probabilities (BYPP)
represented by percentage equal or greater than 50% are shown above
the nodes. The new isolates are in blue, ex-type strains are in bold
123
Fungal Diversity
TEF1-a sequence dataset, two novel species, Keissleriella
caraganae and Murilentithecium lonicerae are introduced.
In addition, Phragmocamarosporium hederae Wijayaw.
et al. associated with leaf spots on Cycas sp. (Cycadaceae) is
reported in Yunnan, China for the first time.
Keissleriella Höhn
We follow the latest treatment and updated accounts of
Keissleriella in Wanasinghe et al. (2018). Although 43
epithets of Keissleriella are listed in Index Fungorum
(2018), only 19 species have been confirmed in Lentitheciaceae based on molecular data (Fig. 7).
Keissleriella caraganae Chaiwan, Phookamsak, Wanas. &
K.D. Hyde, sp. nov.
Index Fungorum number: IF555523; Facesoffungi
number: FoF04965, Fig. 8
Etymology: The specific epithet ‘‘caraganae’’ refers to
the host genus Caragana, from which the holotype was
collected.
Holotype: KUN-HKAS 102236
Saprobic on Caragana arborescens (Fabaceae). Sexual
morph Ascomata 140–175 lm high, 170–235 lm diam.,
scattered, solitary or in groups, semi-immersed, visible as
raised, black dots on host surface, globose to subglobose,
Fig. 6 Vicosamyces venturisporus (VIC 44320, holotype). a, b
Symptoms on naturally infected leaf. c Globose to pyriform
pseudothecium immersed in the leaf tissue. d Immature ascus.
123
glabrous, ostiolate at centre, with minute papilla, filled with
short, brown, aseptate periphyses. Peridium 15–25 lm
wide, thin-walled, of equal thickness, composed of several layers of small, flattened, brown to dark brown
pseudoparenchymatous cells, arranged in a textura angularis
to textura prismatica, intermixed with the host cells. Hamathecium composed of dense, 2–3 lm wide, broad filamentous, distinctly septate, anastomosed pseudoparaphyses,
embedded in a hyaline gelatinous matrix. Asci 39–75 9
10–12 lm (
x = 60.1 9 11.1 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short
pedicellate, apically rounded, with well-developed ocular
chamber.
Ascospores
14–20 9 3–7 lm
(
x = 16.9 9 5.1 lm, n = 20), overlapping 1–2-seriate, pale
yellowish, fusiform to ellipsoidal, with rounded ends, (1–
)3(–4)-septate, slightly constricted at the central septum,
smooth-walled, with small guttules, surrunded by a distinct
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
22–29 mm diam. after 1 week at 20–25 °C, colony from
above, white to cream at the margin, greenish grey in the
centre; from below, white to cream at the margin, greenish
grey in the centre; medium dense, circular, slightly raised,
e Mature ascus. f Immature ascospore. g Brown and smooth-walled
ascospores. Scale bars c = 50 lm, d–g = 10 lm
Fungal Diversity
surface smooth, with edge entire, floccose to velvety, not
producing pigmentation in agar.
Material examined: CHINA, Yunnan Province, Kunming Institute of Botany, on dead hanging branch of
Caragana arborescens Lam. (Fabaceae), 2 November
2017, R. Phookamsak, KIB018 (KUN-HKAS 102236,
holotype),
ex-type
living
culture,
KUMCC
18-0163 = MFLUCC 18-0682 (KIB018A), KUMCC
18-0164 (KIB018B).
GenBank numbers: ITS = MK214368, LSU = MK214371,
SSU = MK214374,
TEF1-a = MK214377
(KUMCC
18-0163); ITS = MK359434, LSU = MK359439, SSU =
MK359444 TEF1-a = MK359073 (KUMCC 18-0164).
Notes: Keissleriella caraganae is similar to other
Keissleriella species in having ascomata with an ostiolar
neck, filled with short, brown, aseptate periphyses, bitunicate, broadly cylindrical to cylindric-clavate asci and septate ascospores, surrounded by distinct mucilaginous
sheath (Tanaka et al. 2015; Wanasinghe et al. 2018).
Multigene phylogenetic analyses (Fig. 7) show that K.
caraganae is sister to K. yonaguniensis Kaz. Tanaka & K.
Hiray. (KT2604). Although it clusters with other species of
Keissleriella and Pleurophoma Höhn. the clade is not wellresolved agreeing with previous studies (Tibpromma et al.
2017; Hyde et al. 2018b; Wanasinghe et al. 2018). Keissleriella caraganae has ellipsoidal to fusiform, pale yellowish, 3-septate ascospores, whereas K. yonaguniensis has
cylindrical, yellowish, 5-septate ascospores, with rounded
ends (Tanaka et al. 2015). Both K. caraganae and K.
rosacearum Phukhams. et al. (MFLU 15-1044) have fusiform, pale yellowish, 3-septate ascospores, but K. rosacearum was collected from Rosa canina L. (Rosaceae) in
Italy (Wanasinghe et al. 2018). Multigene phylogenetic
analysis (Fig. 7) shows that these two species form distinct
lineages in different clades.
Murilentithecium Wanas. et al.
We follow the latest treatment and updated accounts of
Murilentithecium in Wanasinghe et al. (2018). Generic
notes were also provided by Wanasinghe et al. (2014a).
Three species (including our new species) are presently
included in this genus viz. M. clematidis Wanas. et al., M.
lonicerae (in this study) and M. rosae Phukhams. et al.
(Index Fungorum 2019). These three species were collected from Clematis vitalba L. (Italy), Lonicera maackii
(Rupr.) Maxim (Yunnan, China) and Rosa canina L.
(Italy).
Murilentithecium lonicerae Phookamsak, Chaiwan,
Wanas. & K.D. Hyde, sp. nov.
Index Fungorum number: IF555524; Facesoffungi
number: FoF04966, Fig. 9
Etymology: The specific epithet ‘‘lonicerae’’ refers to
the host genus Lonicera, from which the holotype was
collected.
Holotype: KUN-HKAS 102238
Saprobic on Lonicera maackii. Sexual morph Undetermined. Asexual morph Conidiomata 95–150 lm high,
110–170 lm diam., pycnidial, semi-immersed, visible as
raised, black dots on host surface, solitary, globose to
subglobose, glabrous, uni-loculate, ostiolate at centre, with
minute papilla, lacking periphyses. Conidiomata walls 5–
15 lm diam., thin-walled, of unequal thickness, slightly
thickened at the base, composed of 5–7 layers, of flattened,
brown pseudoparenchymatous cells, slightly dark at the
apex, arranged in textura angularis to textura prismatica.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 8–15 9 (3–)4–8 lm (
x = 11 9 5.5 lm,
n = 35), enteroblastic, phialidic, rarely annellidic, discrete,
determinate, hyaline, smooth, aseptate, cylindrical to
doliiform, with narrow channel, minute collarette and
periclinal wall thickening, arising from the inner cavity of
pycnidial wall. Conidia (13.5–)14–17(–18.5) 9 7–10(–12)
lm (
x = 15.6 9 9.4 lm, n = 50), initially light brown to
pale yellowish, aseptate, becoming reddish brown to dark
brown, muriform, subglobose to obovoid, or turbinate, with
truncate base, (1–)2–4 transverse septa, with several longitudinal sectors, not constricted at the septa, smoothwalled with minute guttules.
Culture characteristics: Colonies on PDA reaching
30–35 mm diam. after 3 weeks at 20–25 °C; colony from
above, white-grey at the margin, grey at the centre; from
below, white-grey at the margin, grey to dark grey at the
centre, slightly radiated outwards colony; dense, circular,
slightly raised to umbonate, surface smooth, with edge
entire, floccose; not producing pigmentation in agar.
Material examined: CHINA, Yunnan Province, Kunming Institute of Botany, Lonicera maackii (Rupr.) Maxim.
(Caprifoliaceae), 20 April 2017, R. Phookamsak, KIB035
(KUN-HKAS 102238, holotype), ex-type living culture,
MFLUCC 18-0675 = KUMCC 18-0167 (KIB035IA),
KUMCC 18-0168 (KIB035IB), KUMCC 18-0169
(KIB035IIA), KUMCC 18-0170 (KIB035IIB).
GenBank numbers: ITS = MK214370, LSU = MK214373,
SSU = MK214376, TEF1-a = MK214379 (KUMCC 18-0167);
ITS = MK359436, LSU = MK359441, SSU = MK359446,
TEF1-a = MK359075 (KUMCC 18-0168); ITS = MK359437,
LSU = MK359442, SSU = MK359447, TEF1-a = MK359076
(KUMCC 18-0169); ITS = MK359438, LSU = MK359443,
SSU = MK359448, TEF1-a = MK359077 (KUMCC 18-0170).
Notes: Murilentithecium lonicerae can be distinguished
from M. clematidis and M. rosae in having reddish brown
to dark brown, subglobose to obovoid, or turbinate conidia,
123
Fungal Diversity
123
Fungal Diversity
b Fig. 7 Phylogram generated from maximum likelihood analysis
based on the combined LSU, SSU, ITS and TEF1-a sequence dataset
for taxa in Lentitheciaceae. Related sequences were obtained from
Wanasinghe et al. (2018). Eighty-three strains are included in the
combined sequence analyses, which comprise 3419 characters with
gaps. Single gene analyses were also performed and topology and
clade stability compared from combined gene analyses. Massarina
cisti (CBS 266.62) and M. eburnea (CBS 473.64, H3953) were used
as the outgroup taxa. Bootstrap support value for ML equal to or
greater than 60% and Bayesian posterior probabilities equal to or
greater than 0.95 BYPP are given above the nodes. Newly generated
sequences are in blue. Type strains are in bold
with truncate base, (1–)2–4 transverse septa, with several
longitudinal sectors. Murilentithecium clematidis has pale
brown to brown, oblong to clavate conidia, with 3–5
transverse septa, and 2–5 longitudinal septa (Wanasinghe
et al. 2014a; Wijayawardene et al. 2016). Murilentithecium
rosae has yellowish brown to dark brown, ovoid conidia,
with 3 transverse septa, and 1–2 longitudinal septa
(Wanasinghe et al. 2018). Multigene phylogenetic analyses
Fig. 8 Keissleriella caraganae (KUN-HKAS 102236, holotype).
a Appearance of ascomata on host surface. b, c Section through
ascomata. d, e Section through peridium. f, g Asci embedded in
cellular pseudoparaphyses (g = stained in Indian ink). h, i Asci. j–l
(Fig. 7) show that M. lonicerae forms a distinct lineage
basal to Murilentithecium.
Phragmocamarosporium Wijayaw. et al.
We follow the latest treatment and updated accounts of
Phragmocamarosporium in Wanasinghe et al. (2018).
There are only three species in this genus, P. hederae, P.
platani Wijayaw. et al. and P. rosae Wanas. et al. Phragmocamarosporium hederae and P. rosae were collected
from Hedera helix L. and Rosa canina in Europe (Germany
and Great Britain respectively). Whereas, P. platani was
found on Platanus sp. in Asia (Guizhou, China). In this
study, P. hederae is reported from China on a different
host.
Phragmocamarosporium hederae Wijayaw., R.K. Schumach. & K.D. Hyde, Index Fungorum 370: 1 (2018),
Fig. 10
Holotype: GERMANY, near Berlin, park, on a twig of
Hedera helix L. (Araliaceae), 18 May 2013, Rene Klaus
Schumacher, NNW GER 014/8 (MFLU 15-0165), living
cultures MFLUCC 13-0552, GUCC 8.
Ascospores. m Ascospores stained in Indian ink. n, o Culture on PDA
after one week (n = from above, o = from below). Scale bars
a = 200 lm, b, c = 50 lm, d–i = 20 lm, j–m = 5 lm
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Fungal Diversity
Fig. 9 Murilentithecium lonicerae (KUN-HKAS 102238, holotype).
a Appearance of conidiomata on host surface. b Section through
conidioma. c Section through conidioma wall. d–g Conidiogenous
cells and conidia. h–k Conidia. l Germinating of conidium. m, n
Culture on PDA after 1 week (m = from above, n = from below).
Scale bars a = 200 lm, b = 50 lm, c, l = 20 lm, d–k = 10 lm
Associated with leaf spots on Cycas (Cycadaceae).
Sexual morph Undetermined. Asexual morph Conidiomata 130–170 lm high, 180–270 lm diam., pycnidial,
semi-immersed, visible as raised, black dots on host surface, scattered, solitary to gregarious, globose to subglobose,
glabrous, uni-loculate, ostiolate at centre, with minute
papilla, lacking periphyses. Conidiomata walls 10–20 lm,
thin-walled, of equal thickness, composed of 3–5 layers, of
flattened, brown to dark brown, pseudoparenchymatous cells,
with blackened cells at the papilla, arranged in textura angularis to textura prismatica, difficult to distinguish from conidiogenous cells. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells (2.5–)3–5(–8) 9 (1.5–)2–5(–7)
lm (
x = 4.3 9 3.7 lm, n = 30), holoblastic, phialidic,
hyaline, smooth, aseptate, ampulliform, arising from the
inner cavity of the conidioma wall. Conidia (8–)10–13
(–14) 9 3–4 lm (
x = 12 9 4.2 lm, n = 50), initially light
brown, becoming reddish-brown to brown, oblong to
ellipsoidal, or subclavate with truncate base, 3-septate, not
constricted at the septa, smooth-walled.
Culture characteristics: Colonies on PDA reaching
10–15 mm diam. after 10 days at 25–30 °C; from above,
white to cream at the margin, grey at the centre; from
below, white to cream at the margin, black at the centre;
medium dense, circular, slightly raised, surface slightly
smooth, with edge entire, fluffy to feathery; not producing
pigmentation in agar.
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Fungal Diversity
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, associated with
leaf spots on Cycas (Cycadaceae), 5 April 2017, R.
Phookamsak, KIB020 (KUN-HKAS 102237), living culture,
KUMCC
18-0165
(KIB020A),
MFLUCC
18-0677 = KUMCC 18-0166 (KIB020B).
Known hosts and distribution: Hedera helix L. (Araliaceae; Germany) and associated with leaf spots on Cycas
(Yunnan Province, China) (Wijayawardene et al. 2015 and
this study).
GenBank numbers: ITS = MK214369, LSU = MK214372,
SSU = MK214375, TEF1-a = MK214378 (KUMCC 18-0165);
ITS = MK359434, LSU = MK359439, SSU = MK359444,
TEF1-a = MK359073 (KUMCC 18-0166).
Notes: Multigene phylogenetic analyses (Fig. 7) show
that the strain MFLUCC 18-0677 grouped with Phragmocamarosporium hederae and P. platani in Lentitheciaceae.
A BLASTn search of LSU and SSU sequence data indicates that MFLUCC 18-0677 is identical to P. hederae
(100% and 99% similarities, respectively). We therefore,
identify our isolate as P. hederae and this species was
collected from Cycas in China for the first time. Our new
isolate is similar to P. hederae in having phragmosporous
conidia. Whearas, P. platani has phragmosporous and
muriform conidia at maturity (Wijayawardene et al. 2015).
Compared to the type of P. hederae our new isolate has
shorter and broader conidiogenous cells (8–10 9
1.5–2.5 lm in the type collection) and longer conidia
(9–11 9 3–4.5 lm in the type collection). Phragmocamarosporium platani has smaller conidiogenous cells
(1.5–3 9 1.5–2.5 lm) and narrower conidia (12–13 9
5–7.5 lm) (Wijayawardene et al. 2015). Only LSU and
SSU sequence data for P. hederae and P. platani are
available in GenBank, and sequences of more informative
genes are needed to clarify species in this genus.
Leptosphaeriaceae M.E. Barr
Leptosphaeriaceae was introduced by Barr (1987) and is
typified by Leptosphaeria Ces. & De Not. to accommodate
species having immersed, subglobose, thick-walled ascomata containing interascal filamentous pseudoparaphyses,
with bitunicate, broad asci bearing fusiform, transversely
septate, hyaline to yellow-brown ascospores and
coelomycetous asexual morphs in the order Pleosporales
(Ariyawansa et al. 2015b). Ariyawansa et al. (2015b) recircumscribed the genera in Leptosphaeriaceae based on
morphological characteristics and multigene phylogenetic
analyses, and accepted ten genera with more than 140
species. This is in agreement of the taxonomic outline of
Ascomycota, provided by Wijayawardene et al. (2018a)
and the notes of each genus in this family were provided by
Ariyawansa et al. (2015b) and Wijayawardene et al.
(2017a).
We follow the latest treatment of Leptosphaeriaceae in
Ariyawansa et al. (2015b) and updated accounts of taxa in
Leptosphaeriaceae in Hyde et al. (2016, 2017), Tennakoon
et al. (2017) and Tibpromma et al. (2017). In this paper, we
introduce four new species, Leptosphaeria urticae, Plenodomus artemisiae, P. lijiangensis and Sphaerellopsis
isthmospora in Leptosphaeriaceae. The asexual morph of
Plenodomus sinensis is also introduced from a fern in
China and a new host record of Sphaerellopsis paraphysata
associated with rust on living leaves of Liriope spicata
(Thunb.) Lour (Asparagaceae) is reported.
Leptosphaeria Ces. & De Not.
Leptosphaeria was introduced by Cesati and De Notaris
(1863) and is typified by L. doliolum (Pers.) Ces. & De Not.
(lectotype designated by Shearer et al. 1990). The genus is
characterized by semi-immersed to erumpent, coriaceous
ascomata, which become superficial, a thick-walled peridium composed of scleroplectenchymatous cells, cylindrical
to cylindric-clavate asci, reddish to yellowish brown,
ellipsoidal to fusiform, septate ascospores and
coelomycetous coniothyrium-like and phoma-like asexual
morphs (Ariyawansa et al. 2015b; Dayarathne et al. 2015).
Taxonomic revision of the genus was discussed in Ariyawansa et al. (2015b). Over 1600 epithets are listed for
Leptosphaeria (Index Fungorum 2019), but few species
have been confirmed by phylogenetic analysis. Most Leptosphaeria species lack molecular data to clarify their
phylogenetic placements. Some other Leptosphaeria sensu
lato species have been treated in different genera in Leptosphaeriaceae and other related families (de Gruyter et al.
2013; Ariyawansa et al. 2015b).
Leptosphaeria urticae D. Pem, E.B.G. Jones & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555597; Facesoffungi
number: FoF04370, Fig. 11
Etymology: Name reflects the host from which the fungus was isolated.
Holotype: MFLU 18-0591
Saprobic on dead branches of Urtica dioica. Sexual
morph Ascomata 100–130 high, 70–110 lm diam., solitary, scattered or in small groups, erumpent through host
epidermis to superficial, conical to mammiform, dark
brown to black, coriaceous, smooth, easily removed from
the host substrate, ostiolate with minute papilla. Ostioles
50–70 lm diam., papillate, black, shiny, smooth. Peridium
25–50 lm wide, comprising two cell types, outer layer
composed of small, thick-walled cells of textura angularis
to textura globulosa, surface heavily pigmented termed as
scleroplectenchyma, thinner at the apex, wide at sides,
inner layer composed of subhyaline or light brown
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Fungal Diversity
Fig. 10 Phragmocamarosporium hederae (KUN-HKAS 102237).
a Symptom of leaf spot disease on Cycas. b Appearance of
conidiomata on host surface. c Section through conidioma.
d Section through conidioma wall. e–g Conidiogenous cells with
conidia. h–l Conidia. Scale bars c = 50 lm, d, h = 10 lm, e–g, i–
l = 5 lm
relatively thin-walled cells of textura angularis, cells near
the base comparatively larger. Hamathecium comprising
numerous, dense, 1.5–2 lm wide, filamentous, septate,
cellular pseudoparaphyses, branched and anastomosing,
embedded in gelatinous matrix. Asci 60–140 9 9–11 lm
(
x = 104.5 9 10 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, rounded at the apex, pedicellate,
numerous, with ocular chamber. Ascospores 35–40 9 4–
6 lm (
x = 38.3 9 5.2 lm, n = 20), overlapping 1–2-seriate, initially hyaline, becoming yellowish brown at maturity, long fusiform, (8–)9-septate, constricted at the septa,
narrowly rounded at both ends, smooth-walled, lacking a
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 24 h. Colonies growing on MEA reaching
2 cm diam. in one week at 16 °C. Mycelium superficial,
surface smooth, irregular, slightly raised, edge crenate,
velutinous, from above white to pale yellow, reverse
yellow.
Material examined: UK, Sussex, Singleton, on dead
stem of Urtica dioica L. (Urticaceae), 5 April 2017, E.B
Gareth Jones, 353 UK (MFLU 18-0591, holotype), ex-type
living culture MFLUCC 17-2302.
GenBank
numbers:
ITS = MK123333,
LSU =
MK123332, SSU = MK123329, TEF1-a = MH028391.
Notes: Phylogenetic analyses of a combined LSU, SSU
and ITS sequence dataset (Fig. 12) reveal that Leptosphaeria urticae (MFLU 18-0591) is sister to L. italica
Dayar. et al. (MFLU 15-0174). Leptosphaeria urticae differs from L. italica in having longer asci (L. urticae,
60–140 9 9–11 lm versus 60–112 9 7–12 lm, L. italica), longer ascospores (L. urticae, 35–40 9 4–6 lm
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Fungal Diversity
versus 12–18 9 4–6 lm, L. italica) and more ascospore
septation (L. urticae, (8–)9-septate versus 3-septate, L.
italica) (Dayarathne et al. 2015). Based on the NCBI
BLASTn search of ITS sequence data, L. urticae has 96%
similarity with L. sclerotioides (Preuss ex Sacc.) Gruyter
et al. However, the two species cannot be compared as L.
urticae is represented only by its sexual morph, whereas, L.
sclerotioides is known only by its asexual morph (de
Gruyter et al. 2013). Phylogenetic analysis indicates that
these two species are not conspecific. Hence, we introduce
L. urticae as a new species based on both morphological
and molecular data.
Five Leptosphaeria species have been reported from
Urtica: L. acuta (Fuckel) P. Karst., L. acutiuscula Berl., L.
atropurpurea Petr., L. doliolum (Pers.) Ces. & De Not. and
L. ogilviensis (Berk. & Broome) Ces. & De Not. (Shoemaker 1984; Farr and Rossman 2018). These species can
be distinguished from each other based on ascospore
septation.
Plenodomus Preuss
Plenodomus was introduced by Preuss (1851) and is
typified by P. rabenhorstii. Subsequently, Boerema and
Kesteren (1964) designated P. lingam (Tode) Höhn. as the
type combination over P. rabenhorstii because the type
material of P. rabenhorstii was lost during the World War
II (de Gruyter et al. 2013; Ariyawansa et al. 2015b;
Tennakoon et al. 2017). Based on molecular phylogeny, de
Gruyter et al. (2013) reclassified Phoma section
Plenodomus and synonymized species in Phoma
section Plenodomus under the genus Plenodomus in
Leptosphaeriaceae. The genus was re-circumscribed by
Ariyawansa et al. (2015b) based on study of type and
representative specimens coupled with molecular data.
Marin-Felix et al. (2017) and Tennakoon et al. (2017)
updated the accounts of Plenodomus based on molecular
data. There are 97 epithets available in Index Fungorum
(2019).
Plenodomus artemisiae A. Karunarathna, Phookamsak &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556118; Facesoffungi
number: FoF05696, Fig. 13
Etymology: The specific epithet ‘‘artemisiae’’ refers to
the host genus Artemisia, on which the type species was
collected.
Holotype: KUN-HKAS 102226.
Saprobic on dead branches and stems of Artemisia sp.
Sexual morph Ascomata 140–280 lm high, 300–450 lm
diam., black, shiny on the host surface, solitary to aggregated, immersed at the base, becoming superficial, uniloculate, subglobose to irregular in shape, with truncate
base, glabrous, ostiolate, papillate. Ostioles central, dark
brown, beak-like papilla, ostiolar canal filled with periphyses. Peridium 10–85 lm wide, thick-walled of unequal
thickness, thickened at the based, slightly thin at the apex,
composed of several cell layers of dark brown scleroplectenchymatous cells, arranged in a textura angularis to
textura globulosa. Hamathecium composed of hyaline,
filamentous, 2–4 lm wide, distinctly septate pseudoparaphyses, anastomosing, embedded in a hyaline gelatinous
matrix.
Asci
(64–)70–90(–100) 9 (9.5–)10–13 lm
(
x = 82.7 9 11.2 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical, short, pedicellate, apically rounded
with a distinct ocular chamber. Ascospores (28–)30–
40 9 (4.5–)5–6 lm (
x = 34.4 9 5.5 lm, n = 30), overlapping 2–3-seriate, pale brown, fusiform, 5-septate,
slightly constricted at the septa, enlarge at the third cell
from above, lacking a mucilaginous sheath and appendages. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
30–33 mm diam. after 4 weeks. Colony dense, circular,
low convex, surface smooth, with edge entire, floccose;
from above white; from below, yellowish-grey at the edge,
with white to cream margin, dark yellowish at the centre,
slightly radiating outwards colony; not produced pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead branches
and stems of Artemisia sp., 20 December 2015, R.
Phookamsak, AS003 (KUN-HKAS 102226, holotype). extype living culture, KUMCC 18-0151.
GenBank
numbers:
ITS = MK387920,
LSU =
MK387958, SSU = MK387928, TEF1-a = MK435600,
RPB2 = MK435607 (KUMCC 18-0151).
Notes: Plenodomus artemisiae forms a distinct lineage
and is sister to P. salviae Thambug. et al. (MFLUCC
13-0219) with high support (99% ML and 0.99 BYPP;
Fig. 12). Plenodomus artemisiae can be distinguished from
P. salviae in having shorter and broader, pale brown,
fusiform ascospores, enlarged at the third cell from above.
Plenodomus salviae has longer and thinner, yellowish
brown,
cylindric-fusiform
ascospores
(30–48 9
3.1–4.3 lm), and without the enlarged cell (Ariyawansa
et al. 2015b). Plenodomus artemisiae is also similar to
Leptosphaeria artemisiae (Fuckel) Auersw. in having
5-septate ascospores, with the enlarged third cell and
occurring on Artemisia. However, L. artemisiae has larger
ascomata and narrower ascospores (32–37 9 8.5–10;
Shoemaker 1984). Furthermore, P. artemisiae has pale
brown, fusiform ascospores, whereas, L. artemisiae has
light reddish brown, broadly elliptical ascospores (Shoemaker 1984). Phylogenetic affinity of L. artemisiae could
not be resolved due to lack of molecular data.
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Fungal Diversity
Fig. 11 Leptosphaeria urticae (MFLU 18-0591, holotype). a–c
Appearance of ascomata on host surface. d Section through an
ascoma. e Peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores.
n Spore germination on MEA after 24 h. o Culture from above and
below. Scale bars a, b = 500 lm, c = 200 lm, d = 100 lm, e, g–
i = 50 lm, j–m = 10 lm, f = 5 lm
Plenodomus lijiangensis Phookamsak, A. Karunarathna &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556137; Facesoffungi
number: FoF05697, Fig. 14
Etymology: The specific epithet ‘‘lijiangensis’’ refers to
Lijiang prefecture-level city, of Yunnan Province, China
where the holotype was collected.
Holotype: KUN-HKAS 102249
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Fungal Diversity
Saprobic on dead fronds of fern. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
140–290 lm high, 135–240 lm diam., pycnidial, black,
scattered, gregarious, superficial, uni-loculate, varied in
shape, subconical to ovoid, or subglobose, with truncate
base, widest at the base, glabrous, with indistinct ostiole.
Conidiomata walls 17–100 lm wide, thick-walled, of
unequal thickness, thickened at the apex, comprising several cell layers, outer layer composed of broad, dark brown
to black, scleroplectenchymatous cells of textura angularis
to textura globulosa, inner layer composed of broad, hyaline cells of textura angularis. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 4–9 9 5–8 lm
(
x = 6.4 9 6.5 lm, n = 40), enteroblastic, phialidic, discrete, determinate, ampulliform to doliiform, hyaline,
smooth, with minute collarette, with 1–2 apertures, and
periclinal wall thickening, arising from the inner cavity of
the conidioma wall. Conidia 3–5 9 1.7–2.3 lm
(
x = 4.3 9 2 lm, n = 50), hyaline, oblong to obovoid,
aseptate, smooth-walled, with 1–2 guttules.
Culture characteristics: Colonies on PDA, reaching 57–
58 mm diam. after 3 weeks. Colony dense, circular, flattened,
slightly raised, surface smooth, with edge entire, floccose;
from above white at the margin, cream at the centre, with pale
grey concentric ring near the margin; from below, yellowish
brown at the edge, with paler margin, dark brown to black
at the centre, colony slightly radiating outwards; not producing pigmentation on agar medium. Sporulation on PDA
after three months. Conidiomata 120–250 lm high, 130–
230 lm diam., scattered, solitary to gregarious, semi-immersed in culture colony, or embedded in agar medium,
perithecial, pycnidial, with short stipe (19–49 lm long),
black, glabrous, globose to subglobose, lacking ostioles.
Conidiomata walls 3–8 lm wide, thin-walled, equally
thick, comprising 1–2 cell layers of dark brown to black
pseudoparenchymatous cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2–3 9 1.5–3 lm (
x = 2.5 9 2.2 lm, n = 20), enteroblastic, phialidic, discrete, determinate, oblong to pyriform, hyaline, with minute collarette, and periclinal wall
thickening, arising from the inner cavity of the conidioma
wall, difficult to distinguish from the conidioma wall.
Conidia 3–4(–4.5) 9 1.5–2.5 lm (
x = 3.9 9 2, n = 50),
hyaline, oblong to ellipsoidal, or obovoid, aseptate, smoothwalled, with 1–2 small guttules.
Material examined: CHINA, Yunnan Province, Lijiang,
Yulong, on dead fronds of fern, 1 August 2015, R.
Phookamsak, LJ003 (KUN-HKAS 102249, holotype), extype living culture, KUMCC 18-0186.
GenBank numbers: ITS = MK387921, LSU = MK387959,
SSU = MK387929, TEF1-a = MK435601 (KUMCC 18-0186).
Notes: Based on the NCBI BLASTn search of ITS
sequence data, Plenodomus lijiangensis closest match is P.
deqinensis Qian Chen & L. Cai (CGMCC 3.18221; 98%
similarity). Phylogenetic analyses of a concatenated LSU,
SSU and ITS sequence dataset (Fig. 12) reveal that P.
lijiangensis forms a sister lineage with P. deqinensis and
groups with P. agnitus (Desm.) Gruyter et al., P. fallaciosus (Berl.) Gruyter et al. and P. lupini (Ellis & Everh.)
Gruyter et al. Plenodomus lijiangensis shares a size range
of conidia and conidiogenous cells with P. deqinensis and
was also collected from Yunnan, China (Marin-Felix et al.
2017). However, P. lijiangensis was isolated from dead
fronds of fern, while P. deqinensis was isolated from soil.
In vitro, P. lijiangensis forms a globose to subglobose
conidiomata, inconspicuous ostiole, with a short stipe
which is similar to the asexual morph of P. sinensis Tennakoon et al. (Fig. 16). While, P. deqinensis forms globose
to subglobose, slightly papillate ostiole with a narrow pore
or opening via a rupture (Marin-Felix et al. 2017). A
comparison of ITS sequence shows that P. lijiangensis
differs from P. deqinensis in eight base positions (1.55%/
517 bp). According to the guidelines in Jeewon and Hyde
(2016), we introduce P. lijiangensis as a new species.
Plenodomus sinensis Tennakoon, Phookamsak & K.D.
Hyde, in Tennakoon et al., Phytotaxa 324(1): 76 (2017),
Figs. 15, 16
Holotype: CHINA, Yunnan Province, Xishuangbanna,
Nabanhe, dead branch of Tamarindus indica (Fabaceae),
25 November 2015, D.S. Tennakoon, DXH 015 (MFLU
17-0767).
Saprobic on dead fronds of ferns and dead stems of
Cirsium sp. Sexual morph Ascomata 250–290 lm high,
300–360 lm diam., black, shiny, scattered, gregarious,
semi-immersed to erumpent through host epidermis, subglobose to subconical, uni-loculate, glabrous, ostiolate.
Peridium thick-walled of unequal thickness, thickened at
base, thinner toward sides and apex, composed of three
type cell layers, inner layer 5–20 lm wide, comprising 2–3
strata of flattened, pale brown, thin-walled, pseudoparenchymatous cells, arranged in textura angularis to
textura prismatica, middle layer 25–100 lm wide, comprising several strata, of hyaline, thick-walled, scleroplectenchymatous cells of textura angularis to textura
globulosa, outer layer thin-walled, comprising 1 stratum, of
black, coriaceous cells of textura angularis. Hamathecium
comprising filamentous, septate, 2–4 lm wide, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (75–)80–95(–107) 9 (8.5–)9–11(–12)
lm (
x = 88.1 9 10.3, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, subsessile to short
pedicellate, with knob-like to truncate pedicel, apically
rounded, with well-developed ocular chamber. Ascospores
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Fungal Diversity
123
Fungal Diversity
b Fig. 12 Phylogram generated from maximum likelihood analysis
based on a combined LSU, SSU and ITS sequence dataset of taxa in
Leptosphaeriaceae. The updated sequence data was derived from
Tennakoon et al. (2017). Seventy strains are included in the combined
sequence analyses. Single gene analyses were also performed and
topology and clade stability compared from combined gene analyses.
Phaeosphaeria oryzae (CBS 110110) and Phaeosphaeriopsis glaucopunctata (MFLUCC 13-0265) and Paraphoma radicina (CBS 111.79)
were used as the outgroup taxa. Bootstrap support values for ML
equal to or greater than 60% and Bayesian posterior probabilities
equal to or greater than 0.80 BYPP are indicated at the nodes. Newly
generated sequences are in blue and ex-type strains are in bold
30–39 9 4–6(–6.5) lm (
x = 34.1 9 5.3, n = 40), overlapping 2–3-seriate, fusiform, initially hyaline, becoming
pale brown to pale yellowish at maturity, 6-septate, widest
at the third cell, slightly constricted at the septa, deeply
constricted at the third septum from above, smooth-walled,
inconspicuous minute appendages at both end cells.
Asexual morph Coelomycetous. Conidiomata 50–120 lm
high, 50–110 lm diam., pycnidial, black, shiny, scattered,
gregarious, superficial, uni-loculate, globose to subglobose,
with short stipe (5–10 9 7–11 lm), glabrous, ostiole central, with pore-like opening, apapillate. Conidiomata walls
5–10 lm wide, thin-walled, of equal thickness, comprising
2–3 cell layers, of dark brown pseudoparenchymatous
cells, of textura angularis. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells (3–)4–7
(–8) 9 4–6(–8) lm (
x = 5.6 9 5.1 lm, n = 30), enteroblastic, phialidic, discrete, determinate, ampulliform to
doliiform, hyaline, collarette, and periclinal wall thickening, arising from the inner cavity of the conidioma wall.
Conidia (2.7–)3–4 9 1–2 lm (
x = 3.8 9 1.4, n = 100),
hyaline, oblong, slightly curved, aseptate, smooth-walled.
Culture characteristics: Colonies on PDA reaching
28–30 mm diam. after 4 weeks at room temperature. Colony dense, irregular in shape, slightly raised to low convex,
surface smooth, edge undulate, with margin well-defined;
from above dark grey; from below, black; not produced
pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Lijiang,
Yulong, on dead fronds of fern, 29 July 2015, R.
Phookamsak, LJ001 (KUN-HKAS 102229, sexual morph),
living culture, KUMCC 18-0153; ibid., Baoshan, Shuizai,
Dawazi mountain, on dead fronds of fern, 22 October 2015,
I.D. Goonasekara, BS010 (KUN-HKAS 102228, asexual
morph), living culture, KUMCC 18-0152; Baoshan, Shuizai, Dawazi mountain, on dead stems of Cirsium sp.
(Asteraceae), 22 October 2015, R. Phookamsak, BS023
(KUN-HKAS 102227).
Known hosts and distribution: Plukenetia volubilis L.
(Euphorbiaceae) Tamarindus indica L. (Fabaceae)
(Xishuangbanna, China) (Tennakoon et al. 2017).
GenBank numbers: ITS = MK387922, LSU = MK387960,
SSU = MK387930, TEF1-a = MK435602, RPB2 = MK435608
(KUMCC 18-0153); ITS = MK387923, LSU = MK387961,
SSU = MK387931, TEF1-a = MK435603 (KUMCC 18-0152);
ITS = MK387924, LSU = MK387962, SSU = MK387932
(KUN-HKAS 102227).
Notes: Based on the NCBI BLASTn search of ITS
sequences, our isolates (KUMCC 18-0152, KUMCC
18-0153 and KUN-HKAS 102227) match with Plenodomus sinensis Tennakoon et al. (MFLU 17-0757), with
99% similarity. The sexual morph of KUMCC 18-0153 and
KUN-HKAS 102227 share similar size of ascomata, asci
and ascospores with the type, as well as sharing similar
ascospore characters with fusiform, 6-septate ascospores
(Tennakoon et al. 2017). Phylogenetic analyses of a concatenated LSU, SSU and ITS sequence dataset (Fig. 12)
reveal that our isolates cluster with P. sinensis (MFLU170757) with moderate support (86% ML and 0.90 BYPP).
The asexual morph of P. sinensis, which is reported for
the first time in this study, is similar to the asexual morph
of P. lijiangensis in having globose to subglobose conidiomata with a short stipe. However, these two species are
phylogenetically distinct.
Tennakoon et al. (2017) introduced Plenodomus sinensis
as a saprobic species occurring on Plukenetia volubilis and
Tamarindus indica from Xishuangbanna, Yunnan, China
(tropical rain forest climate). In this study, P. sinensis was
found on ferns and Cirsium sp. from Baoshan (mild subtropical highland climate) and Lijiang (a mild, with abundant rainfall and plenty of sunshine climate), Yunnan,
China. This indicates that P. sinensis may occur on a wide
range of hosts and in different climatic regions.
Sphaerellopsis Cooke
Sphaerellopsis was introduced by Sutton (1977) to
accommodate mycoparasitic taxa occurring on a wide
range of rusts and is typified with S. filum (Biv.) B. Sutton.
Sphaerellopsis was re-circumscribed by Trakunyingcharoen et al. (2014) and Ariyawansa et al. (2015b)
based on molecular phylogeny. The link between the sexual genus Eudarluca Speg. and the asexual genus
Sphaerellopsis is still debated. Eudarluca was synonymized under Sphaerellopsis by Rossman et al. (2015)
based on holomorphic characters of Eudarluca caricis (Fr.)
O.E. Erikss. However, Phookamsak et al. (2014b) re-examined the isotype specimen of Eudarluca australis Speg.
and treated Eudarluca in Phaeosphaeriaceae according to
the generic type, E. australis is not congeneric with E.
caricis. Eudarluca australis is typical of Phaeosphaeriaceae in having uni-loculate ascomata, a thin-walled
peridium, comprising 1–2 layers of brown, pseudoparenchymatous cells, cylindrical asci, with pale brown,
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Fungal Diversity
Fig. 13 Plenodomus artemisiae (KUN-HKAS 102226, holotype).
a Ascomata on host. b, c Vertical section of ascomata. d Ostiole.
e Section through peridium. f Pseudoparaphyses. g–i Asci. j–m
Ascospores. n Ascospore germination. o, p Culture characteristics
(o = from above, p = from below). Scale bars a = 500 lm, b,
c = 100 lm, d, e = 50 lm, f–i, n = 20 lm, j–m = 10 lm
ellipsoidal to fusiform, (1–)2-septate ascospores and this
concurs with the iconotype of E. australis, established by
Spegazzini (1908) (Phookamsak et al. 2014b). Whereas, E.
caricis forms black, multi-loculate ascostroma, with thickwalled peridium (Yuan et al. 1998; confirming the connection of E. caricis and Sphaerellopsis filum). Phylogenetic affinity of Eudarluca australis has not been proved
yet.
Sphaerellopsis isthmospora A. Karunarathna, Phookamsak
& K.D. Hyde, sp. nov.
Index Fungorum number: IF556138; Facesoffungi
number: FoF05698, Fig. 17
Etymology: The specific epithet ‘‘isthmospora’’ refers to
the fungus having isthmospores.
Holotype: KUN-HKAS 102225
Saprobic on dead branches of herbaceous plant. Sexual
morph Ascomata 230–330 lm high, 260–510 lm diam.,
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Fungal Diversity
Fig. 14 Plenodomus lijiangensis (KUN-HKAS 102249, holotype).
a–i Morphological characteristics on natural substrate. l–u Morphological characteristics in vitro. a Conidiomata on host. b Vertical
section of conidioma. c Section through conidioma wall. d–f
Conidiogenous cells. g–i Conidia. j, k Culture characteristics on
PDA (j = from above, k = from below). l–n Conidiomata forming on
PDA after three months. o Squash mount of conidioma. p Vertical
section of conidioma. q Section through conidioma wall stained with
congo red. r, s Conidiogenous cells. t Conidiogenous cells stained
with congo red. u Conidia. Scale bars a = 200 lm, b, n, o = 100 lm,
c, p = 50 lm, u = 10 lm, g, q–t = 5 lm, h, i = 2 lm
123
Fungal Diversity
black, shiny, scattered, solitary to gregarious, erumpent
through host epidermis, becoming semi-immersed to
superficial, varied in shape, subglobose to mammiform,
with flattened, quadrilateral, truncate base, uni-loculate,
glabrous, ostiolate, minutely papillate. Ostioles central,
with obtuse, minute papilla, dark brown to black, ostiolar
canal filled with periphyses. Peridium 18–60 lm wide,
thick-walled of unequal thickness, thicker at the sides towards
the apex, with flattened base, comprising several cell layers of
black, coriaceous, pseudoparenchymatous cells, arranged in
textura angularis to textura prismatica. Hamathecium initially comprising 2–5 lm wide, hyaline, filamentous distinctly septate pseudoparaphyses, laterally becoming,
6–18 lm wide, broadly cellular, hyaline, septate catenophyses, deeply constricted at the septa. Asci (120–)130–150
(–165) 9 (15–)19–23(–28) lm (
x = 141.4 9 21.6 lm, n =
20), 8-spored, bitunicate, fissitunicate, subcylindric-clavate,
subsessile to short pedicellate, with truncate pedicel, apically
rounded with well-developed ocular chamber. Ascospores
(65–)75–95(–118) 9 4–7 lm (
x = 87.1 9 5.9 lm, n = 30),
isthmosporous, overlapping 2–3-seriate, hyaline to yellowish, elongate cylindrical to subcylindric-clavate, bent at the
8th septum, 10–12-septate, slightly constricted at the
septa, deeply constricted at the 8th septum, split into two
part-spores; upper part 40–70(–82) lm long, 5–7-septate,
cylindrical, with rounded end; lower part 28–35(–50) lm
long, 2–3-septate, subcylindric-clavate, with acute end,
guttulate, lacking a mucilaginous sheath. Asexual morph
Undetermined.
Material examined: CHINA, Yunnan Province,
Baoshan, Shuizai, Dawazi mountain, on dead branches of
herbaceous plant, 23 October 2015, R. Phookamsak,
BS012 (KUN-HKAS 102225, holotype).
GenBank numbers: ITS = MK387925, LSU = MK387963,
SSU = MK387933, TEF1-a = MK435604 (HKAS 102225A);
ITS = MK387926, LSU = MK387964, SSU = MK387934,
TEF1-a = MK435605 (HKAS 102225B).
Notes: Sphaerellopsis isthmospora forms a phylogenetically distinct lineage, but clusters with other Sphaerellopsis
species in Leptosphaeriaceae (Fig. 12). Sphaerellopsis
isthmospora can be distinguished from other Sphaerellopsis
species in its sexual morph having isthmosporous ascospores, with 10–12-septate, deeply constricted and bent at the
8th septum. Sphaerellopsis filum (sexual morph: Eudarluca
caricis) has spindle-shaped, slightly inequilateral, 2–3-septate ascospores (Yuan et al. 1998). Based on morphological
difference and phylogenetic affinity, we therefore, introduce
a new species S. isthmospora from herbaceous plant in
Baoshan, China.
123
Sphaerellopsis paraphysata Crous & Alfenas, in Trakunyingcharoen et al., IMA Fungus 5(2): 411 (2014)
Facesoffungi number: FoF04968, Fig. 18
Holotype: BRAZIL, Minas Gerais, Viçosa, Universidade Federal de Viçosa campus, on rust on Pennisetum sp.,
18 November 2012, A.C. Alfenas, CBS H-21848, ex-type
living culture, CPC 21841 = CBS138579.
Associated with rust on living leaves of Liriope spicata
(Thunb.) Lour. Sexual morph Undetermined. Asexual
morph Coelomycetous. Conidiomata 105–160 lm high,
90–150 lm diam., black, pycnidial, solitary, associated
with rust stromatic along the leaf veins, semi-immersed to
superficial on stromata, globose to subglobose, uni-loculate, glabrous, ostiole central, with pore-like opening.
Conidiomata walls 12–30 lm wide, composed of 2–5
layers, of dark brown pseudoparenchymatous cells of textura angularis. Conidiophores 15–21 9 2.4–3 lm
(
x = 18 9 2.7 lm, n = 20), arising from the basal cavity,
1–2-celled, hyaline, curved, cylindrical, or reduced to
conidiogenous cells. Conidiogenous cells 3.5–6 9 2.5–
4.5 lm (
x = 4.7 9 3.5 lm, n = 20), enteroblastic, phialidic, discrete, determinate, cylindrical to ampulliform to
doliiform, hyaline, 0–1-septate, smooth, thin-walled, minute
collarette, with 1–2 apertures, and periclinal wall thickening.
Conidia 14–17 9 3–5 lm (
x = 15.5 9 4.5 lm, n = 20),
hyaline, fusiform to ellipsoidal, mostly 1–3-septate, constricted at the central septum, smooth-walled.
Culture characteristics: Colonies on PDA reaching
25–30 mm diam. after 4 weeks at room temperature. Colony dense, irregular in shape, flattened, slightly raised,
surface slightly rough, heaped and folded at the centre,
with small granular and black, stromatic, edge undulate,
with margin well-defined, felted at the centre, fluffy at the
edge; from above white at the margin, with yellowish grey
to greenish grey at the centre; from below, white to cream
at the margin, black at the centre; not producing pigmentation on agar medium, sporulating on PDA after 3 weeks.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, associated with
rust on living leaves of Liriope spicata (Thunb.) Lour
(Asparagaceae), 6 December 2017, R. Phookamsak,
KIB044 (KUN-HKAS 101483), living culture, KUMCC
18-0195.
Known hosts and distribution: Associated with rust on
Pennisetum sp. (Brazil), on Ravenelia macowania on
Vachellia karroo (South Africa), on leaves of Phragmites
sp. (Australia), and associated with rust on living leaves of
Liriope spicata (Yunnan, China) (Trakunyingcharoen et al.
2014; Crous et al. 2018; this study).
GenBank numbers: ITS = MK387927, LSU = MK387965,
SSU = MK387935, TEF1-a = MK435606.
Fungal Diversity
Fig. 15 Plenodomus sinensis (KUN-HKAS 102229, sexual morph).
a Ascomata on host. b Vertical section of ascoma. c Section through
peridium. d Pseudoparaphyses. e–g Ascospores. h Ascospore
germination. i–k Asci. Scale bars a = 200 lm, b = 100 lm,
c = 50 lm, d, i–k = 20 lm, e–h = 10 lm
Notes: Sphaerellopsis paraphysata was introduced by
Trakunyingcharoen et al. (2014) based on morphological
comparisons and phylogenetic analysis. We made a new
collection from China associated with a rust on living
leaves of Liriope spicata. The new isolate (KUN-HKAS
101483) is similar in morphology with S. paraphysata but
differs from the type of S. paraphysata in having smaller
conidiomata, presence of conidiophores and lacking paraphyses. Phylogenetic analyses of a combined LSU, SSU
and ITS sequence dataset show that our strain (KUMCC
18-0195) forms a sister lineage with S. paraphysata (CPC
21841) with high support (100% ML and 1.00 BYPP). A
comparison of ITS nucleotide base shows that our new
isolate has same base pairs with the type strain of S.
paraphysata. Thus we identify the new isolate as S.
paraphysata and our new collection is a new host record in
China.
Lophiotremataceae K. Hiray. & Kaz. Tanaka
Lophiotremataceae was introduced by Hirayama and
Tanaka (2011) to accommodate the type genus Lophiotrema Sacc. and is typified by L. nucula (Fr.) Sacc.
Lophiotrema shares morphological characters with
Lophiostoma due to its compressed carbonaceous ascomata
with crest-like apex but is distinguished by peridial
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Fungal Diversity
Fig. 16 Plenodomus sinensis (KUN-HKAS 102228, asexual
morph). a Conidiomata on host. b Squash mount of conidioma.
c Squash mount of conidioma showing ostiole. d Section through
conidioma. e Section through conidioma wall. f Stalk of conidioma.
g–i Conidiogenous cells. j–n Conidia. Scale bars a = 500 lm,
c = 50 lm, b, d = 20 lm, e, f, j = 10 lm, g–i = 5 lm, k–n = 2 lm
structure and shape of asci (Zhang et al. 2009b, 2012;
Hirayama and Tanaka 2011; Hyde et al. 2013). We follow
the latest treatment and the updated accounts of
Lophiotremataceae in Hyde et al. (2016) and Hashimoto
et al. (2017). Lophiotremataceae comprises Atrocalyx A.
Hashim. & Kaz. Tanaka, Crassimassarina A. Hashim. &
Kaz. Tanaka, Cryptoclypeus A. Hashim. & Kaz. Tanaka,
Galeaticarpa A. Hashim. & Kaz. Tanaka, Lophiotrema and
Pseudocryptoclypeus A. Hashim. & Kaz. Tanaka (Hashimoto et al. 2017; Wijayawardene et al. 2018a). In the
present study, a new species, Lophiotrema mucilaginosis
collected on dead wood in China, is introduced.
a short stipe, hyaline, ellipsoidal to fusiform, septate
ascospores and pycnidial coelomycetous asexual morphs
(Hirayama and Tanaka 2011; Hirayama et al. 2014;
Hashimoto et al. 2017). The genus has a long taxonomic
history and has always been confused with Lophiostoma
Ces. & De Not. and Massarina Sacc. in previous studies
(Zhang et al. 2009b, 2012; Hirayama and Tanaka 2011;
Hyde et al. 2013; Hirayama et al. 2014). However,
Lophiotrema was re-classified based on molecular data by
Hashimoto et al. (2017) and many species of Lophiotrema
sensu lato were treated as new genera in
Lophiotremataceae.
Lophiotrema Sacc.
Lophiotrema was introduced by Saccardo (1878) and is
typified by L. nucula (Fr.) Ces. & De Not. The genus was
established to accommodate taxa in Pleosporales, characterized by ascomata with a slit-like ostioles, a peridium of
uniform thickness, cylindrical to cylindric-clavate asci with
Lophiotrema mucilaginosis M. Raza & L. Cai, sp. nov.
Index Fungorum number: IF555333; Facesoffungi
number: FoF04941, Fig. 19
Etymology: In reference to the mucilaginous sheath
around spores.
Holotype: HMAS 255437
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Fungal Diversity
Saprobic on dead wood. Sexual morph Ascomata 220–
340 lm high, 240–400 lm diam., black, scattered, solitary,
semi-immersed to erumpent through host surface, conical
to mammiform, with flattened base, uni-loculate, glabrous,
ostiolate, papillate. Ostioles 50–90 9 25–60 lm, apically
with crest-like papilla, filled with periphyses, carbonaceous, with beak-like opening. Peridium 55–80 lm wide,
outer layer thick, composed of dark, coriaceous, pseudoparenchymatous cells of textura epidermoidea, inner layer
comprising light pigmented to hyaline cells of textura
angularis. Hamathecium composed of branched, 1.5–2 lm
wide, filamentous, indistinct septate, anastomosed pseudoparaphyses, embedded in a hyaline, gelatinous matrix.
Asci (83.5–)102–144(–210) 9 (10.5–)13–15(–21.5) lm
(
x = 127.5 9 14.5 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate
with furcate to truncate pedicel, apically rounded, with
well-developed ocular chamber. Ascospores (31.5–)34–
45.5(–48) 9 (5.5–)6–11(–12.5) lm (
x = 39.1 9 8.6 lm,
n = 40), overlapping 2-seriate, hyaline, subfusoid to
Fig. 17 Sphaerellopsis isthmospora (KUN-HKAS 102225, holotype). a Appearance of ascomata on host substrate. b Section through ascoma.
c Section through peridium. d, e Pseudoparaphyses. f–h Asci. i–n Ascospores. Scale bars a = 500 lm, b = 100 lm, c = 50 lm, d–n = 20 lm
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Fungal Diversity
Fig. 18 Sphaerellopsis paraphysata (KUN-HKAS 101483). a Liriope
spicata. b–d Appearance of conidiomata associated with rust on host
substrate. e Section through conidioma wall. f Section through
conidioma. g, h Culture on PDA after 2 weeks (g = from above,
h = from below). i–r in vitro (OA). i Sporulation on OA after
4 weeks. j Section through conidioma. k Section through conidioma
wall. l, m Conidiogenous cells stained in congo red. n–r Conidia.
Scale bars j = 100 lm, c, d, f = 50 lm, k = 20 lm, e, l, m = 10 lm,
n = 5 lm, o–r = 2 lm
fusiform, with rounded or obtuse ends, 1(–3)-septate,
smooth-walled, guttulate when young, with an entire
mucilaginous sheath (9–20.5 lm wide at sides). Asexual
morph Undetermined.
Culture characteristics: Colonies on PDA reaching
2.5–3 mm diam. after 1 week at 25 ± 2 °C, circular,
convex or dome-shaped, rough with entire edge, mucoid,
smooth at the margin; from above, green earth at the
fruiting zone, grey at the productive zone and light grey at
ageing zone, dome, shining black mucoid colony; from
below, light grey at the fruiting zone, light green to
blackish at the productive zone, dark grey at the ageing
zone; cracking and not producing pigmentation in PDA
agar medium.
Material examined: CHINA, Yunnan Province, Baoshan
City, Longling County, on wood litter, October 2015, M.
Raza, BAP 119 (HMAS 255437, holotype), ex-type living
culture, LC12112.
GenBank
numbers:
ITS = MH822889,
LSU =
MH822890, SSU = MH822891, RPB2 = MH822892,
TEF1-a = MH822893.
Notes: Multi-loci phylogenetic analyses based on a
concatenated ITS, LSU, SSU, TEF1-a and RPB2 sequence
dataset show that Lophiotrema mucilaginosis forms a wellsupported lineage (100% ML and 1.00 BYPP; Fig. 20),
sister to L. hydei J.F. Zhang et al. and clusters with L.
neohysterioides M.E. Barr. Lophiotrema mucilaginosis and
L. hydei were collected on wood litter and herbaceous plant
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Fungal Diversity
Fig. 19 Lophiotrema mucilaginosis (HMAS 255437, holotype).
a Blackish ascomata on dead wood. b Vertical section of ascoma.
c Peridial structure. d Pseudoparaphyses. e, f Immature asci. g, h
Mature asci. i Apical ring stained with cotton blue. j Immature
ascospores. k–o Mature ascospores. p Ascospores with mucilaginous
sheath. q Germination of ascospore. r, s Culture characteristics on
PDA (r = from above, s = from below). Scale bars b = 100 lm,
c = 50 lm, d, p, q = 20 lm, e–i = 10 lm, o = 5, j–n = 2 lm
from Yunnan and Guizhou Provinces in China respectively. Lophiotrema mucilaginosis can be distinguished
from L. hydei in having larger asci (78–89(–
99) 9 6.9–8.8 lm, L. hydei) and larger ascospores
(23–28 9 3–4 lm, L. hydei) (Zhang et al. 2018). In addition, L. mucilaginosis has conical to mammiform ascomata,
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Fungal Diversity
Fig. 20 Phylogenetic tree
generated from maximum
likelihood analysis (RAxML)
based on a combined ITS, LSU,
SSU, TEF1-a and RPB2
sequence dataset of genera in
Lophiotremataceae. Maximum
likelihood bootstrap support
values greater than 70% and
Bayesian posterior probabilities
greater than 0.95 BYPP are
indicated on the branches. The
new isolate is in blue. The type
strains are in bold. The tree is
rooted with Cryptocoryneum
akitaense (KT 3019)
with flattened base and 1(–3)-septate ascospores. Whereas,
L. hydei has globose to subglobose ascomata and inconspicuously 0–1-septate ascospores. Lophiotrema mucilaginosis differs from L. neohysterioides by its larger asci
(70–96 9 7–10 lm, L. neohysterioides) and larger ascospores (17–25 9 3–5 lm, L. neohysterioides) and presence
of mucilaginous sheath surrounding the ascospores, a feature not observed in L. neohysterioides (Tanaka and Harada
2003a).
Occultibambusaceae D.Q. Dai & K.D. Hyde
Occultibambusaceae was introduced by Dai et al.
(2017a) and is typified by Occultibambusa D.Q. Dai &
K.D. Hyde with O. bambusae D.Q. Dai & K.D. Hyde being
the type species. The family was introduced to accommodate bambusicola-like taxa, mainly occurring on bamboo
(Dai et al. 2017a; Doilom et al. 2017). Dai et al. (2017a)
accepted four genera in Occultibambusaceae viz. Neooccultibambusa Doilom & K.D. Hyde., Occultibambusa,
Seriascoma Phookamsak et al. and Versicolorisporium Sat.
Hatak. et al. and this is in agreement of Wijayawardene
et al. (2018a). Subsequent authors introduced new taxa in
this family (Hyde et al. 2016, 2018b; Zhang et al. 2017b;
Tibpromma et al. 2018). We follow the latest treatment of
Occultibambusaceae in Dai et al. (2017a) and introduce the
new genus, Brunneofusispora S.K. Huang & K.D. Hyde to
accommodate Brunneofusispora sinensis and other isolates
of Massarina rubi. The updated sequence data were
retrieved from Zhang et al. (2017b).
123
Brunneofusispora S.K. Huang & K.D. Hyde, gen. nov.
Index Fungorum number: IF555599; Facesoffungi
number: FoF04862
Etymology: The generic epithet ‘‘Brunneofusispora’’
refers to the taxon having brown, fusiform ascospores.
Saprobic on dead wood. Sexual morph Ascomata
solitary to scattered, immersed, eventually erumpent, globose to subglobose, uni-loculate, glabrous, dark brown to
black, ostiolate, with long beak. Peridium composed of
brown to hyaline pseudoparenchymatous cells of textura
angularis. Hamathecium composed of numerous, filamentous, septate pseudoparaphyses, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical
to clavate, short pedicellate, rounded at the apex, with an
ocular chamber. Ascospores overlapping 2-seriate, hyaline
to brown, broadly fusiform, 1-septate, constricted at the
septum, smooth-walled, with guttules, surrounded by a
mucilaginous sheath. Asexual morph Undetermined.
Type species: Brunneofusispora sinensis S.K. Huang &
K.D. Hyde
Notes: Multigene phylogenetic analyses reveal that our
new taxon forms a distinct lineage, but clusters with three
isolates of Massarina rubi (Fuckel) Sacc. (current name:
Lophiotrema rubi (Fuckel) Y. Zhang ter et al.) in Occultibambusaceae (Fig. 21). We introduce a new genus
Brunneofusispora to accommodate B. sinensis, which was
collected from a woody plant in Yunnan, China. Brunneofusispora resembles Neooccultibambusa in occurring on
woody plants and forming globose to subglobose ascomata,
Fungal Diversity
Fig. 21 Maximum likelihood phylogenetic tree generated from
analysis of a combined LSU, SSU, ITS, RPB2 and TEF1-a sequence
dataset for 77 taxa of representative families in Pleosporales.
Melanomma pulvispyrius (CBS 124080) was selected as the outgroup
taxon. ML and MP support values greater than 60% and Bayesian
posterior probabilities greater than 0.90 BYPP are indicated above the
nodes as ML/PP/MP. The strain numbers are noted before the species
names. Isolates from this study are indicated in blue. Ex-type strains
are indicated in bold
but Brunneofusispora differs from Neooccultibambusa in
having a long prominent neck (Doilom et al. 2017).
Brunneofusispora resembles Lophiotrema in having
cylindrical, cylindric-clavate asci and fusiform ascospores,
but it differs from Lophiotrema in having globose to subglobose ascomata, with a long prominent neck. Lophiotrema species have minute papilla, with crest-like or slitlike openings (Zhang et al. 2009b; Hashimoto et al. 2017).
Lophiotrema rubi (: Masarina rubi) is an orphan
species in Lophiotremataceae, which was transferred from
Massarina based on phylogenetic analysis by Zhang et al.
(2009b). It is characterized by immersed ascomata with
broadly fusiform ascospores (Saccardo 1883b; Aptroot
1998). In this study, L. rubi strains CBS 691.95, MUT 4323
and MUT 4887 clustered with our strain (KUMCC
17-0030) and they form an independent clade within Occultibambusaceae (Fig. 21). However, the morphological
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Fig. 22 Brunneofusispora sinensis (KUN-HKAS 97451, holotype).
a Habitat. b, c Appearance of ascomata on dead wood. d Ascoma in
vertical section. e Peridium. f Asci with pseudoparaphyses stained in
congo red. g–i Developing stages of asci. j–m Ascospores (note: l, m
stained in congo red). Scale bars c = 500 lm, d = 200 lm,
f = 50 lm, g–i = 20 lm, e, j–m = 10 lm
characteristics of these strains have never been described
(Zhang et al. 2009a; Gnavi et al. 2017). We therefore,
tentatively include these strains of Massarina rubi until
they are clarified base on evidence from morphology and
phylogeny.
Etymology: The specific epithet ‘‘sinensis’’ refers to the
country, China, where the taxon was collected.
Holotype: KUN-HKAS 97451.
Saprobic on dead wood. Sexual morph Ascomata 325–
370 lm diam., perithecial, solitary to scattered, immersed,
eventually erumpent, globose to subglobose, uni-loculate,
glabrous, dark brown to black, ostiole, papillate with long
beak. Ostioles central, lined with periphyses. Peridium 20–
45 lm wide, equally thick-walled, composed of 5–8 strata,
of blackened pseudoparenchymatous cells, arranged in a
Brunneofusispora sinensis S.K. Huang & K.D. Hyde, sp.
nov.
Index Fungorum number: IF555600; Facesoffungi
number: FoF04863, Fig. 22
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Fungal Diversity
textura angularis. Hamathecium composed of numerous,
1.5–3.5 lm wide, filamentous, septate pseudoparaphyses,
embedded in a gelatinous matrix. Asci 53–110 9 9–18 lm
(
x = 75 9 14 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate to clavate, short pedicellate, rounded
at the apex, with an ocular chamber. Ascospores 18–
22 9 5–8.5 lm (
x = 20 9 7 lm, n = 50), overlapping
2-seriate, initially hyaline, becoming light brown to brown
at maturity, broadly fusiform, 1-septate, constricted at the
septum, smooth-walled, with guttules, surrounded by
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Ascospores germinated on PDA
within 2 weeks at 23 °C, colony on PDA reaching 1 cm
diam. after 4 weeks, irregular in shape, surface rough, with
edge umbonate and well-defined margin, velvety to floccose; from above brown; from below cream, not producing
pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Songming, Niulan river, on dead wood, 1 October 2016, S.K.
Huang (KUN-HKAS 97451, holotype), ex-type living
culture, KUMCC 17-0030.
GenBank
numbers:
ITS = MH393558,
LSU =
MH393557, SSU = MH393556, TEF1-a = MH395329.
Parabambusicolaceae Kaz. Tanaka & K. Hiray.
We follow the latest treatment and updated accounts of
Parabambusicolaceae in Tanaka et al. (2015), Li et al.
(2016), Phukhamsakda et al. (2016, 2018) and Wanasinghe
et al. (2017). Two novel genera are introduced based on
molecular phylogeny (Fig. 23) coupled with morphological
characterization viz. Lonicericola and Paratrimmatostroma. In addition, a novel species Parabambusicola
thysanolaenae is introduced.
Lonicericola Phookamsak, Jayasiri & K.D. Hyde, gen.
nov.
Index Fungorum number: IF556139; Facesoffungi
number: FoF04962
Etymology: The generic epithet ‘‘Lonicericola’’ refers to
the host genus Lonicera, from which the type species was
collected.
Saprobic o dead hanging branches of Lonicera maackii.
Sexual morph Ascomata black, scattered, solitary to gregarious, immersed under host epidermis, slightly raised,
globose to subglobose, uni-loculate, glabrous, ostiolate,
papillate. Peridium of equal thickness, composed of several
layers, of flattened to broad, brown to dark brown, pseudoparenchymatous cells, arranged in textura angularis to
textura prismatica. Hamathecium composed of numerous,
filamentous, septate, anastomosed pseudoparaphyses,
embedded in a hyaline gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, broadly cylindrical to cylindricclavate, subsessile to short pedicellate, with furcate to
obtuse pedicel, apically rounded, with ocular chamber.
Ascospores overlapping 2–3-seriate, hyaline, fusiform to
vermiform, with enlarged cell, septate, constricted at the
septa, smooth-walled, surrounded by distinct mucilaginous
sheath. Asexual morph Undetermined.
Type species: Lonicericola hyaloseptispora Phookamsak, Jayasiri & K.D. Hyde
Notes: Lonicericola is similar to the sexual genera
Aquastroma Kaz. Tanaka & K. Hiray., Multiseptospora
Phookamsak & K.D. Hyde, Neoaquastroma Wanas. et al.
and Parabambusicola Kaz. Tanaka & K. Hiray in
Parabambusicolaceae, in having hyaline, fusiform to vermiform, multi-septate ascospores, with an entire sheath
(Liu et al. 2015a; Tanaka et al. 2015). Lonicericola can be
distinguished from these related genera based on habitat,
ascomal shape, ascospore septation and multigene phylogenetic evidence. Lonicericola can be distinguished from
Aquastroma based on its terrestrial habitat, broadly cylindrical to cylindric-clavate asci, with a subsessile to short
pedicel and 9-septate ascospores. Aquastroma was found
on submerged woody plant from aquatic habitat and has
clavate asci, with a longer pedicel and 6–8-septate ascospores (Tanaka et al. 2015). Lonicericola resembles Multiseptospora in having globose to subglobose ascomata
immersed in the host tissue. However, Multiseptospora has
10–11-septate ascospores, the ascomata are covered by
brown to dark brown vegetative hyphal tufts, lack papilla,
and have pore-like openings (Liu et al. 2015a).
Neoaquastroma has 3–7-septate ascospores in N. guttulatum, 4–7-septate in N. bauhiniae and 5–8-septate in N.
krabiense (Wanasinghe et al. 2017; Phukhamsakda et al.
2018). The asexual morph of Neoaquastroma has been
reported as coelomycetous; whereas an asexual morph is
not yet known for Lonicericola. Parabambusicola differs
from Lonicericola in having hemispherical to conical
ascomata, with a flattened base, and 5–6-septate ascospores
as well as occurring on bamboo and stout grasses (Tanaka
et al. 2015).
Multigene phylogenetic analyses reveal that Lonicericola forms a distinct lineage with Aquastroma, Multiseptospora, Neoaquastroma and Parabambusicola and
clusters with the hyphomycetous genera Pseudomonodictys
Doilom et al. and Paratrimmatostroma Jayasiri et al. with
moderate support (77% ML and 0.99 BYPP). A comparison of ITS, LSU, SSU and TEF1-a sequence data indicates
that Lonicericola differs from Pseudomonodictys in
97/593 bp (16.4%, ITS), 14/888 bp (1.6%, LSU),
18/1033 bp (1.7%, SSU) and 36/959 bp (3.7%, TEF1-a).
However, we could not compare the morphological characteristics of Lonicericola with Pseudomonodictys and
Paratrimmatostroma as they are represented by different
morphs.
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Fungal Diversity
Fig. 23 Simplified phylogram showing the best RAxML tree obtained
from a combined multigene (SSU, ITS, LSU and TEF1-a) analyses.
Matrix of 42 taxa including related families of the family Parabambusicolaceae (Phukhamsakda et al. 2018). The matrix comprises 3580
characters with gaps. The best scoring RAxML tree with a final
likelihood value of -14853.890740 is presented. MLBS above 70%
and Bayesian posterior probabilities above 0.90 are given at each
branch. The tree is rooted with Morosphaeria velatispora KH218
(Morosphaeriaceae). Type species are in bold and new isolates are in
blue
Lonicericola hyaloseptispora Phookamsak, Jayasiri &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556140; Facesoffungi
number: FoF04963, Fig. 24
Etymology: The specific epithet ‘‘hyaloseptispora’’
refers to the fungus having hyaline, multi-septate
ascospores.
Holotype: KUN-HKAS 102223
Saprobic dead hanging branches of Lonicera maackii.
Sexual morph Ascomata 170–240 lm high, 165–250 lm
diam., black, scattered, solitary to gregarious, immersed
under host epidermis, slightly raised, globose to subglobose, uni-loculate, glabrous, ostiolate, papillate. Ostioles
centrally located, oblong, with minute papilla, with porelike opening, filled with hyaline periphyses. Peridium
8–25 lm wide, of equal thickness, composed of several
layers, of flattened to broad, brown to dark brown,
pseudoparenchymatous cells, arranged in textura angularis to textura prismatica. Hamathecium composed of
numerous, 2–3.5 lm wide, filamentous, septate, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (90–)100–120(–145) 9 (24–)25–30(–33)
lm (
x = 115.1 9 27 lm, n = 30), 8-spored, bitunicate,
fissitunicate, broadly cylindrical to cylindric-clavate, subsessile to short pedicellate, with furcate to obtuse pedicel,
apically rounded, with ocular chamber clearly visible when
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Fungal Diversity
Fig. 24 Lonicericola hyaloseptispora (KUN-HKAS 102223, holotype). a, b Appearance of ascomata on host substrate. c Section through ascoma. d Ostiole with papilla immersed in the host.
e Section through peridium. f Asci immersed in hyaline, cellular
pseudoparaphyses. g, h Asci. i–k Ascospores. l Ascospore stained
with Indian ink. m Germinating ascospore. Scale bars a, b = 500 lm,
c = 100 lm, d, e = 50 lm, f–m = 20 lm
young. Ascospores overlapping 2–3-seriate, hyaline,
becoming brown when release from the asci, fusiform to
vermiform, enlarged at the 4th cell from the apex
(4–10 9 9–12 lm, l/w), (8–)9-septate, constricted at the
septa, smooth-walled, with small to large guttules, surrounded by entire mucilaginous sheath (3.5–13 lm wide).
Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
50–55 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony dense, circular, flattened, slightly
raised, surface smooth, with entire edge, floccose to fluffy;
from above dark grey to brown; from below, black; not
producing pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Kunming, Kunming Institute of Botany, on dead hanging
branches of Lonicera maackii, 20 April 2017, R.
Phookamsak, KIB034 (KUN-HKAS 102223, holotype),
ex-type living culture KUMCC 18-0149 (KIB034A),
KUMCC 18-0150 (KIB034B).
GenBank
numbers:
ITS = MK098191,
LSU =
MK098197, SSU = MK098203, (KUMCC 18-0149);
ITS = MK098194,
LSU = MK098200,
SSU =
MK098206, TEF1-a = MK098210 (KUMCC 18-0150).
Parabambusicola Kaz. Tanaka & K. Hiray
We follow the latest treatment and updated accounts of
Parabambusicola in Tanaka et al. (2015). Previously, only
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Fungal Diversity
Fig. 25 Parabambusicola thysanolaenae (KUN-HKAS 102222,
holotype). a Appearance of ascomata on host substrate. b Section through ascoma. c Section through peridium. d Ostiole. e Asci
123
immersed in hyaline, cellular pseudoparaphyses, stained in Indian ink.
f Ascus. g–i Ascospores. j, k Ascospores stained in Indian ink. Scale
bars a = 500 lm, b = 100 lm, c, d = 50 lm, e–k = 20 lm
Fungal Diversity
P. bambusina was accommodated in this genus (Index
Fungorum 2019). We introduce the second species, P.
thysanolaenae, collected from Thysanolaena maxima in
Yunnan, China.
Parabambusicola thysanolaenae Goonas., Jayasiri,
Phookamsak & K.D. Hyde, sp. nov.
Index Fungorum number: IF555596; Facesoffungi
number: FoF04964, Fig. 25
Etymology: Named after the host from which the fungus
was isolated.
Holotype: KUN-HKAS 102222
Saprobic on dead stems of Thysanolaena maxima,
appearing as raised, dome-shaped areas on host surface,
covered by brown, vegetative hyphae. Sexual morph Ascomata 130–170 lm high, 430–600 lm wide, mostly
clustered together, sometimes solitary, immersed under
host epidermis, raised, becoming semi-immersed, globose
in surface view, hemispherical with a flattened base in
cross section, uni-loculate, glabrous, ostiole central, with
pore-like opening. Peridium 25–60 lm wide, lateral
walls composed of numerous layers of inner, hyaline,
flattened cells to outer, pale brown to brown, textura
angularis cells and pale brown to brown, globular or
polygonal cells showing no conspicuous layers at the base,
intermixed with host tissue. Hamathecium composed of
numerous, 1.5–3 lm wide, filamentous, septate pseudoparaphyses, anastomosing above the asci, embedded in a
hyaline gelatinous matrix. Asci (50–)80–120 9 (10–)
25–33 lm (
x = 107 9 28 lm, n = 30), 8-spored, bitunicate, fissitunicate, broadly cylindrical to cylindric-clavate,
subsessile, rounded at the apex, with inconspicuous ocular chamber, clearly visible when young. Ascospores
45–55 9 7.5–11 lm (
x = 46.5 9 9 lm, n = 35), overlapping 2–3-seriate, hyaline, fusiform to vermiform, narrower
towards the lower cell, enlarged at the 4th cell from apex,
slightly curved, 5–(6–7)-septate, primary septum mostly
median, constricted at the septa, smooth-walled, with an
entire sheath, large guttules present when immature.
Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
30–32 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony dense, circular, flattened, surface
smooth, with entire edge, velvety to floccose; from above
greenish grey to dark green, paler at the edge; from below,
black; produced dark brown pigmentation around colony
on agar medium. Colonies on MEA reaching 28–30 diam.
after 3 weeks at room temperature (20–30 °C). Colony
dense, circular, flattened, surface smooth, with entire edge,
floccose to cottony; from above cream to pale yellowish;
from below, yellowish brown, paler at the edge; not producing pigmentation on agar medium.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Mengla County, Xishuangbanna Tropical
Botanical Garden (XTBG), on dead stems of Thysanolaena
maxima (Roxb.) Kuntze (Poaceae), 22 April 2017, R.
Phookamsak, IS003 (KUN-HKAS 102222, holotype), extype living culture, KUMCC 18-0147 (IS003A), KUMCC
18-0148 (IS003B).
GenBank numbers: ITS = MK098190, LSU = MK098199,
SSU = MK098205, TEF1-a = MK098209 (KUMCC 18-0147);
ITS = MK098193, LSU = MK098198, SSU = MK098202,
TEF1-a = MK098211 (KUMCC 18-0148).
Notes: Parabambusicola thysanolaenae shares similar
peridial and ascal characters with P. bambusina (Teng)
Kaz. Tanaka & K. Hiray. but can be distinguished by
having larger ascomata (300–500 9 150–300 lm in P.
bambusina), absence of a beak-like structure and wider
peridium (10–20 lm in P. bambusina). Ascospores of P.
thysanolaenae are 5-septate and shorter than the 3–5-septate ascospores of P. bambusina (54.7 9 8.4 lm, Tanaka
and Harada 2003b). Phylogenetically P. thysanolaenae
clusters with P. bambusina forming a well-separated lineage (100% ML and 1.00 BYPP; Fig. 23).
Paratrimmatostroma Jayasiri, Phookamsak, D.J. Bhat &
K.D. Hyde, gen. nov.
Index Fungorum number: IF556153; Facesoffungi
number: FoF04960
Etymology: With reference to similar morphology of
genus ‘‘Trimmatostroma’’
Saprobic on dead fronds of a fern. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Sporodochia effuse or confluent, visible as black powdery,
superficial mass on host substrate, flattened, light brown,
with a membranous base, composed of pseudoparenchymatous cells of textura angularis. Mycelium immersed,
composed of septate, pale brown, branched hyphae. Conidiophores macronematous, or semi-macronematous,
mononematous, prostrate, or erect, usually short, oblong to
cylindrical, straight or flexuous, arising as lateral branches
from creeping hyphae, septate, branched or unbranched,
slightly constricted at the septa so as to give a monilioid
appearance, pale brown, smooth-walled. Conidiogenous
cells holoblastic, mono- to polyblastic, integrated, terminal,
brown, smooth-walled. Conidia solitary, acropleurogenous,
dark brown, paler at the apical cell, branched, straight or
flexuous, variable in shape, helicoid, cylindrical, sigmoid,
or reniform, solitary, tapering near apex and base, rounded
at tip, septate, constricted at the septa, smooth and thickwalled.
Type species: Paratrimmatostroma kunmingensis Jayasiri, Phookamsak, D.J. Bhat & K.D. Hyde
123
Fungal Diversity
Fig. 26 Paratrimmatostroma kunmingensis (KUN-HKAS102224, holotype). a Appearance of sporodochia on host substrate. b, c Sporodochia.
d–g Conidiophores with attached conidia. h–q Conidia. Scale bars a = 200 lm, b = 50 lm, d–q = 10 lm
Notes: Paratrimmatostroma is similar to Trimmatostroma Corda in forming effuse to confluent sporodochia,
semi-macronematous,
mononematous
conidiophores, integrated, terminal conidiogenous cells,
and branched, straight or flexuous, septate, pigmented
conidia, which are variable in shape (Ellis 1971; Crous
et al. 2007a). However, they are phylogenetically distinct
in that Paratrimmatostroma belongs to Parabambusicolaceae (Pleosporales, Dothideomycetes), whereas Trimmatostroma was recently treated in Mollisiaceae Rehm
(Helotiales, Leotiomycetes) (Crous et al. 2007a;
Wijayawardene et al. 2018a). Multigene phylogenetic
analyses show that Paratrimmatostroma forms a well-resolved clade (74% ML and 1.00 BYPP; Fig. 23), and
clusters with Pseudomonodictys and Lonicericola.
Paratrimmatostroma is distinct from Pseudomonodictys in
forming sporodochia on the host substrate and having
branched, straight or flexuous conidia, with variable conidial shape. Pseudomonodictys has muriform, top-shaped to
ellipsoidal conidia and does not form sporodochia on host
123
substrate (Ariyawansa et al. 2015a). Paratrimmatostroma
was found on a fern in Yunnan, China (nonflowering
vascular plants in the low-latitude monsoon climate),
whereas, Pseudomonodictys was collected from teak in
Thailand (flowering plant in tropical climate). A comparison of ITS, LSU, SSU and TEF1-a sequence dataset
indicates that Paratrimmatostroma differs from Pseudomonodictys in 64/540 bp (11.8%, ITS), 23/857 bp
(2.7%, LSU), 22/1060 bp (2.1%, SSU) and 41/928 bp
(4.4%, TEF1-a). Based on phylogenetic analysis and
morphological distinctiveness, we introduce Paratrimmatostroma as a new genus to accommodate a single
species, P. kunmingensis.
Paratrimmatostroma kunmingensis Jayasiri, Phookamsak,
D.J. Bhat & K.D. Hyde, sp. nov.
Index Fungorum number: IF556152; Facesoffungi
number: FoF04961, Fig. 26
Fungal Diversity
Etymology: The specific epithet ‘‘kunmingensis’’ refers
to Kunming City, Yunnan Province, China, where the type
was collected.
Holotype: KUN-HKAS 102224
Saprobic on dead fronds of a fern. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Sporodochia effuse or confluent, visible as black powdery,
superficial mass on host substrate, flattened, light brown,
with a membranous base, composed of pseudoparenchymatous cells of textura angularis. Mycelium immersed,
composed of septate, pale brown, branched hyphae. Conidiophores (6–)15–30(–50) 9 2–4 lm (
x = 19.2 9 3.2 lm,
n = 20),
macronematous
or
semi-macronematous,
mononematous, prostrate, or erect, usually short, oblong to
cylindrical, straight or flexuous, arising as lateral branches
from creeping hyphae, septate, branched or unbranched,
slightly constricted at the septa so as to give a monilioid
appearance, pale brown, smooth-walled. Conidiogenous
cells 3–10(–15) 9 2–4.5 lm (
x = 7.1 9 3.1 lm, n = 30),
holoblastic, mono- to polyblastic, integrated, terminal,
brown, smooth-walled. Conidia solitary, acropleurogenous,
dark brown, paler at the apical cell, branched, straight or
flexuous, variable in shape, helicoid [(8–)10–20(–
27) 9 (6–)10–20 lm (
x = 16.8 9 13.7 lm, n = 40)],
cylindrical to sigmoid, or reniform [(8.5–)15–50 9 6–8(–
10) lm (
x = 29.7 9 7.7 lm, n = 50)], solitary, tapering
near apex and base, rounded at tip, multi-septate, 6–13septate at maturity, constricted at the septa, smooth and
thick-walled.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead fronds of
a fern, 1 April 2017, R. Phookamsak, KIB025 (KUNHKAS 102224, holotype).
GenBank
numbers:
ITS = MK098192,
LSU =
MK098196, SSU = MK098204, TEF1-a = MK098208
(KUN-HKAS 102224A); ITS = MK098195, LSU =
MK098201, SSU = MK098207 (KUN-HKAS 102224B).
Periconiaceae (Sacc.) Nann.
Periconiaceae has long been unused and placed as
members of Massarinaceae Munk until Tanaka et al.
(2015) revised Massarineae and placed it as a distinct
family based on phylogenetic analysis (Tanaka et al. 2015;
Hyde et al. 2017, 2018b). We follow the latest treatment
and updated accounts of Periconiaceae in Tanaka et al.
(2015), Hyde et al. (2017, 2018b), Liu et al. (2017b) and
Thambugala et al. (2017). We report Periconia cortaderiae
Thambug. & K.D. Hyde from Caragana arborescens Lam.
(Fabaceae) in Yunnan, China for the first time.
Periconia Tode
Periconia was introduced by Tode (1791) to accommodate hyphomycetous species having macronematous
conidiophores and 1-celled, pigmented, verruculose to
echinulate conidia and is typified by P. lichenoides Tode
(Tanaka et al. 2015; Thambugala et al. 2017). The genus
was re-circumscribed by Tanaka et al. (2015) and this was
followed by subsequent authors (Hyde et al. 2017, 2018b;
Liu et al. 2017b; Thambugala et al. 2017). We follow the
latest treatment of Periconia in Tanaka et al. (2015). The
updated phylogenetic analyses were retrieved from
Thambugala et al. (2017) and Hyde et al. (2018b) (Fig. 27).
Periconia cortaderiae Thambug. & K.D. Hyde, in Thambugala et al., Mycosphere 8(4): 734 (2017), Fig. 28
Holotype: THAILAND, Chiang Rai, Mae Fah Luang
University, on dead stems and leaves of Cortaderia sp.
(Poaceae), 21 December 2014, K.M. Thambugala, KM
035 (MFLU16–2579), ex-type living culture MFLUCC
15–0457, ICMP 21414
Saprobic on dead, hanging branches of Caragana
arborescens. Sexual morph Undetermined. Asexual
morph Colonies on the substratum superficial, numerous,
effuse, black, floccose. Mycelium immersed on the substrate, composed of septate, branched, smooth, dark
hyphae. Conidiophores 100–225 9 8.5–11 lm, macronematous, mononematous, septate, branched, erect, mostly
slightly flexuous to curved, dark brown, forming sphaerical
heads at apex, arising from a stromatic base. Conidiogenous cells 4.5–5.2 9 4.1–5.6 lm (
x = 4.9 9 5.1 lm,
n = 20), mono- to polyblastic, discrete, terminal, globose,
light brown. Conidia (5.5–)6–9(–12) 9 (4.5–)5–8 lm
(
x = 7.8 9 6.4 lm, n = 50), solitary or catenate, in acropetal chains, subglobose to globose, occasionally ellipsoidal to cylindrical, light brown to moderately brown,
verruculose.
Culture characteristics: Colonies on PDA reaching
68–74 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony medium sparse, circular, flattened,
surface smooth, edge entire, velvety to woolly; from above,
sectoring, with a part of cream to pale yellowish and yellowish-green to dark grey, with small white tufts in another
part; from below, pale yellowish at the margin, sectoring at
the centre, with a part of yellowish to orange-yellow and
dark brown in another part; not producing pigmentation on
agar medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead hanging
branches of Caragana arborescens (Fabaceae), 2 April
2017, R. Phookamsak, KIB017 (KUN-HKAS 102240),
living culture, KUMCC 18-0174 = MFLUCC18-0667
(KIB017A),
KUMCC
18-0175 = MFLUCC18-0668
(KIB017B).
Known hosts and distribution: On dead stems and leaves
of Cortaderia sp. (Poaceae, Thailand) and Caragana
arborescens (Fabaceae, Yunnan, China; in this study).
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Fungal Diversity
Fig. 27 Phylogram generated from the best scoring of the RAxML
tree based on combined ITS, LSU and TEF1-a sequence dataset of
taxa in Periconiaceae and other related families (Massarinaceae,
Didymellaceae and Lentitheciaceae). Taxa in Morosphaeriaceae,
Morosphaeria ramunculicola (KH 220) and M. velatispora (KH 221)
were selected as the outgroup taxa. Bootstrap support values for
maximum likelihood (green) equal to or greater than 70% and the
Bayesian posterior probabilities (blue) equal or higher than 0.95 are
indicated above the nodes. Ex-type and ex-epitype strains are in bold.
Newly generated sequences are indicated in blue
GenBank
numbers:
ITS = MH892348,
LSU =
MH892401 (MFLUCC 18-0667); ITS = MH892349,
LSU = MH892402, TEF1-a = MH908964 (MFLUCC180668).
Notes: Phylogenetic analyses of a concatenated LSU,
ITS and TEF1-a sequence datset show that our strains
(MFLUCC 18-0667 and MFLUCC 18-0668) form a wellresolved clade, clustering with Periconia cortaderiae
(100% ML and 1.00 BYPP; Fig. 27). A comparison of ITS
and TEF1-a nucleotide base pair indicates that our strains
are identical to the type strain of P. cortaderiae, which is
not significantly different (2/520 bp in ITS and 4/830 bp in
TEF1-a). We therefore, identify our isolates as P. cortaderiae and report its occurrence on Caragana arborescens in Yunnan, China for the first time. Our isolate has
shorter conidiophores (400–800 lm high, holotype) and
slightly larger conidia (4–6.6 9 4.1–7.1 lm, holotype)
(Thambugala et al. 2017). Periconia species are morphologically variable in different hosts and geographic distributions, but the molecular data of informative genes can
123
Fungal Diversity
Fig. 28 Periconia cortaderiae (KUN-HKAS102240). a Appearance of fungal colonies on host substrate. b, c Conidiophores. d–f Conidiophores
and conidiogenous cells. g–j Conidia. Scale bars b, c = 50 lm, d–f = 20 lm, g, j = 10 lm, h, i = 5 lm
clarify their conspecificity. Therefore, more informative
genes such as RPB2 and TUB2 are needed in further
studies of this genus.
Phaeosphaeriaceae M.E. Barr
Phaeosphaeriaceae was introduced by Barr (1979) to
accommodate fungal taxa which mainly occurr on monocotyledons, but are also found on some herbaceous plants
(Phookamsak et al. 2014b, 2017). Many additional genera
have been introduced in this family since Phookamsak
et al. (2014b) re-circumscribed the genera in
Phaeosphaeriaceae. Wijayawardene et al. (2018a) listed
52 genera in Phaeosphaeriaceae. Wanasinghe et al. (2018),
Bakhshi et al. (2019) and Maharachchikumbura et al.
(2019) introduced other seven genera in this family, and 59
genera are now accepted in Phaeosphaeriaceae based on
molecular phylogeny coupled with morphological characteristics (Wanasinghe et al. 2018; Wijayawardene et al.
2018a). We follow the latest treatment and updated
accounts of Phaeosphaeriaceae in Phookamsak et al.
Fig. 29 Phylogram generated from maximum likelihood analysis c
based on a concatenated LSU, SSU, TEF1-a and ITS sequence dataset
of Phaeosphaeriaceae. Updated sequence data were obtained from
GenBank (http://www.ncbi.nlm.nih.gov/) and retrieved from Hyde
et al. (2018b) and Wanasinghe et al. (2018). One hundred and fifty-two
strains are included in the analysis. Staurosphaeria rhamnicola
(MFLUCC 17-0813 and MFLUCC 17-0814) were selected as the
outgroup taxon. Bootstrap support values for maximum likelihood (left)
equal to or greater than 60% and Bayesian posterior probabilities (right)
equal or higher than 0.95 are indicated above the nodes. Newly
generated sequences are in blue. Type strains are indicated in bold
123
Fungal Diversity
123
Fungal Diversity
Fig. 29 continued
123
Fungal Diversity
Fig. 29 continued
(2014b), Hyde et al. (2018b) and Wanasinghe et al. (2018).
A new genus Brunneomurispora is introduced to accommodate a single species, B. lonicerae. Five new species are
also introduced viz. Galiicola baoshanensis, Neosetophoma lonicerae, Ophiobolus malleolus, Phaeosphaeria
cycadis and Wojnowiciella kunmingensis. Furthermore,
Wojnowicia rosicola W.J Li et al. is transferred to Wojnowiciella. Amarenomyces dactylidis Mapook et al. is
reported from a fern in China for the first time. An updated
phylogenetic analysis (Fig. 29) was performed following
123
previous phylogenies derived from Hyde et al. (2018b) and
Wanasinghe et al. (2018).
Amarenomyces O.E. Erikss.
Amarenomyces was introduced by Eriksson (1981) and
is typified by A. ammophilae (Lasch) O.E. Erikss., occurring on marine grass (Ammophila arenaria L., Poaceae).
The genus is characterized by immersed to erumpent,
globose to subglobose ascomata, thin-walled peridium,
multi-layered endotunica, broadly cylindrical asci, with
subsessile, knob-like pedicel and large, pigmented, septate,
thick-walled and sheathed ascospores (Phookamsak et al.
Fungal Diversity
2014b). Eriksson (1981) placed the genus in
Botryosphaeriaceae Theiss. & Syd.; however, Zhang et al.
(2009a) treated the genus as a synonym of Phaeosphaeria
I. Miyake in Phaeosphaeriaceae based on molecular phylogeny. Phookamsak et al. (2014b) re-circumscribed the
genera in Phaeosphaeriaceae based on multigene phylogenetic analyses coupled with morphological studies and
thus, Amarenomyces was re-instated. Hyde et al. (2017)
introduced a second species, A. dactylidis Mapook et al.,
collected from dead aerial stems of Dactylis glomerata L.
in Italy. Only two species are presently accommodated in
this genus.
Amarenomyces dactylidis Mapook, Camporesi & K.D.
Hyde, in Hyde et al., Fungal Divers 87: 78 (2017), Fig. 30
Holotype: ITALY, Forlı̀-Cesena Province, Camposonaldo-Santa Sofia, on dead aerial stems of Dactylis glomerata (Poaceae), 10 January 2014, E. Camporesi, MFLU
17-0498, ex-type living culture MFLUCC 14-0207.
Saprobic on dead fronds of a fern. Sexual morph Ascomata 170–230 lm high, 160–260 lm diam., scattered,
solitary, semi-immersed to superficial, visible as raised,
black dots on the host surface, globose to subglobose, uniloculate, glabrous, ostiole central, with minute, papilla (20–
45 9 25–55 lm, l/w), lacking periphyses. Peridium 13–
23 lm wide, thin-walled of equal thickness, composed of
5–6 cell layers, of flattened to broad, pseudoparenchymatous cells; outer layer comprising brown to dark brown
cells of textura angularis; inner layer comprising flattened,
hyaline to pale brown cells, of textura angularis to textura
prismatica. Hamathecium composed of numerous, 1.8–
4.5 lm wide, filamentous, septate, pseudoparaphyses,
anastomosing above the asci, embedded in a hyaline
gelatinous matrix. Asci (70–)75–95(–113) 9 (9.5–)10–
13(–14) lm (
x = 88.9 9 12 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short
pedicellate, with furcate pedicel, apically rounded, with
well-developed ocular chamber. Ascospores (19–)(23–)25–
28(–32) 9 4.5–6 lm (
x = 26.1 9 5.5 lm, n = 50), overlapping 1–2-seriate, yellowish brown to brown, fusiform to
subcylindrical, slightly narrower towards the end cells,
asymmetrical, 7–8-septate, slightly constricted at the septa,
enlarged at the 5th or 6th cell from above, flattened at the
2nd to 4th, or 5th cells, smooth-walled, surrounded by a
distinct
mucilaginous
sheath.
Asexual
morph
Undetermined.
Culture characteristics: Colonies on PDA reaching
35–38 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony dense, irregular in shape, flattened to
slightly raised, surface smooth, with edge undulate, velvety
to floccose; from above, initially white, with pale grey at
the centre, becoming greenish grey after 4 weeks; from
below, white to pale yellowish at the margin, dark brown at
the centre, becoming black after 4 weeks; not producing
pigmentation on agar medium.
Material examined: CHINA, Yunnan Province,
Baoshan, Shuizai, Dawazi mountain, on dead fronds of a
fern, 22 October 2015, I.D. Goonasekara, BS008 (KUNHKAS 102230), living culture, KUMCC 18-0154.
Known hosts and distribution: On dead aerial stems of
Dactylis glomerata (Poaceae, Italy) and on dead fronds of
a fern (Yunnan, China) (Hyde et al. 2017; this study).
GenBank
numbers:
ITS = MK356371,
LSU =
MK356345, SSU = MK356359.
Notes: Multigene phylogenetic analyses show that the
new strain KUMCC 18-0154 is sister to Amarenomyces
dactylidis (MFLUCC 14-0207) with high support (98%
ML and 1.00 BYPP). A comparison of ITS nucleotide base
pairs shows that KUMCC 18-0154 is identical (1/541 bp)
to A. dactylidis and thus, we identify our new isolate as A.
dactylidis. Our isolate (KUN-HKAS 102230) shares a size
range of the ascomata, asci and ascospores as well as the
ascospore septation with the type (MFLU 17-0498).
Although, our isolate is slightly larger in ascomata, asci
and ascospores, but ITS sequence data showed that they are
conspecific (Hyde et al. 2017). Therefore, A. dactylidis is
reported from a fern in Yunnan, China for the first time.
Brunneomurispora Phookamsak, Wanas. & K.D. Hyde,
gen. nov.
Index Fungorum number: IF556165; Facesoffungi
number: FoF05699
Etymology: The generic epithet ‘‘Brunneomurispora’’
refers to the fungus having brown, muriform ascospores.
Saprobic on Lonicera maackii. Sexual morph Ascomata scattered, solitary or in groups, semi-immersed to
erumpent, globose to subglobose, or irregular in shape, uniloculate, glabrous, ostiolate, papillate. Peridium slightly
thick, composed of several layers, of small, flattened to
broad, dark brown, pseudoparenchymatous cells of textura
angularis to textura prismatica. Hamathecium composed
of numerous, broad, filamentous, septate, anastomosed
pseudoparaphyses, embedded in a hyaline gelatinous
matrix. Asci 8-spored, bitunicate, fissitunicate, broadly
cylindrical to cylindric-clavate, or clavate, short pedicellate, apically rounded, with inconspicuous ocular chamber.
Ascospores overlapping 1–2-seriate, dark brown, muriform, fusiform to ellipsoidal, asymmetrical, slightly larger
in the upper part, constricted at the central septum, smoothwalled, lacking mucilaginous sheath. Asexual morph
Undetermined.
Type species: Brunneomurispora lonicerae Phookamsak, Konta, Wanas. & K.D. Hyde
Notes: Phylogenetic analyses of a concatenated LSU,
SSU, TEF1-a and ITS sequence dataset (Fig. 29) show that
our new strains (KUMCC 18-0157 and KUMCC 18-0158)
123
Fungal Diversity
form a well-separated lineage basal to Neosetophoma
Gruyter et al. with high support (94% ML and 0.99 BYPP).
Our new isolate is distinct from Neosetophoma in having
dark brown muriform ascospores. While, the sexual morph
of Neosetophoma has phragmosporous, brown, fusiform
ascospores (Tibpromma et al. 2017; Hyde et al. 2018b).
Hence, we introduce a new genus Brunneomurispora
herein to accommodate B. lonicerae which was isolated
from Lonicera maackii in Yunnan, China.
Brunneomurispora resembles Embarria Wanas. et al.
and Hydeomyces Maharachch. et al. in having immersed to
erumpent, globose or subglobose ascomata, with a minute
papilla, clavate asci, with dark brown, muriform, asymmetrical ascospores and ascospores that are constricted at
the central septum (Wanasinghe et al. 2018;
Maharachchikumbura et al. 2019). However, Brunneomurispora can be distinguished from Embarria in its
peridium structure comprising several layers of brown,
small, flattened to broad pseudoparenchymatous cells and
its ascospores being 4–6 transverse septa, with 1–2 longitudinal septa, sectored, and lacking a mucilaginous sheath.
Embarria has a thin-walled peridium, comprising large,
2–3 cell layers of pseudoparenchymatous cells and its
ascospores are 4–6 transverse septa, with a single longitudinal septum, surrounded by a thick mucilaginous sheath
(Wanasinghe et al. 2018). Hydeomyces differs from
Brunneomurispora in having thicker peridium (35–60 lm
thick), smaller, cylindrical asci (70–85 9 9–17 lm) and
smaller, 1-seriate, muriform ascospores, with 2–4 transverse septa and 1 longitudinal septum (10–15 9 5–6.5 lm)
(Maharachchikumbura et al. 2019). Hawksworthiana lonicerae Wanas. et al. was also isolated from Lonicera in
Italy. This species differs from Brunneomurispora lonicerae in having cylindrical to cylindric-clavate asci, with
yellowish brown, ellipsoidal, muriform, 3 transverse septa,
with 1 longitudinal septum ascospores (Wanasinghe et al.
2018).
Many genera in Phaeosphaeriaceae are characterized by
dictyosporous ascospores viz. Allophaeosphaeria Ariyaw.
et al., Dactylidina Wanas. et al., Dematiopleospora Wanas.
et al., Embarria, Galiicola Tibpromma et al., Hawksworthiana Wanas. et al., Hydeomyces Maharachch. et al.,
Italica Wanas. et al., Muriphaeosphaeria Phukhams. et al.,
Populocrescentia Wanas. et al. and Yunnanensis Karun.
et al. (Wanasinghe et al. 2014b, 2018; Ariyawansa et al.
2015a; Liu et al. 2015a; Phukhamsakda et al. 2015; Karunarathna et al. 2017; Maharachchikumbura et al. 2019).
These genera are represented by a single or a few species
and have very little morphological differences in their
sexual morphs. However, they always form distinct
clades, separate from each other, as well as the asexual
morphs of some different genera. Furthermore, Poaceicola
and Populocrescentia are heterogeneous, forming both
123
phragmosporous and dictyosporous ascospores (Wanasinghe et al. 2018). More sampling of taxa in these genera
are needed for a better understanding.
Brunneomurispora lonicerae Phookamsak, Konta, Wanas.
& K.D. Hyde, sp. nov.
Index Fungorum number: IF556166; Facesoffungi
number: FoF05700, Fig. 31
Etymology: The specific epithet ‘‘lonicerae’’ refers to
the host genus Lonicera, from which the fungus was
collected.
Holotype: KUN-HKAS 102232
Saprobic on dead hanging branches of Lonicera
maackii. Sexual morph Ascomata 170–280 lm high, 230–
330 lm diam., scattered, solitary, or in groups, semi-immersed to erumpent, visible as raised, black dot on the host
surface, globose to subglobose, occasionally irregular in
shape, uni-loculate, glabrous, ostiole central, with minute,
mammiform papilla, lacking periphyses. Peridium 15–
40 lm wide, of unequal thickness, composed of several
layers, of small, flattened to broad, pseudoparenchymatous
cells; outer layer comprising brown to dark brown cells of
textura angularis; inner layer comprising flattened, hyaline
cells of textura angularis to textura prismatica. Hamathecium composed of numerous, 2–5.5 lm wide, filamentous,
septate, anastomosed pseudoparaphyses, embedded in a
hyaline gelatinous matrix. Asci (67–)80–110(–132) 9 (13–
)16–20(–24) lm (
x = 98 9 18.8 lm, n = 25), 8-spored,
bitunicate, fissitunicate, broadly cylindrical to cylindricclavate, or clavate, short pedicellate, with truncate pedicel,
apically rounded, with inconspicuous ocular chamber,
clearly seen when immature. Ascospores (12–)14–18(–
20)(–23) 9 (4–)5–8(–13) lm (
x = 17 9 7.7 lm, n = 50),
overlapping 1–2-seriate, dark brown, muriform, fusiform to
ellipsoidal, with acute or rounded ends, or acute at the
upper cells, asymmetrical, slightly larger in the upper part,
straight, sometimes bent, mostly 4–6 transverse septa, with
1–2 longitudinal septa in each cell, becoming many sectors,
constricted at the central septum, smooth-walled, lacking a
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
28–30 mm diam. after 1 week at room temperature
(20–30 °C). Colony medium dense, circular, flattened,
surface smooth, with edge entire, velvety to woolly; from
above, white to cream at the margin towards the centre,
with sectored, greenish grey to dull green or light green at
the centre; from below, cream to pale yellowish at the
margin, yellowish at the middle, brown-green at the centre;
not producing pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead hanging
branches of Lonicera maackii, 20 April 2017, R.
Phookamsak, KIB030 (KUN-HKAS 102232, holotype),
Fungal Diversity
Fig. 30 Amarenomyces dactylidis (KUN-HKAS 102230). a Appearance of ascomata on host. b Section through ascoma. c Section through
peridium. d Cellular pseudoparaphyses. e–h Asci. i–n Ascospores.
o Ascospore stained with Indian ink. p Ascospore germination. Scale
bars a = 200 lm, b = 50 lm, c–h = 20 lm, i–p = 10 lm
ex-type living culture KUMCC 18-0157 (KIB030A),
KUMCC 18-0158 (KIB030B).
GenBank
numbers:
ITS = MK356372,
LSU =
MK356346, SSU = MK356360, TEF1-a = MK359064,
RPB2 = MK359079
(KUMCC
18-0157);
ITS =
MK356373,
LSU = MK356347,
SSU = MK356361,
TEF1-a = MK359065 (KUMCC 18-0157).
Tibpromma et al. which was found as a saprobe on Galium
in Italy. The genus is characterized by semi-immersed to
erumpent, globose to subglobose ascomata, fissitunicate,
cylindric-clavate asci, orange-brown, elongate fusiform
ascospores with 4–5 transverse septa, some with 1–2 longitudinal septa; its asexual morph has not been found
(Ariyawansa et al. 2015a). We introduce a second species,
G. baoshanensis which is represented by its asexual morph.
Galiicola Tibpromma et al.
Galiicola was introduced by Ariyawansa et al. (2015a)
to accommodate a single species G. pseudophaeosphaeria
Galiicola baoshanensis Phookamsak, Wanas. & K.D.
Hyde, sp. nov.
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Fungal Diversity
Index Fungorum number: IF556167; Facesoffungi
number: FoF05701, Fig. 32
Etymology: The specific epithet ‘‘baoshanensis’’ refers
to Baoshan prefecture-level city of Yunnan Province,
China, where the holotype was collected.
Holotype: KUN-HKAS 102234
Saprobic on dead branches of herbaceous plant. Sexual
morph Undetermined. Asexual morph Coelomycetous,
amarenographium-like. Conidiomata 90–125 lm high, 90–
120 lm diam., pycnidial, scattered, solitary, immersed to
semi-immersed, uni-loculate, globose to subglobose,
glabrous, dark brown, visible as small black dot on the host
surface, ostiolate, apapillate, with pore-like opening.
Conidiomata walls 8–25 lm wide, thin-walled, composed
of flattened to broad, brown to dark brown pseudoparenchymatous cells, arranged in a textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–7(–12) 9 (2–)3–6 lm (
x = 5.7 9 4.7 lm,
n = 30), enteroblastic, phialidic, discrete, determinate,
ampulliform to doliiform, occasionally cylindrical,
unbranched, aseptate, hyaline, smooth, minute collarette
with periclinal wall thickening, arising from the inner
cavity of the conidioma wall, difficult to distinguish from
the conidioma wall. Conidia (30–)35–45 9 (10–)11–14(–
15.5) lm (
x = 39.6 9 12.9 lm, n = 50), muriform, brown
to dark brown, paler at the end cells, ellipsoidal to broadly
fusiform, or subclavate, apex rounded, base truncate or
obtuse, 8–9 transverse septa, with 2–4 longitudinal septa,
not constricted at the septa, rough-walled, echinulate,
lacking mucilaginous sheath surrounding conidia.
Material examined: CHINA, Yunnan Province,
Baoshan, Shuizai, Dawazi mountain, on dead branches of
herbaceous plant, 23 October 2015, R. Phookamsak,
BS018 (KUN-HKAS 102234, holotype).
GenBank
numbers:
ITS = MK356374,
LSU =
MK356348, SSU = MK356362, TEF1-a = MK359066.
Notes: Multigene phylogenetic analyses of a combined
LSU, SSU, TEF1-a and ITS sequence dataset show that
Galiicola baoshanensis forms a sister lineage with the
generic type of Galiicola, G. pseudophaeosphaeria with
high support (100% ML and 1.00 BYPP; Fig. 29). A
comparison of TEF1-a sequences indicates that G.
baoshanensis differs from G. pseudophaeosphaeria in
14/730 bp (1.9%). However, we could not compare the
morphological characters of G. baoshanensis with G.
pseudophaeosphaeria as they are represented by different
morphs. Galiicola baoshanensis is introduced as the
asexual species in Galiicola for the first time.
Galiicola baoshanensis is similar to Amarenographium
ammophilae Wanas. et al. in having conidia that are
muriform, clavate, ellipsoidal, ovoid or fusoid conidia,
with rounded apex, acute or truncate base, and 7–9 transverse septa (Wijayawardene et al. 2016). However, G.
123
baoshanensis can be distinguished from A. ammophilae in
having brown to dark brown conidia, with 2–4 longitudinal
septa and lacking appendages at the apex and the base.
Whereas, A. ammophilae has yellowish brown to brown
conidia, with 1–3 longitudinal septa and appendages at the
apex and the base (Wijayawardene et al. 2016). A comparison of ITS and TEF1-a sequences indicates that G.
baoshanensis differs from A. ammophilae in 71/570 bp
(12.5%) and 58/899 bp (6.5%), respectively. Muti-gene
phylogenetic analyses also supported their distinctiveness
(Fig. 29).
Neosetophoma Gruyter et al.
We follow the latest treatment and updated accounts of
Neosetophoma in Hyde et al. (2018b) and Wanasinghe
et al. (2018). Seventeen species are known in this genus
(Hyde et al. 2018b; Index Fungorum 2019).
Neosetophoma lonicerae Phookamsak, Wanas. & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556168; Facesoffungi
number: FoF05702, Fig. 33
Etymology: The specific epithet ‘‘lonicerae’’ refers to
the host genus Lonicera, from which the holotype was
collected.
Holotype: KUN-HKAS 102231
Saprobic on Lonicera maackii. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
110–160 lm high, 80–160 lm diam., pycnidial, scattered,
solitary to gregarious, immersed to semi-immersed, uniloculate, globose to subglobose, glabrous, dark brown to
black, visible as small black dot on the host surface,
associating with other fungal taxa, ostiole central, occasionally near the centre, minutely mammiform papilla.
Conidiomata walls 5–12 lm wide, equally thin-walled,
composed of 2–3 cell layers, of broad, brown to dark
brown pseudoparenchymatous cells, arranged in a textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous
cells
3.5–7 9 (3–)5–10 lm
(
x = 6.2 9 6.7 lm, n = 20), enteroblastic, phialidic, discrete, determinate, ampulliform to doliiform, unbranched,
aseptate, hyaline, smooth, minute collarette with periclinal
wall thickening, arising from the inner cavity of the conidioma
wall.
Conidia
(8.5–)9–12(–14) 9 4–5 lm
(
x = 11 9 4.8 lm, n = 50), yellowish brown, ellipsoidal,
1–3-septate, not constricted at the septa, smooth-walled,
lacking mucilaginous sheath surrounding conidia.
Culture characteristics: Colonies on PDA reaching
33–35 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony medium dense, circular, flattened to
raised, surface slightly rough with hyphal tufts, edge entire,
velvety to fluffy; from above, white to white yellowish at
the margin, light green to yellowish green at the centre;
from below, radiating outwards colony, white to cream at
Fungal Diversity
Fig. 31 Brunneomurispora lonicerae (KUN-HKAS 102232, holotype). a Appearance of ascomata on host. b Close up of ascoma on
host. c Section through ascoma. d Section through peridium. e Cellular
pseudoparaphyses. f–i Asci. j–o Ascospores. Scale bars a = 500 lm,
b = 200 lm, c = 100 lm, d = 50 lm, e–i = 20 lm, j–o = 10 lm
the margin, dark green to black at the middle, orangish
brown at the centre; producing yellowish pigment on agar
medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead hanging
branches of Lonicera maackii, 20 April 2017, R.
Phookamsak, KIB033 (KUN-HKAS 102231, holotype),
ex-type living culture, KUMCC 18-0155 (KIB033A),
KUMCC 18-0156 (KIB033B).
GenBank
numbers:
ITS = MK356375,
LSU =
MK356349, SSU = MK356363, TEF1-a = MK359067
(KUMCC
18-0155);
ITS = MK356376,
LSU =
MK356350, SSU = MK356364, TEF1-a = MK359068
(KUMCC 18-0156).
Notes: In the NCBI BLASTn search of ITS sequence,
Neosetophoma lonicerae has a closest match with fungal
endophyte species (M16-3161) with 100% similarity and is
closely related to N. italica W.J. Li et al., N. rosarum R.H.
Perera et al., N. samarorum (Desm.) Gruyter et al. and N.
rosigena Wanas. et al. with 98% similarity. Multigene
phylogenetic analyses based on a combined LSU, SSU,
TEF1-a and ITS sequence dataset show that N. lonicerae
forms a separate lineage, clustering with N. rosigena, N.
samarorum and N. garethjonesii Tibpromma et al. with
moderate support (76% ML and 0.98 BYPP; Fig. 29). A
comparison of ITS nucleotide base pairs shows that N.
lonicerae differs from N. rosigena, N. samarorum and N.
garethjonesii in 14/555 bp (2.5%), 15/555 bp (2.7%) and
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Fungal Diversity
Fig. 32 Galiicola baoshanensis (KUN-HKAS 102234, holotype).
a Appearance of conidiomata on host. b Section through conidioma.
c Ostiole with pore-like opening. d Section through conidioma wall.
e–g Conidiogenous cells. h Conidium when immature. i–m Conidia.
Scale bars a = 200 lm, b = 50 lm, d = 20 lm, c, g–m = 10 lm, e,
f = 5 lm
16/522 bp (3%), respectively. We therefore, introduce N.
lonicerae as a new species following the guidelines of
Jeewon and Hyde (2016).
Neosetophoma lonicerae is similar to some other
Neosetophoma species in having pale brown to brown,
oblong to ellipsoidal, or subfusoid, 1–3-septate conidia;
such as in N. iranianum Papizadeh et al., N. italica, N.
rosae R.H. Perera et al., N. rosarum and N. shoemakeri
Senwanna et al. However, they can be distinguished based
on their conidial size, host occurrence and phylogenetic
distance (Liu et al. 2015a; Karunarathna et al. 2017; Hyde
et al. 2018b; Wanasinghe et al. 2018).
Ophiobolus Riess
We follow the latest treatment and updated accounts of
Ophiobolus in Phookamsak et al. (2017).
123
Ophiobolus malleolus S.K. Huang, Bulgakov & K.D.
Hyde, sp. nov.
Index Fungorum number: IF554782; Facesoffungi
number: FoF04686, Fig. 34
Etymology: The specific name ‘‘malleolus’’ refers to the
hammer-shaped at top of the ascospores.
Holotype: MFLU 15-2230.
Saprobic on dead stems of Cirsium arvense (Asteraceae). Sexual morph Ascomata 230–270 lm high, 265–
310 lm diam., scattered, solitary to gregarious, immersed,
eventually erumpent, uni-loculate, globose to subglobose,
Fungal Diversity
Fig. 33 Neosetophoma lonicerae (KUN-HKAS 102231, holotype).
a Appearance of conidiomata on host. b Section through conidioma.
c, d Section through conidioma wall. e–h Conidiogenous cells. i–n
Conidia. o Germinated conidium. p, q Culture characteristics on PDA
(p = from above, q = from below). Scale bars a = 200 lm,
b = 50 lm, c–h, o = 10 lm, i–n = 5 lm
dark brown to black, papillate. Ostiole central, short
papilla, filled with periphyses. Peridium 25–60 lm wide,
thick-walled, composed of several cell layers of brown to
dark brown cells, paler to hyaline towards the inner layers,
outer layer comprising black, coriaceous stratum, arranged
in a textura angularis. Hamathecium composed of
numerous, 1–3 lm wide, filamentous, septate pseudoparaphyses, anastomosing at the apex, embedded in a gelatinous matrix. Asci 125–155 9 12–15 lm (
x = 140 9
12.5 lm, n = 20), 8-spored, bitunicate, cylindric-clavate,
subsessile to short pedicellate, with knob-like or truncate
pedicel, rounded at the apex, with a distinct ocular chamber. Ascospores 100–112 9 2.5–4.5 lm (
x = 107.5 9
3.5 lm, n = 50), fasciculate, in parallel or spiral, initially
hyaline, becoming brown at maturity, guttulate, filiform,
enlarged at the first cell with hammer-like, tapering
towards the end cell, up to 15 septa, curved, slightly constricted at the septa at maturity, smooth-walled, with apical
mucilaginous cap. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on PDA,
colony reaching 10 mm diam. after 2 weeks at room temperature (20–30 °C). Colony dense, irregular in shape, flattened to umbonate, surface smooth, with edge erose, velvety
to floccose; from above, cream; from below, cream to pale
yellowish; not producing pigmentation on agar medium.
Material examined: RUSSIA, Rostov region, Krasnosulinsky District, Donskoye forestry, arboretum
(47.8547249̊N, 40.2318907̊E), on dead stems of Cirsium
arvense (Asteraceae), 28 June 2015, T.S. Bulgakov, T-526
(MFLU 15-2230, holotype), ex-type living culture,
MFLUCC 15-1077.
GenBank
numbers:
ITS = MH399730,
LSU =
MH399731, SSU = MH399729.
Notes: Multigene phylogenetic analyses reveal a close
phylogenetic affinity between Ophiobolus malleolus and O.
disseminans (Fig. 29, 68% ML and 1.00 BYPP).
Ophiobolus malleolus is distinct from O. disseminans in its
ascospores being filiform with an enlarged at the apical cell
similar to a hammer-like, with apical mucilaginous cap,
123
Fungal Diversity
and not splitting into two part spores. Ophiobolus disseminans has filiform ascospores with two swollen cells near
the centre, lacks an apical mucilaginous cap, and splits into
two part spores at the central septum (Phookamsak et al.
2014b, 2017).
Ophiobolus anguillides (Cooke) Sacc. also has filiform
ascospores enlarged hammer-like at the apical cell and an
apical mucilaginous cap. However, O. anguillides has
larger ascomata and ascospores (500–600 lm diam., and
120–130 9 2.5–3 lm; Shoemaker 1976). Ophiobolus
anguillides has been reported from many hosts, mainly on
Artemesia in Europe and North America (Shoemaker 1976;
Farr and Rossman 2018). However, the species has never
been reported from Cirsium arvense. Unfortunately, there
is no molecular data available for O. anguillides.
Phaeosphaeria I. Miyake
We follow the latest treatment and updated accounts of
Phaeosphaeria in Phookamsak et al. (2014b), Hyde et al.
(2017) and Tibpromma et al. (2017). More than 200 epithets are listed under Phaeosphaeria in Index Fungorum
(2019); however, the phylogenetic affinities of few species
have been confirmed based on molecular data. Some species listed under Phaeosphaeria have already been transferred to other related genera in Phaeosphaeriaceae and
other related families (Phookamsak et al. 2014b; Ariyawansa et al. 2015a; Tennakoon et al. 2016).
Phaeosphaeria cycadis Wanas., Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556169; Facesoffungi
number: FoF05703, Fig. 35
Etymology: The specific name ‘‘cycadis’’ refers to the
host family Cycadaceae, of which the holotype was
collected.
Holotype: KUN-HKAS 102235
Associated with leaf spots on Cycas sp. (Cycadaceae).
Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata (60–)100–160 lm high, (60–)
135–200 lm diam., pycnidial, visible as black dot on host
surface, scattered to clustered, solitary to gregarious, semiimmersed to erumpent, uni-loculate, subglobose to ampulliform, or irregular in shape, ostiolate, with minute papilla.
Conidiomata walls 10–20(–30) lm wide, composed of 4–5
cell layers, of brown to dark brown pseudoparenchymatous
cells, of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells (3.5–)4.5–8 9 4–8
(–10) lm (
x = 6.4 9 5.7 lm, n = 30), ampulliform to
broadly conical, gradually tapering toward the apex,
holoblastic, phialidic, hyaline, smooth-walled, with periclinal wall thickening, arising from the inner cavity of the
conidioma wall. Conidia (10–)12–14(– 16) 9 3–5 lm
(
x = 12.9 9 3.9 lm, n = 50), pale brown to light yellowish,
oblong to ellipsoidal, or subcylindrical, (0–)1–2-septate,
123
truncate to obtuse base, with obtuse apex, not constricted at
the septa, smooth-walled.
Culture characteristics: Colonies on PDA reaching
50–52 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony dense, irregular in shape, flattened,
surface slightly smooth, edge undulate, with entire margin,
cottony to floccose, slightly sparse near the margin; from
above, white to cream, slightly radiated outwards colony;
from below, slightly radiating, cream at the margin, pale
brown to yellowish-brown at the centre, sectering with
golden brown; not producing pigmentation on agar
medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, associated with
leaf spots on Cycas sp. (Cycadaceae), 5 April 2017, R.
Phookamsak, KIB022 (KUN-HKAS 102235, holotype),
ex-type living culture, KUMCC 18-0161 (KIB022A),
KUMCC 18-0162 (KIB022B).
GenBank
numbers:
ITS = MK356378,
LSU =
MK356352, SSU = MK356366, TEF1-a = MK359069
(KUMCC
18-0161);
ITS = MK356379,
LSU =
MK356353, SSU = MK356367, TEF1-a = MK359070
(KUMCC 18-0162).
Notes: In the NCBI BLASTn search of ITS sequences,
Phaeosphaeria cycadis most closely matches P. acaciae
Tennakoon et al. with 99% similarity. Multigene phylogenetic analyses based on a combined LSU, SSU, TEF1-a
and ITS sequence dataset show that P. cycadis forms a
sister lineage with P. acaciae with high support (100% ML
and 1.00 BYPP; Fig. 29). The ITS gene could not clarify
the novelty of P. cycadis in this study as the species is not
significantly different from P. acaciae in a comparison of
ITS sequences (5/452 bp). Phookamsak et al. (2014b)
mentioned that Phaeosphaeria contains species complexes
that cannot be resolved based only on the ITS gene and that
a combination of protein coding genes, such as TEF1-a and
RPB2, is necessary to clarify species in this genus. Based
on morphological characteristics, P. cycadis differs from P.
acaciae in having larger conidia (P. cycadis, (10–)12–14
(– 16) 9 3–5 lm versus 8–12 9 2.4–3.5 lm, P. acaciae;
Hyde et al. 2017), with (0–)1–2 conidial septa and is
associated with leaf spots on Cycas sp. Phaeosphaeria
acaciae has 1–3 conidial septa and occurs on dead stems of
Acacia sp. as a saprobe (Hyde et al. 2017). Furthermore, P.
acaciae produced a pink pigment on PDA, but this is
absent in P. cycadis. We therefore, introduce P. cycadis as
a new species in this study based on its morphological
distinctiveness.
Wojnowiciella Crous et al.
Wojnowiciella was introduced by Crous et al. (2015b)
and is typified by W. eucalypti Crous et al. Crous et al.
(2015a) treated Wojnowicia as a synonym of Septoriella
Fungal Diversity
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Fungal Diversity
b Fig. 34 Ophiobolus malleolus (MFLU 15-2230, holotype) a Herbar-
ium label. b Appearance of ascomata on Cirsium arvense stems.
c Ascomata on host. d Ascoma in vertical section. e Peridium.
f Pseudoparaphyses. g–i Developing stages of the asci. j–m Ascospores (note: m stained in Indian ink). Scale bars c = 500 lm,
d = 100 lm, e = 50 lm, f–m = 20 lm
Oudem. based on nomenclature study and this is in
agreement of Wijayawardene et al. (2017a). Subsequently,
all identified Wojnowicia species in Phaeosphaeriaceae
were synonymized under Wojnowiciella (Crous et al.
2015b; Hernandez-Restrepo et al. 2016). Wojnowiciella
can be distinguished from Wojnowicia Sacc. in having
apapillate, glabrous conidiomata and dark brown conidia
(Crous et al. 2015b). Wojnowicia was introduced by Saccardo (1892) and is characterized by setose conidiomata
(Crous et al. 2015a; Wijayawardene et al. 2016). Seven
species are accommodated in this genus (Index Fungorum
2019).
Wojnowiciella kunmingensis Phookamsak, Wanas. &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556170; Facesoffungi
number: FoF05704, Fig. 36
Etymology: The specific epithet ‘‘kunmingensis’’ refers
to Kunming Institute of Botany, Kunming City, Yunnan
Province, China, where the holotype was collected.
Holotype: KUN-HKAS 102233
Saprobic on Lonicera maackii. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
110–190 lm high, 110–190 lm diam., pycnidial, scattered,
solitary to gregarious, immersed, slightly raised, visible as
small black dot on host surface, uni-loculate, globose to
subglobose, glabrous, dark brown to black, ostioles central,
apapillate, with pore-like opening. Conidiomata walls 5–
12 lm wide, thin-walled, of equal thickness, composed of
1–3 cell layers, of flattened, dark brown, pseudoparenchymatous cells, arranged in a textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells (3–)4–6.5(–9) 9 (2.5–)3.5–6(–8) lm l/w
(
x = 5 9 5.3 lm, n = 40), holoblastic, phialidic, discrete,
determinate, ampulliform to doliiform, unbranched, aseptate, hyaline, smooth, arising from the inner cavity of the
conidioma wall. Conidia (16–)18–24(–27.5) 9 4–6(–7)
lm l/w (
x = 22 9 5.6 lm, n = 50), dark brown, subcylindrical to fusiform, or reniform, slightly curved, with
acute to rounded apex, and rounded to truncate base, 3–7septate, not constricted at the septa, thick-walled, smoothwalled, with guttules, having flattened, mucous caps at both
ends.
Culture characteristics: Colonies on PDA reaching
35–38 mm diam. after 2 weeks at room temperature
(20–30 °C). Colony medium dense, slightly irregular in
123
shape, flattened to slightly raised, surface slightly rough
with greyish-green hyphal tufts, edge undulate, with entire
margin, floccose; from above, slightly radiating, white to
cream at the margin, pale brown at the middle, separated
from the margin with brown to dark green concentric ring
near the edge, greyish green to dark green at the centre;
from below, white to cream margin, brown to dark brown
at the centre; not producing pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead hanging
branches of Lonicera maackii, 20 April 2017, R.
Phookamsak, KIB031 (KUN-HKAS 102233, holotype),
ex-type living culture, KUMCC 18-0159 (KIB031A),
KUMCC 18-0160 (KIB031B).
GenBank
numbers:
ITS = MK356380,
LSU =
MK356354, SSU = MK356368, TEF1-a = MK359071
(KUMCC
18-0159);
ITS = MK356381,
LSU =
MK356355, SSU = MK356369, TEF1-a = MK359072,
RPB2 = MK359078 (KUMCC 18-0160).
Notes: In the NCBI BLASTn search of ITS sequences,
Wojnowiciella kunmingensis most closely matches W.
dactylidis (Wijayaw. et al.) Hern.-Restr. & Crous, W.
spartii (W.J. Li et al.) Hern.-Restr. & Crous and Wojnowicia italica Qing Tian et al. with 99% similarity. ITS
and TEF1-a genes could not resolve the novelty of the
Wojnowiciella species in this study. A comparison of ITS
and TEF1-a nucleotide bases shows that Wojnowiciella
kunmingensis is not significantly different from W. dactylidis and other Wojnowiciella species as well as Wojnowicia italica. However, a comparison of RPB2 sequence
data shows that Wojnowiciella kunmingensis differs from
W. dactylidis and Wojnowicia italica in 21/856 bp (2.4%)
and 20/954 bp (2.1%), respectively.
Multigene phylogenetic analyses based on a combined
LSU, SSU, TEF1-a and ITS sequence data show that
Wojnowiciella kunmingensis forms a sister lineage with W.
dactylidis and clusters with other Wojnowiciella species
(Fig. 29). However, W. kunmingensis differs from W.
dactylidis in having 3–7-septate conidia, while W. dactylidis has 7–11-septate conidia (Liu et al. 2015a). Wojnowiciella leptocarpi Crous et al. also has 3–7-septate
conidia, with mucous caps at both ends. However, W.
kunmingensis has longer conidia ((16–)18–24(–
27.5) 9 4–6(–7) lm versus 26–36 9 4–6 lm; HernandezRestrepo et al. 2016), that are brown to dark brown, and
smooth-walled. The conidia of W. leptocarpi are orange
brown, thick-walled, and verruculose (Hernandez-Restrepo
et al. 2016). A comparison of RPB2 sequence data shows
that W. kunmingensis differs from W. leptocarpi in
29/851 bp (3.4%). Wojnowiciella lonicerae (Wijayaw.
et al.) Hern.-Restr. & Crous was also collected from Lonicera, but it has 8–11-septate conidia (Liu et al. 2015a).
Fungal Diversity
Fig. 35 Phaeosphaeria cycadis (KUN-HKAS 102235, holotype).
a Appearance of conidiomata on host. b, c Section through conidiomata. d Section through conidioma wall. e, f Conidiogenous cells.
g–k Conidia. Scale bars a = 100 lm, b = 50 lm, c, d = 20 lm, e,
g = 10 lm, f, h–k = 5 lm
Wojnowiciella rosicola (W.J. Li et al.) Wanas.,
Phookamsak & K.D. Hyde, comb. nov.
Index Fungorum number: IF556171; Facesoffungi
number: FoF05705
Basionym: Wojnowicia rosicola W.J. Li, Camporesi &
K.D. Hyde, in Wanasinghe et al., Fungal Divers.: https://
doi.org/10.1007/s13225-018-0395-7, [144] (2018)
Holotype: ITALY, Arezzo [AR], Montemezzano, on
dead aerial branch of Rosa sp. (Rosaceae), 25 August 2014,
E. Camporesi, IT 2200 (MFLU 17-2785); ex-type living
culture, MFLUCC 15-0128.
Morphological description: See Wanasinghe et al.
(2018) (Fig. 93, pp. 144–147).
Notes: Wojnowiciella rosicola was introduced by
Wanasinghe et al. (2018) as Wojnowicia rosicola W.J. Li
et al. The species clustered with other Wojnowiciella species in Phaeosphaeriaceae that were previously treated in
Wojnowicia and recently treated as a synonym of Wojnowiciella by Crous et al. (2015a, b) and HernandezRestrepo et al. (2016). Based on morphological characteristics and phylogenetic analyses, the species is congeneric
with Wojnowiciella. We therefore, treat W. rosicola as a
new combination.
Pseudoastrosphaeriellaceae Phookamsak & K.D. Hyde
Pseudoastrosphaeriellaceae was introduced by Phookamsak
et al. (2015b) to accommodate a monotypic genus
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Fungal Diversity
Fig. 36 Wojnowiciella kunmingensis (KUN-HKAS 102233, holotype). a Appearance of conidiomata on host. b Section through
conidioma. c Section through conidioma wall. d–g Conidiogenous
cells. h–k Conidia. l Germinated conidium. Scale bars a = 200 lm,
b = 50 lm, c, i = 20 lm, h, j–l = 10 lm, d–g = 5 lm
Pseudoastrosphaeriella. This family is characterized by
hemispherical to lenticular ascostromata, globose to subglobose ascomata with a flattened or rounded base, immersed
beneath host epidermis, erumpent through host surface by a
papilla, with short to long necks, trabeculate pseudoparaphyses, short pedicellate, cylindric-clavate to clavate asci,
and hyaline or brown, septate ascospores and coelomycetous
asexual morphs (Phookamsak et al. 2015b). Based on close
morphological characteristics and phylogenetic support,
Hyde et al. (2017) accommodated Carinispora K.D. Hyde in
Pseudoastrosphaeriellaceae and this was followed by
Wijayawardene et al. (2018a). In this study, we introduce a
123
Fungal Diversity
new genus Pseudoastrosphaeriellopsis typified by P. kaveriana in Pseudoastrosphaeriellaceae based on molecular
phylogeny coupled with morphological characteristics.
Pseudoastrosphaeriellopsis Devadatha, Wanas., Jeewon &
V.V. Sarma, gen. nov.
Index Fungorum number: IF555790; Facesoffungi
number: FoF05706
Etymology: Generic epithet in resemblance to
Pseudoastrosphaeriella
Saprobic on decaying stems and twigs of Avicennia
marina and Suaeda monoica, black spots, with short necks
on host surface. Sexual morph Ascomata light brown,
solitary to gregarious, immersed to erumpent, globose to
subglobose with a rounded base, uni-loculate, coriaceous,
brown, short papillate, ostiolate. Ostioles central, cylindrical, straight to oblique, with ostiolar canal lined by
hyaline periphyses. Peridium thin- to thick-walled,
unequally thickened, composed of two layers, inner stratum with hyaline to pale brown compressed cells of textura
angularis, outer stratum with compact brown polygonal
cells of textura angularis, fused with the host tissue. Hamathecium composed of numerous, filamentous, septate,
branched, hyaline pseudoparaphyses. Asci 8-spored, bitunicate, cylindric-clavate to clavate, with a short, thick
pedicel, apically rounded and thickened with an ocular
chamber. Ascospores overlapping 1–2-seriate, brown,
fusiform to broadly fusiform, slightly curved with round to
acute ends, slightly constricted at the 3rd septum and rarely
at other septa, broader in the middle and tapering towards
the ends, septate, with hyaline apical and terminal ends,
smooth-walled, lacking guttules and mucilaginous sheath.
Asexual morph Undetermined.
Type species: Pseudoastrosphaeriellopsis kaveriana
Devadatha, Wanas., Jeewon & V.V. Sarma
Notes: The genus Pseudoastrosphaeriella Phookamsak
et al. was introduced by Phookamsak et al. (2015b) to
accommodate P. aequatoriensis (K.D. Hyde & J. Fröhl.)
Phookamsak & K.D. Hyde, P. africana (D. Hawksw.)
Phookamsak & K.D. Hyde, P. bambusae Phookamsak &
K.D. Hyde, P. longicolla Phookamsak & K.D. Hyde, P.
papillata (K.D. Hyde & J. Fröhl.) Phookamsak & K.D.
Hyde and P. thailandensis Phookamsak et al. based on
morphology and multigene phylogenetic analyses. Some of
the species were previously treated in Astrosphaeriella
Syd. & P. Syd. Maximum likelihood tree (Fig. 37) generated based on a combined LSU, SSU, ITS, RPB2 and
TEF1-a dataset reveals that our new taxon clusters within
Pseudoastrosphaeriellaceae sister to Pseudoastrosphaeriella with 63% ML and 0.95 BYPP statistical support. Tree topologies generated under ML and Bayesian
criteria from combined datasets were congruent, whereas
maximum parsimonious tree was varied, which showed
that our taxon forms a clade between Testudinaceae Arx
and Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray. without any statistical support. Further comparison of LSU
nucleotides between our taxon and Pseudoastrosphaeriella
thailandensis (GenBank: KT955478) resulted in 62/871
(7.1%) base pair differences which confirms its placement
as a new genus in Pseudoastrosphaeriellaceae.
Pseudoastrosphaeriellopsis shares morphological similarity with Pseudoastrosphaeriella in having immersed
ascomata underneath the host epidermis, erumpent, uniloculate, coriaceous, brown, and short papillate asci, with
trabeculate pseudoparaphyses (Phookamsak et al. 2015b).
However, Pseudoastrosphaeriellopsis can be distinguished
from Pseudoastrosphaeriella in having globose to subglobose ascomata, immersed in host tissue, with fusiform
to broadly fusiform ascospores.
Pseudoastrosphaeriellopsis kaveriana resembles Neotrematosphaeria biappendiculata (Kaz. Tanaka et al.)
Thambug. et al., but differs in having immersed ascomata,
with pore-like ostiole, and ascospores lacking appendages,
whereas, the latter has semi-immersed ascomata, with
crest-like ostiole, and ascospores with appendages at both
ends (Thambugala et al. 2015). Biappendiculispora
japonica Thambug. et al. differs from Pseudoastrosphaeriellopsis kaveriana in having slit-like ostioles, bulbous pedicels and 7–9 transverse septa ascospores with
appendages (Thambugala et al. 2015). Pseudoastrosphaeriellopsis kaveriana also shares similarities with
Trematosphaeria wegeliniana L. Holm & K. Holm, T.
hydrophila Sacc. and T. crassisepta Kaz. Tanaka et al. in
having 5-septate ascospores and overlapping ascospore
dimensions, but is clearly distinct in having immersed
ascomata beneath the host epidermis, short papilla and
light brown, fusiform to broadly fusiform, slightly curved
ascospores (Tanaka et al. 2015). Trematosphaeria
wegeliniana and T. hydrophila lack a detailed description
from type material and their taxonomic position remains
unclear (Tanaka et al. 2015).
Thambugala et al. (2015) transferred Trematosphaeria
terricola G.S. Gong to Alpestrisphaeria terricola (G.S.
Gong) Thambug. & K.D. Hyde and Trematosphaeria
biappendiculata (KTC 1124) to the new genus Neotrematosphaeria biappendiculata based on their morphological resemblance to Lophiostomataceae and multigene
phylogeny. Our combined multiloci phylogenetic analysis
also reveals that taxonomic position of some species in
Trematosphaeria remains unclear and this needs further
collection and revision.
Pseudoastrosphaeriellopsis kaveriana Devadatha, Wanas.,
Jeewon & V.V. Sarma, sp. nov.
123
Fungal Diversity
123
Fungal Diversity
b Fig. 37 Phylogram generated from maximum likelihood analysis
based on LSU, SSU, ITS, RPB2 and TEF1-a sequence dataset of
representative families in Pleosporales showing phylogenetic affinities of Pseudoastrosphaeriellopsis kaveriana. Gloniopsis calami
(MFLUCC 15-0739) and Hysterium rhizophorae (NFCCI-4250) were
selected as the outgroup taxa. Bootstrap support values for maximum
likelihood (green), equal to or greater than 60% and Bayesian
posterior probabilities (purple) equal to or greater than 0.95 are given
above each branch, respectively. The new isolate is in blue. Type
strains are in bold
Index Fungorum number: IF555791; Facesoffungi
number: FoF05707, Fig. 38
Etymology: Specific epithet in reference to the river
Kaveri.
Holotype: AMH-9912
Saprobic on decaying stems and twigs of Avicennia
marina and Suaeda monoica, black spots ascomata, with
short necks on host surface. Sexual morph Ascomata 225–
335 lm high, 220–345 lm diam., immersed to erumpent,
globose to subglobose, solitary to gregarious, coriaceous,
brown, short papillate, ostiolate. Ostioles 110–165 lm
long, 50–90 lm diam., central, cylindrical, straight to
oblique, with ostiolar canal lined by hyaline periphyses.
Peridium 15–25 lm wide, less developed at the base
compared to the sides and top, composed of two layers,
inner stratum with 3–5 layers of hyaline to pale brown
compressed cells of textura angularis, outer stratum with
compact brown polygonal cells of textura angularis, fused
with the host cells and fungal tissue. Hamathecium composed of numerous, 1–3 lm wide, filamentous, septate,
branched, hyaline pseudoparaphyses, embedded in a
gelatinous matrix, anastomosing between and above the
asci. Asci 85–145 9 15–18 lm (
x = 108 9 17 lm,
n = 40), 8-spored, bitunicate, cylindric-clavate to clavate,
with a short, thick pedicel, apically rounded and thickened,
with an ocular chamber. Ascospores 25–40 9 5–10 lm
(
x = 34 9 7 lm, n = 50), overlapping 1–2-seriate, brown,
fusiform to broadly fusiform, slightly curved with round to
acute ends, broader in the middle and tapering towards the
ends, 5-septate, slightly constricted at the 3rd septum and
rarely at other septa, smooth-walled, hyaline at both ends,
from the central septum upper half is shorter and lower half
is longer, lacking guttules and mucilaginous sheath.
Asexual morph Undetermined.
Culture characteristics: Ascospores germinated on
SWA within 24 h, germ tubes arising from terminal ends of
the ascospore. Colonies on MEA reaching 30–45 mm
diam. after 25 days of incubation at room temperature, pale
olive buff to olive buff, reverse brown, velvety, surface
raised, irregular, with light yellow exudates.
Material examined: INDIA, Tamil Nadu, Tiruvarur,
Muthupet mangroves (10.4°N 79.5°E), on decaying stems
and twigs of Avicennia marina (Forssk.) Vierh. (Acanthaceae), 28 November 2015, B. Devadatha, AMH-9912
(holotype); isotype at Pondicherry University, Puducherry), ex-type living culture, NFCCI-4221; ibid. on stems
and twigs of Suaeda monoica Forssk. ex J.F. Gmel), 28
November 2015, B. Devadatha, PUHD33.
GenBank
numbers:
ITS = MG947599,
LSU =
MG947595, SSU = MG947598, TEF1-a = MG968955,
RPB2 = MG948547.
Roussoellaceae Jian K. Liu et al.
We follow the latest treatment and updated accounts of
Roussoellaceae in Liu et al. (2014), Dai et al. (2017a),
Tibpromma et al. (2017), Hyde et al. (2018b), Jayasiri et al.
(2019) and Jiang et al. (2019). Phylogenetic affinities of the
family were discussed by Jaklitsch and Voglmayr (2016)
where Roussoellaceae was treated as a synonym of
Thyridariaceae. However, Tibpromma et al. (2017) reinstated Roussoellaceae based on multigene phylogenetic
analysis which showed Roussoellaceae is a well-resolved
family in Pleosporales and this was concurred by Wanasinghe et al. (2018), Wijayawardene et al. (2018a), Jayasiri
et al. (2019) and Jiang et al. (2019). Seven genera were
accepted in this family based on molecular data and morphological characteristics (Wijayawardene et al. 2018a).
Taxonomic status of genera in Roussoellaceae is
doubtful and needs to be clarified due to taxa in these
genera have similar conidial characters and phylogeny has
always shown that they are monophyletic (Liu et al. 2014;
Dai et al. 2017a; Tibpromma et al. 2017; Hyde et al. 2018b;
Jiang et al. 2019). However, Neoroussoella Phookamsak
et al. and Roussoellopsis I. Hino & Katum. are distinct
from Roussoella but these two genera usually cluster with
Roussoella species (Liu et al. 2014; Phookamsak et al.
2014a; Dai et al. 2017a; Tibpromma et al. 2017; Hyde et al.
2018b; Jiang et al. 2019). Wanasinghe et al. (2018) introduced Pseudoneoconiothyrium (Phukhams. et al.) Phukhams. et al. and Pararoussoella to accommodate
roussoella-like taxa in Thyridariaceae. We use increased
taxon sampling in our phylogenetic analysis (Fig. 39) and
show that the two genera erected by Wanasinghe et al.
(2018), cluster with Roussoella species in Roussoellaceae.
Jayasiri et al. (2019) clarified Neoroussoella based on
multigene phylogenetic analysis coupled with morphological characteristics, N. entadae Jayasiri et al. and N. leucaenae Jayasiri et al. were introduced. Roussoella solani
Crous & M.J. Wingf. was transferred to Neoroussoella as
N. solani (Crous & M.J. Wingf.) Jayasiri & K.D. Hyde. In
addition, Roussoella mukdahanensis Phookamsak et al.
was transferred to Pararoussoella as P. mukdahanensis
(Phookamsak et al.) Jayasiri & K.D. Hyde. In this study,
Neoroussoella heveae Senwanna et al. and Roussoella
elaeicola Konta & K.D. Hyde are introduced as novel
123
Fungal Diversity
species based on morphological characteristics coupled
with multigene phylogenetic analysis. In addition, Neoroussoella leucaenae is described on Hevea brasiliensis
Müll.Arg. from Thailand for the first time.
Neoroussoella J.K. Liu et al.
We follow the latest treatment and updated accounts of
Neoroussoella in Jayasiri et al. (2019).
Neoroussoella heveae Senwanna, Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF555287; Facesoffungi
number: FoF04825, Fig. 40
Etymology: Name refers to the host genus Hevea, from
which the fungus was collected.
Holotype: MFLU 17-1983
Saprobic on Para rubber (Hevea brasiliensis). Sexual
morph Undetermined. Asexual morph Coelomycetous.
Conidiomata 90–130 lm high, 115–180 lm diam., pycnidial, immersed, visible as small, brown to black dots on the
host surface, solitary, scattered, globose, uni-loculate,
glabrous, ostiole not observed. Conidiomata walls 7–
15 lm wide, thick-walled, composed of several cell layers
of dark brown to black, pseudoparenchymatous cells of
textura angularis. Conidiophores reduced to conidiogenous
cells.
Conidiogenous
cells
3–7 9 2–5 lm
(
x = 5.2 9 3.4 lm, n = 15), lining the inner cavity, difficult to distinguish from pycnidial wall, enteroblastic,
phialidic, integrated, simple, short, hyaline, ampulliform to
doliiform, thin-walled, smooth. Conidia 2.5–5 9 2–4 lm
(
x = 4 9 3, n = 50), globose or subglobose to ellipsoidal,
initially hyaline, becoming brown to dark brown, aseptate,
with one or two guttules, rough-walled, verruculose.
Culture characteristics: Colonies on MEA reaching
20–25 mm diam. after 2 weeks at 25–30 °C, colony from
above, circular, medium dense, edge entire, velvety to
woolly, white at the margin, white to yellowish in the
middle, geenish grey to grey at the centre; from below,
white at the margin, greenish grey at the centre, with
concentric rings radiate.
Material examined: THAILAND, Phayao Province,
Muang District, Mae Ka, on twig of Hevea brasiliensis
(Euphorbiaceae), 5 December 2016, C. Senwanna,
RBPY018 (MFLU 17-1983, holotype), ex-type living
culture, MFLUCC 17-0338.
GenBank
numbers:
ITS = MH590693,
LSU =
MH590689, SSU = MH590691.
Notes: Neoroussoella heveae is introduced based on
morphological and phylogenetic evidence. Our strain forms
a separate lineage, clusters with Neoroussoella species
(78% ML; Fig. 39). Neoroussoella heveae differs from the
asexual morph of Neoroussoella species in having verruculose conidia, whereas, other Neoroussoella species have
smooth-walled conidia.
123
Neoroussoella leucaenae Jayasiri, E.B.G. Jones & K.D.
Hyde, Mycosphere 10(1): 1–186 (2019), Fig. 41
Holotype: THAILAND, Krabi Province, Mueang Krabi
District, on decaying pod carpel of Leucaena sp. (Fabaceae), 31 August 2017, S.C. Jayasiri, C 356 (MFLU
18-2159; MFLU 18-2160, isotype), ex-type living culture
MFLUCC 18-1544, KUMCC 18-0266.
Saprobic on dead twigs of Hevea brasiliensis. Sexual
morph Ascomata 130–195 lm high, 150–170 lm diam.,
visible as raised, aggregated, small, dark brown to black
dots on the host surface, immersed, erumpent through host
epidermis by minute papilla, scattered, solitary, sometimes
gregarious, globose to subglobose, occasionally irregular in
shape, uni-loculate, glabrous, ostiole central with minute
papilla. Peridium (7–)10–18 lm, composed of several
layers, of hyaline to dark brown pseudoparenchymatous
cells, inner layer comprising flattened, hyaline to brown
cells, arranged in a textura prismatica to textura angularis,
outer layer comprising thick, dark brown to black cells,
arranged in a textura angularis. Hamathecium composed of
dense, 1–3 lm wide, hyaline, septate, filamentous, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (35–)42–68(–75) 9 4–5(–6) lm
(
x = 53 9 5.5, n = 15), 8-spored, bitunicate, cylindrical,
with a short pedicel, apically rounded, with an indistinct
ocular chamber. Ascospores 7–10(– 15) 9 3–4.5(– 6.5)
lm (
x = 9.4 9 4.4, n = 40), overlapping 1-seriate, hyaline
when young and medium to dark brown when mature,
ellipsoidal to ovoid, with rounded ends, 1-septate, constricted at the septum, rough-walled, with two guttules, and
longitudinal striations. Asexual morph See Jayasiri et al.
(2019).
Culture characteristics: Colonies on PDA reaching
15–25 mm diam. after 3 weeks at 25–30 °C, circular,
medium dense, edge entire, velvety; from above, light grey
at the margin, white to light grey at the centre; from below,
white at the margin, light to dark brown at the centre.
Material examined: THAILAND, Phayao Province,
Muang District, Mae Ka, on dead twigs of Hevea
brasiliensis, 3 January 2017, C. Senwanna, RBPY028
(MFLU 17-1985), living culture, MFLUCC 17-0346.
Known hosts and distribution: On decaying pod carpel
of Leucaena sp. and Pterocarpus sp., Hevea brasiliensis
(Thailand) (Jayasiri et al. 2019; this study).
GenBank
numbers:
ITS = MH590694,
LSU =
MH590690, SSU = MH590692, TEF1-a = MH590688.
Notes: Neoroussoella leucaenae (MFLU 17-1985) is
found on Para rubber in Thailand for the first time. Phylogenetically our strain (MFLUCC 17-0346; Fig. 39)
clusters with the type strain of N. leucaenae (MFLUCC
18-1544) and strain MFLUCC 17-0927 with moderate
support (76% ML and 1.00 BYPP; Fig. 39). There is different only one and three nucleotide base positions in ITS
Fungal Diversity
Fig. 38 Pseudoastrosphaeriellopsis kaveriana (AMH-9912, holotype). a Ascomata semi-immersed and released ascospores on the
decaying wood of Avicennia marina. b, c Longitudinal sections of
ascoma. d Ostiole. e Section of peridium comprising hyaline to pale
brown cells of textura angularis. g Cellular pseudoparaphyses. h–k
Immature and mature asci. f, l–o Ascospores. p Germ tubes developed
from terminal ends of ascospore. Scale bars b = 100 lm, c,
d = 50 lm, e–p = 10 lm
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Fungal Diversity
Fig. 39 Maximum likelihood phylogenetic tree based on a combined
LSU, SSU, ITS, TEF1-a and RPB2 sequence dataset. The tree is
rooted to Torula herbarum (CBS 111855). Bootstrap support values
for ML (left) equal to or greater than 60% and the Bayesian posterior
123
probabilities (right), equal to or greater than 0.95 BYPP are indicated
above the nodes. Ex-type strains are in bold and the newly generated
sequences are indicated in blue
Fungal Diversity
Fig. 40 Neoroussoella heveae (MFLU 17-1983, holotype). a Conidiomata immersed in host substrate. b Section through the conidioma.
c Conidioma wall. d–f Conidiogenous cells. g, h Colony on MEA
(g = from above, h = from below). i Conidia. j Germinated conidium.
Scale bars a = 200 lm, b = 100 lm, c = 20 lm, d–j = 5 lm
and TEF1-a among N. leucaenae strains MFLUCC
18-1544, MFLUCC 17-0927 and MFLUCC 17-0346,
which confirms that they are conspecific.
We follow the latest treatment and updated accounts of
Roussoella in Liu et al. (2014), Tibpromma et al. (2017),
Hyde et al. (2018b) and Jiang et al. (2019).
Roussoella Sacc.
Roussoella elaeicola Konta & K.D. Hyde, sp. nov.
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Fungal Diversity
Index Fungorum number: IF555291; Facesoffungi
number: FoF04834, Fig. 42
Etymology: Name reflects the host genus Elaeis.
Holotype: MFLU 15-0022
Saprobic on dead petiole of Elaeis guineensis. Sexual
morph Ascomata 315–410 lm high, 325–350 lm diam.,
solitary, immersed in host tissue, erumpent through the
host surface with minute papilla, globose to subglobose,
uni-loculate, ostiole central with papilla, lacking periphyses. Peridium 25–70 lm wide, thin-walled, composed of
several layers of small, compressed, hyaline to light brown
pseudoparenchymatous cells of textura angularis, outer
layers fusing with the host. Hamathecium composed of
dense, 0.9–2.8 lm broad, filamentous, septate, trabeculate
pseudoparaphyses, embedded in mucilage. Asci 70–
140 9 6–9 lm (
x = 100 9 8 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, pedicellate, rounded apex
with an indistinct ocular chamber. Ascospores 10–15 9 3–
6 lm (
x = 12 9 4 lm, n = 30), slightly overlapping 1-seriate, hyaline to light brown when immature, ellipsoid with
obtuse ends, becoming dark brown at maturity, 1-septate,
with large guttules in each cell, constricted at the septum,
rough-walled with poroid ornamentation, surrounded by
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Ascospores germinated on
MEA within 24 h with germ tube was produced from both
cells. Colony on MEA, at first whitish, felty, azonate, with
fluffy margin, after incubation for 3–4 weeks appear circular, with fluffy, dense, pale brown mycelium in the
middle and dense mycelium in the outer ring on the surface
with smooth margin; from below, brown to dark brown in
the middle and pale brown to yellow at the margin hyphae,
septate, branched, and smooth.
Material examined: THAILAND, Chiang Rai Province,
on dead petiole of Elaeis guineensis (Arecaceae), 25
November 2014, S. Konta, HR02d (MFLU 15-0022,
holotype), ex-type living culture, MFLUCC 15-0276.
GenBank
numbers:
ITS = MH742329,
LSU =
MH742326 (MFLUCC 15-0276a); ITS = MH742330,
LSU = MH742327 (MFLUCC 15-0276b); LSU =
MH742328, SSU = MH742331 (MFLUCC 15-0276c).
Notes: Roussoella elaeicola (MFLU 15-0022) is introduced as a novel species and it was collected from oil palm
(Elaeis guineensis) in Chiang Rai, Thailand for the first
time. Multigene phylogenetic analyses show that R. elaeicola has a close relationship with R. euonymi Crous &
Akulov and Pseudoneoconiothyrium rosae Phukhams et al.
(Fig. 39) which were collected from Euonymus europaeus
L. (Celastraceae) in Ukraine and on dead aerial spines of
Rosa canina L. (Rosaceae) in Italy, respectively (Crous
et al. 2018; Wanasinghe et al. 2018). However, we could
not compare the morphological characteristics of our new
species with these two species as Roussoella euonymi and
123
Pseudoneoconiothyrium rosae are only known as asexual
morph species, while our new taxon is represented by the
sexual morph. One of the distinctive characters of Roussoella elaeicola is the distinctive trabeculate pseudoparaphyses (sensu Liew et al. 2000). A comparison of ITS
nucleotide base pairs indicates that Roussoella elaeicola
differs from R. euonymi and Pseudoneoconiothyrium rosae
in 37/513 bp (7.2%) and 38/514 bp (7.4%). Roussoella
elaeicola can be distinguished from other Roussoella species by its ascospores having poroid ornamentation, similar
to R. scabrispora (Liu et al. 2014). However, R. elaeicola
and R. scabrispora form a distinct lineage in
Roussoellaceae.
Sulcatisporaceae Kaz. Tanaka & K. Hiray.
Sulcatisporaceae was introduced by Tanaka et al. (2015)
to accommodate three genera: Magnicamarosporium Kaz.
Tanaka & K. Hiray., Neobambusicola Crous & M.J. Wingf.
and Sulcatispora Kaz. Tanaka & K. Hiray. and is typified
by Sulcatispora. The family is characterized by globose to
subglobose, ostiolate, papillate ascomata, a thick-walled
peridium with a poorly developed base, and trabeculate,
anastomosed pseudoparaphyses, bitunicate, clavate asci,
and hyaline, broadly fusiform, 1-septate ascospores with an
entire sheath. The asexual morphs are coelomycetous, with
ellipsoid to subglobose, hyaline to dark brown, muriform or
phragmosporous conidia (Crous et al. 2014b; Tanaka et al.
2015; Phukhamsakda et al. 2017). Five species from three
genera are accommodated in this family. We introduce a
new species, Neobambusicola brunnea (MFLU 18-1393)
based on a phylogenetic analysis of a combined LSU and
ITS sequence dataset (Fig. 43).
Neobambusicola Crous & M.J. Wingf.
Neobambusicola was introduced as a monotypic genus
by Crous et al. (2014b) and is typified by N. strelitziae
Crous & M.J. Wingf., isolated from leaves of Strelitzia
nicolai in South Africa. The genus was represented by its
asexual morph and is characterized by erumpent, globose,
dark brown, ostiolate conidiomata, a thin-walled of peridium, subcylindrical to ampulliform, phialidic conidiogenous cells and hyaline to olivaceous, fusoid-ellipsoid,
smooth-walled, 1-septate conidia with hyaline, smooth,
guttulate to granular, aseptate, subglobose to subcylindrical
microconidia (Crous et al. 2014b). Crous et al. (2014b)
treated the genus in Bambusicolaceae. Tanaka et al. (2015)
introduced a new family Sulcatisporaceae to accommodate
Neobambusicola, Magnicamarosporium and Sulcatispora.
Only Neobambusicola strelitziae was accommodated in
Neobambusicola, but we introduce a second species N.
brunnea which is represented by the sexual morph.
Neobambusicola brunnea Y. Chen & Norphanphoun, sp.
nov.
Fungal Diversity
Index Fungorum number: IF555293; Facesoffungi
number: FoF05708, Fig. 44
Etymology: The specific epithet ‘‘brunnea’’ refers to the
brownish ascospores.
Holotype: MFLU 18-1393
Saprobic on dead stem of herbage. Sexual morph Ascomata 80–205 lm high, 90–260 lm diam., semi-immersed, blackish, irregular in shape, scattered on surface of
stem, uni-loculate, glabrous, ostiolate, apapillate. Ostioles
35.5–68 lm diam., dark, circular and sunken at the apex of
ascoma. Peridium 30–40 lm thick, two layered, thickwalled, outer layer irregular, comprising dark brown cells
of textura angularis and inner layer slightly, irregular of
light brown cells. Hamathecium comprising 1–2 lm wide,
filamentous, branched or simple, septate, anastomosed,
cellular pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci 70–90 9 8–10 lm (
x = 83.6 9 9 lm,
n = 10), 4- or 8-spored, bitunicate, cylindrical to cylindricclavate, short pedicellate, apically rounded, with an ocular
chamber. Ascospores 11–18 9 5–6 lm (
x = 12.9 9 5.8 lm,
n = 20), 1-seriate, brown to dark brown, oblong to ellipsoidal,
with rounded ends, 1-septate, slightly constricted at the septum, smooth-walled, with small guttules. Asexual morph
Undetermined.
Material examined: CHINA, Guizhou Province, Qiandongnan Miao and Dong Autonomous Prefecture, Huangping District, on dead stem of herbage, 10 September 2017,
Y. Chen, QDN001 (MFLU 18-1393, holotype).
GenBank numbers: ITS = MH644792, LSU = MH644791.
Notes: DNA was extracted directly from the ascomata,
and a phylogenetic analysis of combined ITS and LSU
sequence dataset shows that Neobambusicola brunnea is
closely related to N. strelitziae with moderate support (87%
ML; Fig. 43). A comparison of ITS and LSU pair wise
shows that N. brunnea differs from N. strelitziae in 31 and
40 base positions, respectively. Therefore, N. brunnea is
introduced as a novel species and this is the first report of
the sexual morph of Neobambusicola.
Thyridariaceae Q. Tian & K.D. Hyde
Thyridariaceae was introduced by Hyde et al. (2013) to
accommodate the genus Thyridaria Sacc. and is typified by
T. broussonetiae (Sacc.) Traverso. The familial concept
was solitary or gregarious, immersed to erumpent, globose,
coriaceous ascomata, in valsoid configurations, with stromatic, pigmented, prosenchymatous tissues and ostioles
with a disc-like ostiolar canal. Asci are fissitunicate,
cylindrical to subclavate, pedicellate, with trabeculate
pseudoparaphyses and ascospores are pigmented, ellipsoidal to fusiform and verruculose, with transverse eusepta
or distosepta, and form coelomycetous Cyclothyrium
asexual morphs (Hyde et al. 2013; Jaklitsch and Voglmayr
2016). Jaklitsch and Voglmayr (2016) excluded
Cyclothyrium from Thyridariaceae and the genus was
tentative placed in Pleosporales, genera incertae sedis
(Wijayawardene et al. 2018a). Furthermore, they also
synonymized Roussoellaceae under Thyridariaceae and
accepted Neoroussoella, Thyridaria, Roussoella, Roussoellopsis and Parathyridaria Jaklitsch & Voglmayr.
However, Tibpromma et al. (2017) reinstated Roussoellaceae and this was followed by subsequent authors (Hyde
et al. 2018b; Wanasinghe et al. 2018; Wijayawardene et al.
2018a; Jayasiri et al. 2019; Jiang et al. 2019). Recently,
Wanasinghe et al. (2018) introduced three new genera in
Thyridariaceae viz. Cycasicola Wanas et al., Pseudoneoconiothyrium and Pararoussoella Wanas et al. However, in
the present study Pseudoneoconiothyrium and Pararoussoella cluster with other Roussoella species in Roussoellaceae. Devadatha et al. (2018) also introduced a new
genus Thyridariella Devadatha et al. in Thyridariaceae.
Thyridariella clustered with Cycasicola in our study
(Fig. 39). However, these two genera are represented by
different morphs and we therefore, treat them as different
genera until a link between these two genera is proven. In
this study, we introduce a novel genus Liua to accommodate a novel species Liua muriformis in Thyridariaceae
based on morphological distinctiveness and phylogenetic
support.
Liua Phookamsak & K.D. Hyde, gen. nov.
Index Fungorum number: IF556175; Facesoffungi
number: FoF05709
Etymology: In honour of Jian-Kui Liu, for his excellent
work on taxonomic revision of Dothideomycetes.
Saprobic on Lonicera maackii. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
pycnidial, scattered, solitary, immersed, slightly raised,
visible as small black dot on host surface, uni-loculate,
globose to subglobose, glabrous, dark brown to black,
ostioles central, apapillate, with pore-like opening. Conidiomata walls thin-walled, of equal thickness, composed of
3–5 cell layers, of flattened, brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis to
textura prismatica. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, phialidic,
discrete, determinate, ampulliform to cylindrical, unbranched, aseptate, occasionally 1–2-septate, hyaline, smooth,
arising from the inner cavity of the conidioma wall.
Conidia phragmosporous to muriform, dark brown, oblong
to ellipsoidal, or obovoid, (2–)3-transversely septate, with
(0–)3 longitudinal septa, slightly constricted at the septa,
smooth-walled.
Type species: Liua muriformis Phookamsak, H.B. Jiang
& K.D. Hyde
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Fungal Diversity
Fig. 41 Neoroussoella leucaenae (MFLU 17-1985). a–c Habit on substrate. d Section through ascoma. e Peridium. f Pseudoparapyses. g Asci.
h Ascospores. i Germinated ascospores. Scale bars b–d = 200 lm, g = 30 lm, e = 20 lm, f, h, i = 10 lm
Notes: In the NCBI BLASTn search of ITS sequences,
Liua muriformis most closely matches with leaf litter
ascomycete (strain its072) with 95% similarity and Cycasicola goaensis Wanas. et al. (MFLU 17-0581) with 93%
similarity. Multigene phylogenetic analyses based on a
combined LSU, SSU, ITS, TEF1-a and RPB2 sequence
dataset show that L. muriformis forms a sister clade with C.
goaensis and C. leucaenae Jayasiri et al. with high support
(100% ML and 1.00 BYPP) and clusters with other two
Thyridariella species in Thyridariaceae. Liua muriformis
123
differs from C. goaensis and C. leucaenae in having dark
brown, muriform or phragmosporous conidia, whereas, C.
goaensis and C. leucaenae have pale yellowish, aseptate
conidia (Wanasinghe et al. 2018; Jayasiri et al. 2019).
Fig. 42 Roussoella elaeicola (MFLU 15-0022, holotype). a, b Asco- c
mata on host substrate. c Section of ascoma. d Peridium. e Pseudoparaphyses. f–h Asci. i–l Immature ascospores. m, n Mature ascospores.
o, p Ascospores with poroid ornamentation. q Germinated ascospore.
r Culture characteristic on MEA from above and below. Scale bars a,
b = 1000 lm, c = 100 lm, d = 50 lm, e–h, q = 20 lm, i–p = 5 lm
Fungal Diversity
123
Fungal Diversity
Fig. 43 Phylogram generated from RAxML analysis based on a
combined LSU and ITS sequence dataset. Tree is rooted with
Camarosporium aloes (CPC 21572) and C. quaternatum (CBS
483.95). Bootstrap values C 50% are indicated at the nodes. The
new species is indicated in blue. Ex-type strains are indicated in bold
Therefore, we introduce Liua as a new genus to accommodate L. muriformis based on its morphological distinct
with Cycasicola.
Etymology: The specific epithet ‘‘muriformis’’ refers to
the holotype having muriform ascospores
Holotype: KUN-HKAS 102241
Saprobic on Lonicera maackii. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
80–150 lm high, 110–190 lm diam., pycnidial, scattered,
solitary, immersed, slightly raised, visible as small black
dot on host surface, uni-loculate, globose to subglobose,
glabrous, dark brown to black, ostioles central, apapillate,
Liua muriformis Phookamsak, H.B. Jiang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556176; Facesoffungi
number: FoF05710, Fig. 45
123
Fungal Diversity
with pore-like opening. Conidiomata walls 7–17 lm wide,
thin-walled, of equal thickness, composed of 3–5 cell
layers, of flattened, brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis to
textura prismatica. Conidiophores reduced to conidiogenous cells. Conidiogenous cells (3–)5–12.5(–
17.5) 9 (2–)5–10 lm l/w (
x = 8.2 9 5.4 lm, n = 30),
holoblastic, phialidic, discrete, determinate, ampulliform to
cylindrical, unbranched, aseptate, occasionally 1–2-septate,
hyaline, smooth, arising from the inner cavity of the
conidioma wall. Conidia (12.5–)13–15(–17) 9 7–9(–10)
lm l/w (
x = 14.4 9 8.2 lm, n = 50), phragmosporous to
muriform, dark brown, oblong to ellipsoidal, or obovoid,
(2–)3-transversely septate, with (0–)3 longitudinal septa,
slightly constricted at the septa, smooth-walled.
Culture characteristics: Colonies on PDA, reaching
30–32 mm diam. after 3 weeks at room temperature
(20–30 °C). Colony dense, circular, flattened to slightly
raised, surface rough, radially furrowed at the centre,
smooth at the margin, with edge entire, velvety; from
above, greyish green at the margin, pale yellowish to yellowish green at the centre; from below, dark green to
greenish grey; not producing pigmentation on agar
medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, on dead hanging
branches of Lonicera maackii, 20 April 2017, R.
Phookamsak, KIB032 (KUN-HKAS 102241, holotype),
ex-type living culture, KUMCC 18-0177.
GenBank
numbers:
ITS = MK433599,
LSU =
MK433598, SSU = MK433595, TEF1-a = MK426798,
RPB2 = MK426799
(KUMCC
18-0177);
ITS =
MK433600, LSU = MK433592 (KUN-HKAS 102241,
KIB0032F).
Dothideomycetes, orders incertae sedis
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf.
Asterinaceae was introduced by Hansford (1946) to
accommodate obligate biotrophic, or epiphytic fungi which
formed web-like, black colonies on the upper and lower
surfaces of leaves, or stems, with or without appressoria of
external mycelium, superficial, dimidiate ascomata with
radiating star-like openings, fissitunicate asci, cylindrical,
2–6-celled, yellowish to brown ascospores and forming
both coelomycetous and hyphomycetous asexual morphs
(Hyde et al. 2013; Hongsanan et al. 2014, 2016; Guatimosim et al. 2015). Twenty-one genera are accepted in the
family (Wijayawardene et al. 2018a). Updated molecular
phylogeny and their evolutionary relationships based on
molecular clock evidence were provided by Hongsanan
et al. (2016) and Liu et al. (2017a).
Lembosia Lév.
The genus Lembosia was described by Léveillé (1845),
based on the type species, L. tenella Lév. The genus is
characterized by having lirelliform or V–Y-shaped ascomata, opening by a longitudinal fissure, absence of
hypostroma, adhering to the host by superficial hyphae
with lateral appressoria (hyphopodia), bitunicate asci disposed as an upright palisade layer, and 2-celled brownish
ascospores (Hongsanan et al. 2014). More than 260 epithets
are listed under Lembosia in Index Fungorum (2019).
However, few species have molecular data. In this manuscript we introduce the new species Lembosia brigadeirensis (Fig. 46).
Lembosia brigadeirensis Firmino, A.R. Machado & O.L.
Pereira, sp. nov.
MycoBank number: MB822355; Facesoffungi number:
FoF04108, Fig. 47
Etymology: Name derived from the mountain range
where the fungus was collected, Serra do Brigadeiro.
Holotype: VIC 44208
Epiphyllous on Epidendrum sp. Sexual morph Colonies
4–6 mm diam., amphigenous, circular to irregular, single
to confluent, dark brown, black. Hyphae 4–5 lm diam.,
straight to flexuous, irregularly branched, brown, septate,
hyphal cells cylindrical, smooth. Appressoria 5.5–7.5
9 6–8 lm, few, entire to lobate, sessile, lateral, alternate to
unilateral, never opposed, globose, unicellular, straight,
brown, penetration peg central on the appressorial cell.
Ascomata 530–1180 9 140–230 lm, superficial, hysterothecia, lirelliform, V–Y-shaped, mostly linear, on the
top of mycelial mat, single to confluent, fringed at margins,
massed in the centre of the colony, opening by longitudinal fissures, dark brown to black, wall of textura radiate
to irregulata, cells isodiametric to cylindrical. Pseudoparaphyses up to 2.5 lm wide, cylindrical, filiform, septate, unbranched, hyaline. Asci 30–47.5 9 15–22 lm,
8-spored, bitunicate, fissitunicate, subclavate to cylindrical,
disposed as an upright palisade layer. Ascospores
17.5–19.5 9 5.5–8 lm, cylindrical to oblong-clavate, ends
rounded, straight or slightly arched, 1-septate, constricted
at the median septum, hyaline, becoming brown at maturity, smooth-walled. Asexual morph Undetermined.
Material examined: BRAZIL, Minas Gerais, Araponga,
Parque Estadual da Serra do Brigadeiro, on leaves of
Epidendrum sp. (Orchidaceae), 10 September 2014, A.L.
Firmino (VIC 44208, holotype).
GenBank numbers: ITS = MF667946, LSU = MF664531.
Notes: Lembosia brigadeirensis is a distinct species
when compared with many other Lembosia species reported on Orchidaceae (Léveillé 1845; Horne 1905; Sydow
1939; Silva and Pereira 2008; Hosagoudar et al. 2009;
Firmino and Pereira 2014). Lembosia brigadeirensis is
123
Fungal Diversity
Fig. 44 Neobambusicola brunnea (MFLU 18-1393, holotype). a–c
Appearance of ascomata semi-immersed in the host. d–f Vertical
section of ascoma. g–j Mature asci with ascospores. k Hamathecium.
123
l Apex of the ascus. m–q Ascospores. Scale bars d = 50 lm,
f = 20 lm, e, g–k = 10 lm, l–q = 5 lm
Fungal Diversity
Fig. 45 Liua muriformis (KUN-HKAS 102241, holotype). a Appearance of conidiomata on host. b Section through conidioma.
c Section through conidioma wall. d–g Conidiogenous cells. h–l
Conidia. m Germinated conidium. n, o Culture characteristics on
PDA (n = from above, o = from below). Scale bars a = 200 lm,
b = 50 lm, c = 20 lm, l, m = 10 lm, d–k = 5 lm
most similar to L. sertiferae Syd., which has epiphyllous
colonies, lobate appressoria, ellipsoid to oblong asci, and
fusiform to oblong ascospores (Firmino and Pereira 2014).
Lembosia bezerrae Firmino & O.L. Pereira has epiphyllous
colonies, smaller appressoria, saccate to ovoid asci, and
fusiform and smaller ascospores than L. brigadeirensis
(Firmino and Pereira 2014). Lembosia epidendri Meir.
Silva & O.L. Pereira has narrow hyphae, wider appressoria,
smaller hysterothecia, saccate to ovoid asci, branched
pseudoparaphyses, and larger and fusiform ascospores
(Silva and Pereira 2008; Firmino and Pereira 2014). Lembosia dendrochili Lév. differs from L. brigadeirensis in
having smaller hysterothecia, and larger asci (Léveillé
1845; Firmino and Pereira 2014). Lembosia rolfsii W.T.
Horne differs from L. brigadeirensis in the subcuticular
mycelium and conidia on superficial mycelium, and probably belonging to Maheshwaramyces (Hosagoudar et al.
2009; Firmino and Pereira 2014).
Based on LSU sequence data, Lembosia brigadeirensis
is 97% similarity to Prillieuxina baccharidincola (Rehm)
Petr. (GenBank no. KP143735), 95% similarity to Asterina
melastomatis Lév. (GenBank no. KP143739) and 94%
similarity to Alysidiella suttonii Cheew. & Crous (GenBank no. HM628777). Based on ITS sequence data, L.
brigadeirensis is 88% similarity to Blastacervulus eucalypti H.J. Swart (GenBank no. GQ303271), Alysidiella
suttonii (GenBank no. HM628774), and Heteroconium
kleinzeense Crous & Z.A. Pretorius (GenBank no.
EF110616). Phylogenetic analysis of LSU sequence dataset
(Fig. 46) shows that Lembosia brigadeirensis forms a sister
lineage with Prillieuxina baccharidincola (VIC42817)
with high support (100% BYPP). Lembosia differs from
Prillieuxina G. Arnaud and Asterina Lév. in having elliptical to cylindrical ascomata with longitudinal fissure.
Alysidiella Crous and Blastacervulus H.J. Swart are asexual morph genera belonging to the same family as Lembosia (Asterinaceae), but with no known connection to a
123
Fungal Diversity
sexual morph. Heteroconium Petr. is also an asexual
morph,
but
belonging
to
Herpotrichiellaceae
(Chaetothyriales).
Phylogenetic analyses based on a combined ITS, LSU,
TEF1-a and TUB2 sequence dataset are provided for the
genera Dothiorella (Fig. 48) and Sphaeropsis (Fig. 51).
Botryosphaeriales Schoch et al.
Dothiorella Sacc.
We follow the latest treatment and updated accounts of
Dothiorella in Yang et al. (2017). Updated phylogenetic
analysis was retrieved from Dissanayake et al. (2017) and
Wanasinghe et al. (2018).
Botryosphaeriaceae Theiss. & P. Syd.
Botryosphaeriaceae can be found as endophytes, saprobes and plant pathogens on various substrates worldwide
(Liu et al. 2012b; Slippers et al. 2013; Dissanayake et al.
2016; Phillips et al. 2018; Jayawardena et al. 2019). The
family comprises 28 genera and more than 190 species
(Phillips et al. 2018; Tibpromma et al. 2018; Wijayawardene et al. 2018a). We follow the latest treatments and
updated accounts in Dissanayake et al. (2016), Hyde et al.
(2016), Yang et al. (2017), Phillips et al. (2018), Wanasinghe et al. (2018) and Jayawardena et al. (2019).
Fig. 46 Phylogenetic tree was obtained by Bayesian inference
methods using the sequences of the LSU region. The posterior
probability values are indicated at the nodes. Strain numbers are
indicated after species names. New sequence is in blue bold. The
123
Dothiorella acericola Phookamsak, Tennakoon & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556178; Facesoffungi
number: FoF05711, Fig. 49
Etymology: The specific epithet ‘‘acericola’’ refers to
the host genus Acer, on which the holotype was collected.
Holotype: KUN-HKAS 102213
analyses included 30 strains including representative genera of
Asterinales sensu stricto and Asterinales sensu lato. The tree is
rooted with Venturia populina (CBS 256.38) and V. inaequalis (CBS
815.69)
Fungal Diversity
Fig. 47 Lembosia brigadeirensis (VIC 44208, holotype). a Colony
with open hysterothecia and surface mycelium. b Cross section of the
ascomata. c Globose to lobate unicelular appressoria. d Parallel
bitunicate asci. e Immature hyaline ascospores. f Brown and smooth
ascospores. Scale bars a = 200 lm, b–f = 10 lm
Saprobic on dried twigs of Acer palmatum. Sexual
morph Undetermined. Asexual morph Coelomycetous.
Conidiomata 220–360 lm high, 190–310 lm diam., pycnidial, aggregated, clustered, semi-immersed to erumpent,
dark brown to black, globose to subglobose, uni- to biloculate, ostiole central, with minute papilla. Conidiomata
walls 15–40 lm wide, composed of several layers of broad
to flattened, dark brown to black, pseudoparenchymatous
cells of textura angularis to textura prismatica, with flattened, hyaline cells towards the inner layers. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells
(7–)9–15(–24) 9 3–6(–7) lm l/w (
x = 13.2 9 4.5 lm,
n = 30), holoblastic, phialidic, discrete, determinate,
ampulliform to cylindrical, unbranched, aseptate, hyaline,
smooth, arising from the inner cavity of the conidioma wall.
Conidia 17–22(–23) 9 7–10(–13) lm l/w (
x = 20.8 9
9.2 lm, n = 50), dark brown, oblong to ellipsoidal, 1-septate, slightly constricted at the septum, smooth-walled.
Culture characteristics: Colonies on PDA reaching
70–73 mm diam. after 1 week at 20–30 °C; initially
medium sparse to dense, circular, or slightly irregular in
shape, surface smooth, with edge entire to lobate; from
above, initially white, becoming white-grey to grey; from
below, grey to pale yellowish; not producing pigmentation
on agar medium.
Material examined: CHINA, Yunnan Province, Kunming, Panlong, Ciba, on dead hanging twigs of Acer palmatum Thunb. (Sapindaceae), 28 November 2015, R.
Phookamsak, COE009 (KUN-HKAS 102213, holotype),
ex-type living culture KUMCC 18-0137.
GenBank
numbers:
ITS = MK359449,
TEF1a = MK361182.
Notes: In the NCBI BLASTn search of ITS and TEF1-a
sequences, Dothiorella acericola is most similar to D.
viticola A.J.L. Phillips & J. Luque, with 99% and 98%
similarities, respectively. Phylogenetic analyses of a combined ITS and TEF1-a sequence dataset show that D.
acericola is sister to Spencermartinsia alpina Y. Zhang ter
& Ming Zhang and distinct from D. viticola (Fig. 48).
Dothiorella acericola, Spencermartinsia alpina and S.
yunnana Zhang ter & Ming Zhang were collected from
Yunnan, China, but they are phylogenetically distant
(Zhang et al. 2016). Dothiorella acericola has the same
size range of conidia as Spencermartinsia alpina and S.
yunnana (Zhang et al. 2016). A comparison of TEF1-a
nucleotide bases shows that D. acericola differs from
Spencermartinsia alpina and S. yunnana in 11/225 bp
(4.9%) and 13/225 bp (5.8%), respectively. Therefore, we
introduce a new species, D. acericola in this study based on
the guidelines of Jeewon and Hyde (2016). Yang et al.
(2017) treated Spencermartinsia as a synonym of
123
Fungal Diversity
Dothiorella. Spencermartinsia alpina and S. yunnana
should perhaps be transferred to the genus Dothiorella.
Dothiorella sarmentorum (Fr.) A.J.L. Phillips, A. Alves &
J. Luque, Mycologia 97(2): 522 (2005)
Facesoffungi number: FoF04836, Fig. 50
Saprobic on a wide range of hosts. Sexual morph
Undetermined. Asexual morph Conidiomata 300–440 lm
high, 215–300 lm diam., stromatic, solitary or scattered in
small groups, immersed, uni-loculate, individual or
aggregated, black, with globose to subglobose, ostiole.
Conidiomata walls comprising several layers; outer layers
thick-walled, dark brown cells of textura angularis; inner
layers of thin-walled, lightly pigmented or hyaline cells.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells lining inner the conidioma cavity, holoblastic,
hyaline, subcylindrical, proliferating at the same level
giving rise to periclinal thickenings. Conidia 18–23 9 8–
10 lm (
x = 21 9 9.4 lm, n = 30), ovoid, with a broadly
rounded apex and truncate base, initially hyaline to lightly
pigmented and aseptate, becoming dark brown, 1-septate,
slightly constricted at the septum, smooth-walled.
Material examined: RUSSIA, Rostov region, Shakhty
City, Alexandrovsky Park, on dead twigs of Platycladus
orientalis (L.) Franco (Cupressaceae), 14 March 2016, T.S.
Bulgakov (MFLU 16-1627).
Known hosts: Acer platanoides L., Aesculus hippocastanum L., Armeniaca vulgaris Lam., Cedrus atlantica
(Endl.) Manetti ex Carrière, Chamaecyparis lawsoniana
(A. Murray) Parl., Cornus sanguinea L., Coronilla emerus
L., Crataegus sp., Cryptomeria japonica (L.f.) D.Don,
Cupressus lusitanica Mill., Eriobotrya japonica (Thunb.)
Lindl., Euonymus europaeus L., Forsythia europaea Degen
& Bald., Malus pumila Miller, Menispermum canadense
L., Paliurus spina-christi Mill., Persica vulgaris Mill.,
Pistacia spp., Populus nigra L., Prunus spp., Pyrus communis L., Quercus spp., Salix sp., Thuja spp., Ulmus spp.,
Vitis spp. (Farr and Rossman 2018).
Known distribution: Iran, Italy, Netherlands, New
Zealand, Norway, Poland, Portugal, Serbia, Spain, Sweden,
UK (Great Britain), Ukraine, the USA (California, Florida,
Oregon, Washington) (Farr and Rossman 2018).
GenBank
numbers:
ITS = MH571673,
TEF1a = MH628155.
Notes: Dothiorella sarmentorum was introduced by
Phillips et al. (2005). This species is a cosmopolitan distribution including many economical important trees
(Phillips et al. 2005, 2013; Dissanayake et al. 2017). We
isolated D. sarmentorum from Platycladus orientalis (Cupressaceae) for the first time (Farr and Rossman 2018).
The morphological characters such as conidia shape, size
and colour are similar to the type as described. However,
we could not obtain a living culture from the isolated
123
spores. Therefore, the morphology of the species is based
only on characters on the host. Phylogenetic analyses of a
combined ITS and TEF1-a sequence dataset (Fig. 48)
show that our isolate (MFLU 16-1627) clusters with the
type strain of D. sarmentorum (IMI63581b) and strain CBS
115038.
Sphaeropsis Sacc.
Sphaeropsis was introduced by Saccardo (1880b) to
accommodate diplodia-like taxa and is typified by S. visci
(Alb. & Schwein.) Sacc. with Phaeobotryosphaeria sexual
morph (Phillips et al. 2008, 2013; Dissanayake et al. 2016;
Wijayawardene et al. 2017a). Sphaeropsis has a cosmopolitan distribution on various hosts (Farr and Rossman
2018). The genus is characterized by pseudothecial, brown
to black, uni-loculate ascomata, thick endotunica, bitunicate asci, with cellular pseudoparaphyses, brown, aseptate
ascomata and asexual morph forms with stromatic conidiomata, with paraphyses and oval, oblong or clavate,
aseptate conidia (Phillips et al. 2013). More than 600
species are listed under Sphaeropsis in Index Fungorum
(2019). However, Phillips et al. (2013) re-circumscribed
the genus and only five species were accepted based on
morphological characteristics of the sexual and asexual
morph connections and phylogenetic evidence (Phillips
et al. 2008, 2013; Doilom et al. 2015, 2017; Dissanayake
et al. 2016; Wijayawardene et al. 2017a). In this study, we
report a new host record of S. eucalypticola from Bauhinia
purpurea (L.) Benth. in Thailand.
Sphaeropsis eucalypticola A.J.L. Phillips, in Phillips et al.,
Stud Mycol 76: 158 (2013)
Facesoffungi number: FoF00169, Fig. 52
Holotype: THAILAND, Chiang Rai Province, Muang
District, on dead twig of Eucalyptus sp., 8 August 2011, M.
Doilom, MFLU 12-0753.
Saprobic on dead twigs. Sexual morph Ascostromata
250–350 lm high, 170–250 lm diam. (ascostromata with
papilla, not including subiculum or hypostroma), black,
convex on host tissue, appearing through cracks in bark,
scattered or clustered in small to large groups on a
subiculum or hypostroma, 185–260 lm high at the base,
aggregated, initially immersed, becoming erumpent, when
cut horizontally locules visible as white contents and dark
ascospore dots, uni-loculate or multi-loculate, globose to
subglobose or flask-shaped. Papilla 60–95 lm long, 65–
85 lm diam., ostiole with periphyses. Peridium 35–80 lm
wide, thick-walled, composed of several layers of dark
brown to black, coriaceous cells of textura angularis.
Hamathecium comprising 2.5–4 lm wide, hyphae-like,
hyaline, numerous, septate pseudoparaphyses, constricted
at the septa. Asci 102–175 9 22–32 lm (
x = 130 9 27 lm,
n = 15), 8-spored, bitunicate, fissitunicate, cylindric-clavate
or clavate, with a short or long pedicel, apically rounded
Fungal Diversity
Fig. 48 Phylogenetic tree generated from maximum likelihood
(RAxML) based on a combined ITS and TEF1-a. Maximum
likelihood bootstrap value C 70% and Bayesian posterior
probabilities C 0.95 BYPP are given at the nodes. The ex-type
strains are in bold. The newly generated sequences are in blue. The
tree is rooted with Neofusicoccum parvum (CMW9081)
with an ocular chamber. Ascospores 27–33 9 11–14 lm
(
x = 30 9 13 lm, n = 20), overlapping 2-seriate, hyaline
when young, becoming pale brown or reddish brown when
mature, ellipsoidal to ovoid, aseptate, sometime 2-septate,
broader in the centre, with an apiculus at both ends, thickwalled, echinulate. Asexual morph Undetermined.
123
Fungal Diversity
Fig. 49 Dothiorella acericola (KUN-HKAS 102213, holotype).
a Appearance of conidiomata on host. b, c Section through conidiomata. d Section through conidioma wall. e–g Conidiogenous cells
(g = stained with congo red). h–j Conidia. k Germinated conidium.
l Culture characteristics on PDA from above and below. Scale bars b,
c = 200 lm, d = 50 lm, h = 20 lm, e–g, i–k = 10 lm
Culture characteristics: Ascospores germinating on
PDA after 5–10 h. Germ tubes produced from germ pore of
ascospores. Colonies on PDA, reaching the edge of the
Petri dish after 5 days, flat or effuse, undulate, initially
white, after 3 days becoming brownish grey to olive.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Tha Sud Subdistrict, Mae Fah Luang
University campus grounds, on dead twigs of Bauhinia
purpurea L. (Leguminosae), 14 March 2012, M. Doilom,
MKBB031 (MFLU 18-1857), living culture, MFLUCC
12-0171.
Known hosts and distribution: Eucalyptus sp., Bauhinia
purpurea, Tectona grandis L.f. (Thailand) (Liu et al.
2012b; Phillips et al. 2013; Doilom et al. 2017).
GenBank
numbers:
ITS = MK108956,
TEF1a = MK108958, TUB2 = MK108957.
Notes: A new isolate of Sphaeropsis eucalypticola
(MFLUCC 12–0171) was collected from dead twigs of
Bauhinia purpurea in Thailand. This new isolate shares a
close phylogenetic affinity to the type of S. eucalypticola
(100% ML, 100% MP and 1.00 BYPP) in our combined
phylogeny of ITS, TEF1-a and TUB2 sequence data
(Fig. 51). Sphaeropsis eucalypticola has been reported
from Eucalyptus sp. and Tectona grandis in Thailand (Liu
et al. 2012b; Doilom et al. 2017), but it has not been previously reported from Bauhinia purpurea (Fig. 52).
123
Microthyriales G. Arnaud
Microthyriales, genera incertae sedis
Parazalerion Madrid, Gené & Cano, gen. nov.
MycoBank number: MB824747; Facesoffungi number:
FoF04480
Etymology: The name reflects the superficial morphological similarity between this genus and species of Zalerion sensu lato
Saprobic in soil. Sexual morph Undetermined. Asexual
morph Hyphomycetous. Vegetative hyphae septate, branched, subhyaline to light olivaceous. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells subcylindrical to narrowly clavate, smooth-walled, light olivaceous. Conidia produced terminally or laterally on
Fungal Diversity
Fig. 50 Dothiorella sarmentorum (MFLU 16-1627). a, b Appearance of conidiomata on host surface. c Section through conidioma wall. d, e
Conidiogenous cells. f–h Conidia. Scale bars c, d = 20 lm, e–h = 10 lm
undifferentiated hyphae, or on short conidiophores. Conidial filament curved, sinuous or irregularly coiled, multiseptate, light olivaceous brown to mid brown, slightly to
strongly constricted at the septa; groups of conidia often
appear compactly intertwined, forming irregularly shaped
masses of cells.
Type species: Parazalerion indica Madrid, Gené &
Cano
Notes: Parazalerion is introduced as a novel monotypic
conidial genus of Ascomycota. Morphologically, it
resembles the marine genus Zalerion R.T. Moore &
Meyers (Sordariomycetes, genera incertae sedis) in the
production of irregularly coiled, dematiaceous, multi-septate conidia which often form knots of cells (Ellis 1976;
Goos 1985; Campbell et al. 2005). The new genus, however, was found in a terrestrial habitat and is phylogenetically related to Spirosphaera minuta Hennebert (Fig. 53),
which belongs in Dothideomycetes, relatively close to
Microthyriaceae (Voglmayr et al. 2011). Furthermore,
knots of cells originate from a single conidial filament in
Zalerion, whereas in Parazalerion they originate from the
intertwining of groups of conidia (Fig. 54). This particular
development of knots of cells also distinguish Parazalerion
from other similar genera, including Cirrenalia Meyers &
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Fungal Diversity
R.T. Moore, Cumulospora I. Schmidt, Glomerulispora
Abdel-Wahab & Nagah., Halazoon Abdel-Aziz et al., Hiogispora Abdel-Wahab & Nagah., Hydea K.L. Pang &
E.B.G. Jones, Matsusporium E.B.G. Jones & K.L. Pang,
Moheitospora Abdel-Wahab et al., Moleospora AbdelWahab et al. and Moromyces Abdel-Wahab et al. (Ellis
1971; Abdel-Wahab et al. 2010). The LSU-based phylogenetic tree demonstrates that Parazalerion is phylogenetically clearly distinct from those hyphomycete genera
(Fig. 53).
The closest match in the BLASTn search with ITS ?
LSU sequences of Parazalerion indica strain CBS 125443
is Spirosphaera minuta strain CBS 476.66 (GenBank no.
HQ696659, 92% similarity, 92% query coverage). In our
study, Spirosphaera minuta and Parazalerion indica
appeared as sister taxa in the LSU-based phylogenetic tree
(Fig. 53) and this relationship received 97% bootstrap
support. Since the type species of Spirosphaera, S. floriformis Beverw., is a phylogenetically distant species
belonging to Helotiales (Voglmayr et al. 2011), Spirosphaera minuta apparently needs to be reallocated to a
different genus. Its definitive phylogenetic position
deserves further study, but the fungus is also clearly different from Parazalerion. It occurs on submerged plant
material and produces more or less globose conidia formed
by a complexly coiled hyaline, branching filament (Hennebert 1968). Spirosphaera is a polyphyletic genus that
needs to be revised. Another member of this genus, S.
cupreorufescens Voglmayr is a member of Pleosporales
(Voglmayr et al. 2011) and also needs to be segregated
from Spirosphaera sensu stricto.
Parazalerion indica Madrid, Gené & Cano, sp. nov.
MycoBank number: MB824748; Facesoffungi number:
FoF04481, Fig. 54
Etymology: Refers to the country where this fungus was
collected, India.
Holotype: IMI 397928
Saprobic in soil. Sexual morph Undetermined. Asexual
morph Hyphomycetous. Vegetative hyphae 1–1.5 lm
wide, septate, branched, subhyaline to light olivaceous,
smooth- and thin-walled. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2.5–5 9 1.5–2 lm,
subcylindrical to narrowly clavate, smooth-walled, light
olivaceous. Conidia produced terminally or laterally on
undifferentiated hyphae, or on short conidiophores. Conidial filament curved, sinuous or irregularly coiled, with
1–3(–6) usually thick and dark septa, light olivaceous
brown to mid brown, thick-walled, smooth to verruculose,
slightly to strongly constricted at the septa, 8–19(–26) lm
long, 4–5.5 lm wide at the widest part, with an obtuse
apex; groups of conidia often appearing tightly intertwined,
123
forming compact, irregularly shaped, light olivaceous
brown to dark brown, 8–23 lm wide masses of cells.
Culture characteristics: Colonies reaching 21 mm.
diam. on OA and 25 mm. diam. on PCA and PDA after
14 days at 25 °C. On OA light grey and floccose at the
centre, light brown to dark brown and almost glabrous
towards the periphery, with a fimbriate margin; reverse
dark brown at the centre, light brown towards the periphery. On PCA white to cream, funiculose at the centre,
almost glabrous towards the periphery, with a fimbriate
margin; reverse concolorous with obverse. On PDA velvety to floccose, cream to dark brown, with a fimbriate
margin; reverse concolorous with obverse. No diffusible
pigments observed in any of the tested media.
Material examined: INDIA, Delhi, isolated from soil, 2
February 1997, H.C. Gugnani (IMI 397928, holotype), extype living cultures CBS 125443, FMR 9690.
GenBank numbers: ITS ? LSU = MH100803.
Muyocopronales Mapook et al.
Muyocopronaceae K.D. Hyde
Muyocopronaceae was invalidly introduced by Luttrell
(1951) [as ‘Myiocopronaceae’], and is typified by Muyocopron Speg. with M. corrientinum Speg. as the type
species. The genus was introduced to accommodate epiphytic fungi, characterized by black, superficial, dimidiatescutate, subcarbonaceous, ostiolate ascomata, without
mycelium and bitunicate, ovoid to obclavate asci, containing subglobose to ellipsoidal, hyaline ascospores
(Mapook et al. 2016; Tibpromma et al. 2016). Muyocopron
has long historical discussion of its taxonomic placement
(Saccardo 1883a; von Arx and Müller 1954, 1975; Eriksson and Hawksworth 1993; Lumbsch and Huhndorf 2007;
2010), until Hyde et al. (2013) re-defined the family
Muyocopronaceae to accommodate Muyocopron. Mapook
et al. (2016) introduced the order Muyocopronales to
accommodate this family based on molecular data coupled
with morphological characteristics. Only Muyocopron is
accommodated in the family with more than 60 epithets
listed (Hyde et al. 2013; Mapook et al. 2016; Tibpromma
et al. 2016; Wijayawardene et al. 2018a; Index Fungorum
2019). In this study, a new host and geographical records of
Muyocopron lithocarpi Mapook et al. is reported (Figs. 55,
56).
Muyocopron Speg.
We follow the latest treatment and updated accounts of
Muyocopron in Mapook et al. (2016) and Tibpromma et al.
(2016).
Muyocopron lithocarpi Mapook, Boonmee & K.D. Hyde,
in Mapook et al., Phytotaxa 265(3): 235 (2016), Fig. 56
Holotype: THAILAND, Chiang Rai Province, on fallen
leaves of Lithocarpus lucidus (Fagaceae), 30 September
Fungal Diversity
Fig. 51 Phylogram generated
from parsimonious tree based
on combined ITS, TEF1-a and
TUB2 sequence dataset. The
tree is rooted to Barriopsis
tectonae (MFLUCC 12-0381)
and B. fusca (CBS 174.26).
Maximum parsimony and
maximum likelihood bootstrap
values C 70% and Bayesian
posterior probabilities C 0.95
are given at the nodes. The extype strains are in bold. The
newly generated sequence is in
blue bold
2014, A. Mapook (MFLU 15-1133), ex-type living culture
MFLUCC 14-1106.
Saprobic on dead stems of herbaceous plant. Sexual
morph Ascostromata dry, black, circular, raised to
umbonate on the host surface, without a subiculum, easily
removed from the host, clustered, gregarious, or in groups
of 2–3 locules, ostiolate. Ascomata 55–110 lm high, 175–
380 lm diam., clustered, gregarious or in groups, superficial, black, with a central irregular ostiole. Peridium 10–
27 lm wide, slightly thick-walled of unequal thickness,
poorly developed at the base, slightly thick at the sides
towards the apex, comprising two types of cell layers; outer
layer composed of black carbonaceous, brittle cells, inner
layer composed of hyaline to brown, pseudoparenchymatous cells of textura angularis to textura prismatica. Hamathecium comprising numerous, 2–3 lm wide, filiform,
septate, anastomosed pseudoparaphyses. Asci (41–)47–
78(–85) 9 (18–)21–28(–29) lm (
x = 59.6 9 24.5 lm,
n = 45), 8-spored, bitunicate, ovoid to obclavate, or
ampulliform, short pedicellate, apically rounded, apex
thick with small ocular chamber. Ascospores (12–)14–
19(–20) 9 (7–)8–10(–12) lm (
x = 17.2 9 9.8 lm, n = 60),
overlapping 1–3-seriate, hyaline, subglobose to obovoid,
with obtuse ends, 1-celled, rough-walled with small granules, and 1–3 large guttules. Asexual morph Undetermined.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Jinghong, Nabanhe, on dead stems of
herbaceous plant, 21 November 2015, R. Phookamsak,
XB016 (KUN-HKAS 102243).
Known hosts and distribution: Lithocarpus lucidus,
Peltophorum sp. (Thailand); Cercis chinensis (Guizhou,
China) (Mapook et al. 2016; Jayasiri et al. 2019).
GenBank numbers: LSU = MK447738, SSU = MK447740.
Notes: In molecular phylogenetic analysis our isolate
clusters with Muyocopron lithocarpi Mapook et al.
(Fig. 55). The morphology of our isolate is similar to M.
lithocarpi described by Mapook et al. (2016), although our
isolate has larger asci than in the original description
123
Fungal Diversity
123
Fungal Diversity
b Fig. 52 Sphaeropsis eucalypticola (MFLU 18-1857). a Ascostromata
on dead twig of Bauhinia purpurea. b Ascostromata cut through
horizontally showing the white contents with dark spots corresponding to asci with ascospores. c–e Vertical section of ascostromata.
f Immature and mature asci with immature and mature ascospores. g–
i Immature asci. j, k Asci with ascospores. l, m Immature ascospores.
n, o Ascospores. p Germinated ascospore. q, r Colony on PDA after
2 months (q = above view, r = below view). Notes h, i, l stained in
lactophenol cotton blue. Scale bars c, e, f = 100 lm. d = 50 lm. g, l–
p = 10 lm. h–k = 20 lm
(41–85 9 18–29 lm versus 45–65 9 15–28 lm) (Mapook
et al. 2016). The species was collected from herbaceous
plant in Yunnan, China for the first time.
Tubeufiales Boonmee & K.D. Hyde
Tubeufiaceae M.E. Barr
We follow the latest treatment and updated accounts of
Tubeufiaceae in Brahmanage et al. (2017), Chaiwan et al.
(2017), Liu et al. (2018), Lu et al. (2018a, b), Phookamsak
et al. (2018), Tibpromma et al. (2018) and Jayasiri et al.
(2019). There are 27 genera accommodated in this family
based on molecular data coupled with morphological
characteristics (Lu et al. 2018b; Tibpromma et al. 2018;
Wijayawardene et al. 2018a; Jayasiri et al. 2019). In this
study, a new species Pseudohelicomyces menglunicus is
introduced from a rotten seed coat in Yunnan, China
(Fig. 57).
Pseudohelicomyces Y.Z. Lu et al.
We follow the latest treatment and updated accounts of
Pseudohelicomyces in Lu et al. (2018b) and Jayasiri et al.
(2019).
Pseudohelicomyces menglunicus J.F. Li, Phookamsak &
K.D. Hyde, sp. nov.
Index Fungorum number: IF555770; Facesoffungi
number: FoF05763, Fig. 58
Etymology: The specific epithet ‘‘menglunicus’’ refers to
the Menglun Town, Xishuangbanna, Yunnan, China, where
the holotype was collected.
Holotype: KUN-HKAS 85795
Saprobic on seed coat. Sexual morph Undetermined.
Asexual morph Mycelium immersed on the substrate,
composed of septate, branched, smooth, thin-walled, subhyaline to dark brown hyphae. Conidiophores (106–)130–
220 9 9–13(–13.5) lm, macronematous or micronematous, paler towards the apex, hyaline to light brown, thinwalled, smooth, septate, branched, straight or flexuous,
cylindrical, tapering towards the apex. Conidiogenous cells
(20–) 23.5–32(–36) 9 (4–)4.5–5.5(–6) lm (
x = 4.5 9 5.3 lm,
n = 10), holoblastic, integrated, intercalary and sometimes terminal, determinate or sympodial, occasionally small and
discrete, denticulate, denticles cylindrical, often narrow,
hyaline, smooth. Conidia (18–)20–33(–34) 9 (21.5–)25–
30(–40) lm (
x = 22.5 9 27.2 lm, n = 20), pleurogenous or
acropleurogenous, solitary, simple, subhyaline to paler yellowish brown, septate, slightly constricted at the septa,
planate to cochleate, smooth, thin-walled, hygroscopic.
Culture characteristics: Conidia germinating on PDA
within 14 h and germ tubes produced from all cells.
Colonies growing on PDA, hairy, brown to dark brown,
reaching 5 mm in 15 days at 23 °C, mycelium partly
superficial, partly immersed, slightly effuse, radially striate, with irregular edge, subhyaline to dark brown; conidia
sporulating within 15 days on PDA.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, on unidentified seed, 15 August 2014, J.F.
Li, H-26 (KUN-HKAS 85795, holotype), ex-type living
culture, MFLUCC14-0689.
GenBank numbers: ITS = MK335914, SSU = MK335915,
TEF1-a = MK335916.
Notes: Pseudohelicomyces menglunicus resembles species of Helicosporium Nees and Neohelicosporium Y.Z. Lu
et al. in morphological characters, but is obviously unique
in conidiophores and conidia. Pseudohelicomyces
menglunicus differs from Helicosporium and Neohelicosporium species in having flexuose, branched and
hyphae-like conidiophores, and smooth-walled conidia. In
the phylogenetic analysis (Fig. 57), P. menglunicus forms a
separated lineage, sister to P. aquaticus Y.Z. Lu et al. with
moderate support (76% ML and 0.97 BYPP). Jayasiri et al.
(2019) introduced a new sexual morph species, P. quercus
Jayasiri et al. on fruit of Quercus sp. from Thailand. The
species also formed a clade with P. aquaticus and P.
menglunicus (pre-analysis, data not shown). A comparison
of ITS and TEF1-a nucleotide bases shows that P.
menglunicus differs from P. quercus in 37/569 bp (6.5%)
of ITS and 28/912 bp (3.1%) of TEF1-a. Hence, P.
menglunicus is introduced as a new asexual morph species.
Class Eurotiomycetes O.E. Erikss. & Winka
We follow the latest treatment and updated account of
Eurotiomycetes in Gueidan et al. (2014) and Geiser et al.
(2015). The outline and notes of the genera in Eurotiomycetes was provided by Wijayawardene et al.
(2017a, 2018a).
Subclass Chaetothyriomycetidae Doweld
Chaetothyriales M.E. Barr
Cyphellophoraceae Réblová & Unter.
Cyphellophoraceae was introduced by Réblová et al.
(2013) to accommodate a monotypic genus Cyphellophora
G.A. de Vries and is typified by C. laciniata G.A. de Vries
123
Fungal Diversity
Fig. 53 Maximum likelihood tree based on partial LSU region,
showing the phylogenetic relationships of Parazalerion and other
ascomycetes including morphologically similar hyphomycetous genera. Bootstrap support values [ 70% are shown near the internodes.
The tree is rooted to Fibulochlamys chilensis (Agaricales). The
original isolate numbers and GenBank accession numbers of LSU
sequences are noted after the species names. Ex-type strains are
indicated in bold
which was isolated from human skin in Switzerland. Taxa
have been reported as pathogens or endophytes on plants,
or as soil borne, as well as infection on humans and animals (i.e., nails and skin) (Réblová et al. 2013; Feng et al.
2014; Gao et al. 2015; Yang et al. 2018a). Twenty-six
epithets are listed in Index Fungorum (2019) with 23
possible species are accepted in this genus.
Cyphellophora G.A. de Vries
Most Cyphellophora species have been reported in their
asexual morph. However, Yang et al. (2018a) reported the
sexual morph of Cyphellophora on living leaves of Alnus
nepalensis D. Don (Betulaceae) from China for the first
time. We introduce a new sexual species, C. filicis, which
was collected from dead fronds of a fern in Thailand. The
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Fungal Diversity
Fig. 54 Parazalerion indica (CBS 125443, holotype). a Colony on PDA after 14 days at 25 °C. b, c Conidiogenous cells and conidia. d, e
Conidia and knots of cells. Scale bars a–e = 10 lm
new species is introduced based on its morphological distinctiveness and phylogenetic support (Fig. 59).
Cyphellophora filicis Hongsanan, Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556179; Facesoffungi
number: FoF05712, Fig. 60
Etymology: The specific epithet ‘‘filicis’’ (Latin: fern)
refers to the host, from which the holotype was collected.
Holotype: KUN-KHAS 102220
Saprobic on dead fronds of a fern, without dark superficial hyphae, appearing as black dots on host surface.
Sexual morph Ascomata 40–60 lm high, 55–75 lm
diam., immersed in host tissue, becoming erumpent, solitary, subglobose to globose, dark brown, uni-loculate,
glabrous, ostiolate, with minute papilla. Peridium 3–10 lm
wide, thin-walled, composed of 2–5 layers of flattened,
brown pseudoparenchymatous cells of textura angularis.
Asci 30–50 9 (10–)12–15(–19) lm (
x = 41.5 9 13.6 lm,
n = 30), 8-spored, bitunicate, ovoid to ampulliform, short
pedicellate, with an ocular chamber. Ascospores (15–)17–
20(–22) 9 (3–)4–5(–6) lm (
x = 18.6 9 4.7 lm, n = 30),
overlapping 2–3-seriate, hyaline, ellipsoidal to fusiform,
inconspicuously 3-septate, not constricted at the septa,
narrowly round at the ends, sometimes curved at the
middle, with a guttule in each cell. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA reaching
34–36 mm diam. after 3 weeks at 20–25 °C, colonies circular, dense, flat, slightly raised, surface dull, with edge
entire, woolly to velvety; from above, dark grey at the
margin, white-grey at the centre, separating from the
margin by convex, concentric ring; from below dark
greenish at the margin, with green yellowish at the centre;
not producing pigmentation in agar.
Material examined: THAILAND, Chiang Rai Province,
Doi Pui, on dead fronds of a fern, 2 February 2017, R.
Phookamsak, DP002 (KUN-KHAS 102220, holotype), extype living culture, KUMCC 18-0144 (DP002A),
KUMCC18-0145 (DP002B).
GenBank
numbers:
ITS = MK404056,
LSU =
MK404052, SSU = MK404054 (KUMCC 18-0144);
ITS = MK404057, LSU = MK404053, SSU = MK404055
(KUMCC18-0145).
Notes: The phylogenetic tree (Fig. 59) shows that
Cyphellophora filicis is closely related to a fungus in
Chaetothyriales (T222) that is associated with ants nest and
their runway galleries (Nepel et al. 2014). It is also related
to C. fusarioides (B. Sutton & C.K. Campb.) Decock, C.
laciniata G.A. de Vries, C. suttonii (Ajello et al.) Decock
and C. vermispora A. Walz & de Hoog, but as a distinct
new species in Cyphellophora (Fig. 59). We could not
compare morphological characters of our new species and
those Cyphellophora species due to the fact that C. filicis is
123
Fungal Diversity
Fig. 55 RAxML tree based on a combined LSU and SSU sequence
dataset. Bootstrap support values for ML equal to or greater than 60%
and Bayesian posterior probabilities equal to or greater than 0.95
123
BYPP are defined as ML/BYPP above the nodes. The tree is rooted to
Lichenothelia convexa (L1607). Newly generated sequence is in blue
and ex-type strains are in bold
Fungal Diversity
Fig. 56 Muyocopron lithocarpi (KUN-HKAS 102243). a Ascomata
on host substrate. b Squash mounts showing upper wall of ascoma.
c Section through the ascostroma. d Peridium. e Pseudoparaphyses. f–
j Development of asci. k–p Ascospores. Scale bars c = 200 lm, f–
j = 30 lm, b, d = 20 lm, e, k–p = 10 lm
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Fungal Diversity
123
Fungal Diversity
b Fig. 57 Phylogram generated from the best scoring of the RAxML
tree based on a combined ITS, LSU, TEF1-a and RPB2 sequence
dataset of Pseudohelicomyces and related genus Helicomyces in
Tubeufiaceae. Tubeufia tectonae (MFLUCC 17-1985) was selected as
the outgroup taxon. Bootstrap support values for maximum likelihood
(left) equal to or greater than 60% and the Bayesian posterior
probabilities (right) equal or higher than 0.90 BYPP are indicated
above the nodes. Ex-type and ex-epitype strains are in bold. Newly
generated sequences are indicated in blue
sexual morph species, while those other species in
Cyphellophora were found as asexual morph. The first
record of the sexual characters in Cyphellophora (C.
jingdongensis) was provided by Yang et al. (2018a).
However, C. jingdongensis differs from C. filicis in
growing on honey dew excretions from insects, with dark
superficial mycelium, scattered, superficial ascomata,
without short necks, ellipsoidal to cylindrical asci and 1–3septate fusoid ascospores. Thus, we introduce C. filicis as a
new species based on the sexual morph characters and
phylogenetic evidence (Figs. 59, 60).
Herpotrichiellaceae Munk
The family Herpotrichiellaceae was introduced by
Munk (1953) and placed in the order Chaetothyriales (Barr
1976; Réblová et al. 2013; Gueidan et al. 2014;
Wijayawardene et al. 2018a). The family is characterized
by small, superficial, inconspicuous, setose ascomata, fissitunicate asci and greenish grey to brown, phragmosporous or dictyosporous ascospores (Munk 1953; Hyde
et al. 2016). Species of this family can be saprobes on
decaying wood, bark and leaves and are also found as
pathogens on humans and living plants, as well as parasites
on fungi or lichens worldwide (Crous et al. 2007b;
Untereiner et al. 2011; Wijayawardene et al. 2017a). Fifteen genera are accepted in this family (Wijayawardene
et al. 2018a).
Capronia Sacc.
Capronia is a poorly understood ascomycete genus
characterized by very small, setose ascomata, lacking
paraphyses, fissitunicate asci, and septate, or muriform,
hyaline or pigmented ascospores (Munk 1957a, b; Müller
et al. 1987; Untereiner et al. 2011; Friebes 2012). Species
of this genus are associated with a wide range of hosts as
saprobes on rotting wood or bark and decaying stems and
leaves of herbaceous plants, or pathogenic on plants as well
as parasites on other fungi, or lichens (Untereiner 2000;
Untereiner et al. 2011; Wijayawardene et al. 2017a). We
introduce a new species Capronia camelliae-yunnanensis,
collected on Camellia yunnanensis in China (Figs. 61, 62).
Capronia camelliae-yunnanensis M. Raza, Z.F. Zhang &
L. Cai, sp. nov.
Index Fungorum number: IF555356; Facesoffungi
number: FoF04884, Fig. 62
Etymology: Named after the epithet of Camellia yunnanensis, the host of which the holotype was collected.
Holotype: HMAS 255435
Saprobic on Camellia yunnanensis. Sexual morph Ascomata 175–200 lm high, 215–220 lm diam., scattered or
clustered, solitary, superficial on decorticated bark of host,
with papilla, globose to subglobose, lodged on a basal
subiculum, which form loose hyphae penetrating the
underlying cells, setose around the surface of the wall,
setae up to 30 lm long. Peridium 20–30 lm wide, thickwalled, of equal thickness, composed of several layers of
small, pseudoparenchymatous cells, inner layers comprising hyaline cells, arranged in a textura angularis, outer
layers comprising brown to dark brown becoming blackened cells of textura prismatica. Hamathecium composed
of dense, 3–5 lm wide, broad, filamentous paraphyses with
indistinct septate, not constricted at the septa, embedded in
a gelatinous matrix. Asci (51–)57–78(–80) 9 (5–)6–10(–
12) lm (
x = 67.8 9 8.7 lm, n = 40), 8-spored, bitunicate,
fissitunicate, broadly cylindrical, short pedicellate with
knob-like pedicel, apically broad rounded, with a blunt
ocular chamber. Ascospores (10–)13–17(–19) 9 (2–)3–
5 lm (
x = 15.1 9 4.5 lm, n = 40), overlapping 1–3-seriate, hyaline and aseptate when young, becoming light
brown to yellowish, ellipsoidal to fusiform, muriform, with
3–7 transverse septa and 1–3 longitudinal septa, slightly
constricted at the septa, smooth-walled. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA reaching
15–20 mm diam. after 5 weeks at 25 ± 2 °C, colonies
circular, umbonate, smooth to woolly with entire edge,
velvety, smooth at the margin; from above, light green at
the fruiting zone, whitish green at the productive zone and
ageing zone; from below, light green at the fruiting zone,
dark green to blackish at the productive zone, grey at the
ageing zone; not producing pigmentation in PDA.
Material examined: CHINA, Yunnan Province, Baoshan
City, Longling County, on decorticated bark of Camellia
yunnanensis Cohen Stuart (Theaceae), October 2015, M.
Raza, HMAS 255435 (holotype), ex-type living culture,
CGMCC3.19061.
GenBank
numbers:
ITS = MH807377,
LSU =
MH807378, SSU = MH807379.
Notes: Capronia camelliae-yunnanensis forms a wellsupported clade, sister to C. pilosella (P. Karst.) E. Müll.
et al. (100% ML and 1.00 BYPP; Fig. 61). Capronia
camelliae-yunnanensis differs from C. pilosella in the shape
and size of the ascospores (C. camelliae-yunnanensis,
(10–)13–17(–19) 9 (2–)3–5 lm versus 12–14 9 4–4.5 lm,
C. pilosella; Karsten 1873). Capronia pilosella has phragmosporous, fusoid to clavate-fusoid, 3-septate ascospores
123
Fungal Diversity
Fig. 58 Pseudohelicomyces menglunicus (KUN-HKAS 85793, holotype). a, b Colonies on rotten seed coat. c Conidiophores with
conidiogenous cells. d, e Conidiophores. f Conidiophores bearing
123
conidia. g Conidiogenous cells. h–r Conidia. s Germinated conidium.
Scale bars a = 0.5 cm, b, d, e = 100 lm, f, s = 50 lm, c, g–
r = 20 lm
Fungal Diversity
Fig. 59 RAxML phylogenetic tree generated from a combined ITS,
LSU and SSU sequence dataset. Bootstrap support values equal or
above 50% and Bayesian values equal or above 0.90 BYPP are shown
at each node. Vonarxia vagans (CBS 123533) was used as the
outgroup taxon. Newly generated sequences are in blue
(Karsten 1873; Müller et al. 1987), while C. camelliaeyunnanensis has muriform, ellipsoidal to fusiform, ascospores with 3–7 transverse septa and 1–3 longitudinal septa.
Aspergilloides Dierckx, Furcatum Pitt., Penicillium, and
Biverticillium Dierckx, and 25 sections. Among the sections of Penicillium, section Citrina was introduced by
Houbraken and Samson (2011) based on phylogenies
derived from RPB1, RPB2, Tsr1, and Cct8 sequence data.
Currently, section Citrina contains 40 species (Houbraken
et al. 2011; Visagie et al. 2014a, b). Members of section
Citrina are found in soil, leaf litter, indoor environments,
and food (Pitt 1979; Pitt and Hocking 2009; Samson et al.
2010). The species of section Citrina are characterized by
the production of relatively small conidia, symmetrically
biverticillate conidiophores and ampulliform phialides.
They are also known for their ability to produce a variety of
extrolites, including mycotoxins, citrinin, and citreoviridin
(Houbraken et al. 2011). While, evaluating fungal diversity
in soil samples in Korea, a new species was isolated and is
described here based on morphological characteristics and
phylogenetic analyses (Fig. 63).
Subclass Eurotiomycetidae Geiser & Lutzoni
Eurotiales G.W. Martin ex Benny & Kimbr.
Trichocomaceae E. Fisch.
The family Trichocomaceae was introduced by Fischer
(1897). This is a large saprobic family in nature with the
most well-known genera including Aspergillus P. Micheli
ex Haller, Penicillium, and Paecilomyces Bainier. Species
belonging to this family have the ability to produce secondary metabolites (mycotoxins or extrolites), and
enzymes (Pitt and Hocking 2009; Samson et al. 2010;
Houbraken et al. 2011).
Penicillium Link
The genus Penicillium was first described by Link in
1809. Species of Penicillium are well known and found
abundantly in the soil, air, indoor environments and in
contaminated foods (Frisvad and Samson 2004; Samson
et al. 2010). According to Houbraken and Samson (2011),
the genus Penicillium was divided into four subgenera:
Penicillium dokdoense Hyang B. Lee & T.T.T. Nguyen,
sp. nov.
Index Fungorum number: IF554459; Facesoffungi
number: FoF013606, Fig. 64
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Fungal Diversity
Fig. 60 Cyphellophora filicis (KUN-KHAS 102220, holotype). a, b
Appearance of ascomata on host surface. c Section through ascoma.
d Section through peridium. e, f Asci embedded in a hyaline
gelatinous matrix. g–l Ascospores. m Ascospore germination. n, o
Culture characteristics on PDA (n = from above, o = from below).
Scale bars a = 500 lm, b = 100 lm, c, d = 20 lm, e, f = 10 lm, g–
l = 5 lm
Etymology: Named after the place where it was collected, Dokdo Island
Holotype: CNUFC-DDS11-1
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Colonies grow slowly on CYA, reaching
23.5–26 mm diam. at 25 °C in 7 days, grey-green, floccose
in centre, soluble pigment absent, margin entire; reverse
yellowish white with yellow-brown in centre. Conidiophores mostly biverticillate, sometimes monoverticillate,
or divaricate, stipes smooth, vary greatly in length, septate,
2–3.5 lm wide. Metulae cylindrical, 2–5 per stipe,
10.5–17.5 9 2.5–4.2 lm. Phialides ampulliform, 3–9 per
metula, 6.7–11.5 9 2–3.5 lm. Conidia roughened,
globose to subglobose, or ellipsoidal, and dark blue-green,
2–3.5 9 2–3 lm.
Culture characteristics: The isolate grew over a wide
range of temperatures with varying growth rates on MEA,
CYA, YES and CREA. The average growth rates of
CNUFC-DDS11-1 on MEA, CYA, YES, and CREA
medium at 25 °C were 34.5, 24.5, 22, and 14.5 mm per
7 days, respectively. Optimal growth was observed around
25 °C, slow growth was observed below 10 °C, and no
growth at 37 °C.
Material examined: REPUBLIC OF KOREA, from soil
in Dokdo Island in the East Sea of Korea, June 2014,
CNUFC-DDS11-1 (holotype); isotype in Korean
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Fungal Diversity
Fig. 61 Phylogenetic tree
generated from maximum
likelihood (RAxML) based on a
combined ITS, LSU and SSU
sequence dataset of
Herpotrichiellaceae. Maximum
likelihood bootstrap support
values greater than 50% and
Bayesian posterior probabilities
greater than 0.75 BYPP are
indicated on the notes. The new
isolate is in blue. The tree is
rooted with Vonarxia vagans
(CBS 123533)
Collection for Type Cultures (KCTC, Daejeon, Korea), extype living culture, JMRC:SF:013606.
GenBank numbers: ITS = MG906868, TUB2 =
MH243037, CMD = MH243031 (CNUFC-DDS11-1);
ITS = MG906869,
TUB2 = MH243038,
CMD =
MH243032 (CNUFC-DDS11-2).
Notes: The data from combined sequence analyses of
the two loci (Fig. 63) reveal that Penicillium dokdoense,
P. terrigenum Houbraken et al., P. cf. terrigenum and
P. copticola Houbraken et al. are closely related. Penicillium dokdoense shares several similarities with P. terrigenum as its growth on CREA is poor with no acid
production, but this species differs from P. terrigenum in
terms of reverse colour on CYA and YES, and colony
features on CYA. Furthermore, P. dokdoense produces
globose to subglobose, or ellipsoid conidia and
biverticillate, monoverticillate, or divaricate conidiophores,
in contrast to the mostly ellipsoid conidia and biverticillate
sporangiophores of P. terrigenum. Penicillium terrigenum
and P. copticola grew at a similar range of temperatures
from 25 °C to 30 °C, whereas P. dokdoense grew slowly.
Penicillium dokdoense grows and sporulates at 35 °C,
while, P. cf. terrigenum does not grow above 30 °C. At
5 °C, P. dokdoense and P. copticola were both capable of
growth, whereas P. terrigenum was not. When grown on
CYA, colonies of P. dokdoense are weakly wrinkled, while
colonies of P. terrigenum are strongly wrinkled. The results
of morphological and comparative sequence analyses of
P. dokdoense indicate that it is a distinct species from P.
terrigenum, P. cf. terrigenum, and P. copticola. Thus, P.
dokdoense is proposed.
Class Lecanoromycetes O.E. Erikss. & Winka
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Fungal Diversity
Fig. 62 Capronia camelliae-yunnanensis (HMAS 255435, holotype). a Blackish ascomata on decorticated bark of Camellia
yunnanensis. b Vertical section of ascoma. c Peridial structure.
d Paraphyses. e–h Asci. i Asci stained with Melzer’s reagent. j Apical
ring stained with Melzer’s reagent. k, l Immature ascospores. m, n
Mature ascospores. o Germinated ascospore. p, q Culture characteristics on PDA (p = from above, q = from below). Scale bars
b = 50 lm, c, d = 20 lm, e–i = 10 lm, o = 5, j–n = 2 lm
We follow the latest treatment and updated accounts of
Lecanoromycetes in Miadlikowska et al. (2014) and
Kraichak et al. (2018). The updated outline and notes of the
genera in Lecanoromycetes was provided by Wijayawardene et al. (2017a, 2018a).
Micropeltidaceae Clem. & Shear
The family Micropeltidaceae was introduced by Clements and Shear (1931) as ‘Micropeltaceae’ and is typified
by Micropeltis Mont. with M. applanata Mont. as the type
species. The family comprises foliar, biotrophic epiphytes,
which are mostly found on the lower leaf surface as small
black dots. Micropeltidaceae species are characterized by
Lecanoromycetes, families incertae sedis
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Fungal Diversity
superficial, flattened, black-blue or greenish to black
thyriothecia, easily removed from the host surface, poorly
developed at the base, the wall comprising interwoven
hyphae, with a central ostiole and ascospores are septate
and hyaline (Clements and Shear 1931; Wu et al. 2011;
Hyde et al. 2013; Hongsanan et al. 2015; Hongsanan and
Hyde 2017). Micropeltis phetchaburiensis sp. nov. is
introduced based on its morphological characteristics
coupled with phylogenetic analyses of a combined LSU
and SSU sequence dataset (Fig. 65). The new species was
collected from living leaves in Thailand.
Micropeltis Mont.
Fig. 63 Phylogenetic tree based on maximum likelihood analysis of a
combined beta tubulin (TUB2) and calmodulin (CMD) dataset for
Penicillium dokdoense and related species within the sect. Citrina.
Sequence of Penicillium corylophilum was used as outgroup taxon.
We follow the latest treatment and updated account of
Micropeltis in Hongsanan and Hyde (2017).
Micropeltis phetchaburiensis Dayarathne, Hongsanan &
K.D. Hyde, sp. nov.
Index Fungorum number: IF555294; Facesoffungi
number: FoF04841, Fig. 66
Etymology: Name reflects Phetchaburi Province in
Thailand, from where the species was collected.
Holotype: MFLU 18-1408
Epiphytic appearing as small black dots, superficial, on
the upper surface of living leaves, superficial hyphae
absent. Sexual morph Thyriothecia 75–90 9 140–160 lm
diam. (
x = 80 9 150 lm, n = 5), solitary, superficial on
Numbers at the nodes indicate the bootstrap values (C 50%) from
1000 replications. The bar indicates the number of substitutions per
position. New taxa are in blue and ex-type strains in bold
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Fungal Diversity
Fig. 64 Penicillium dokdoense (CNUFC-DDS11-1, holotype). a, e
Colonies in yeast extract sucrose agar (YES). b, f Colonies in malt
extract agar (MEA). c, g Colonies in Czapek yeast autolysate agar
(CYA). d, h Colonies in creatine sucrose agar (CREA). (a–d: obverse
view, e–h: reverse view). i–m Verticillate conidiophores and conidia
on phialides. n Conidia. Scale bars i = 20 lm, j–l = 10 lm,
m = 5 lm, n = 3 lm
the surface of hosts, circular, membranous, black, easy to
detached, base poorly developed, with a central, irregular
ostiole. Upper walls comprising an irregular, meandering
arrangement of hyphae, from the central ostiole to the
outside. Peridium 35–50 lm wide, composed of two strata,
the outer stratum having bluish to black, occluded walls,
inner stratum of greenish to hyaline, flattened cells. Hamathecium with evanescent pseudoparaphyses. Asci 50–
68 9 11–16 lm (
x = 60 9 14 lm, n = 10), 8-spored,
bitunicate, broadly cylindrical to fusiform, with a short
pedicel, apically rounded with ocular chamber. Ascospores
16–20 9 2–4 lm (
x = 18 9 3 lm, n = 20), overlapping
2–3-seriate, hyaline, clavate, 3-septate, constricted at the
septa, narrowly rounded at both ends, smooth-walled,
surrounded by a mucilaginous sheath. Asexual morph
Undetermined.
Material examined: THAILAND, Phetchaburi Province,
Prachuap Khiri Khan, 77230 Bang Saphan, Ron Thong, on
living leaves of an unidentified plant, 14 December 2015,
M. Dayarathne, KLAP011 (MFLU 18-1408, holotype;
HKAS102010, isotype).
GenBank numbers: LSU = MH656405, SSU = MH656406.
Notes: Micropeltis phetchaburiensis resembles M. dendrophthoes Hongsanan & K.D. Hyde and M. zingiberacicola H.X. Wu & K.D. Hyde in forming broadly cylindrical
to fusiform asci and hyaline, clavate, septate ascospores
(Wu et al. 2011; Hongsanan et al. 2015). Micropeltis
phetchaburiensis can be distinguished from M. dendrophthoes and M. zingiberacicola in having 3-septate ascospores with the 4 cells being equal in length and width,
while ascospores of the other two species are 4–5-septate
and enlarged at the first cell and relatively longer lower end
cells. In our phylogenetic analyses of a combined LSU and
SSU sequence dataset, M. phetchaburiensis forms a basal
lineage to M. dendrophthoes and M. zingiberacicola with
moderate bootstrap support (97% ML and 93% MP;
Fig. 65).
123
Class Leotiomycetes O.E. Erikss. & Winka
We follow the latest treatment and updated accounts of
Leotiomycetes in Zhang and Wang (2015), Jaklitsch et al.
(2016a) and Wijayawardene et al. (2018a) for the taxonomic outline of this class.
Helotiales Nannf. ex Korf & Lizoň
Fungal Diversity
Lachnaceae Raitv.
Raitviir (2004) raised Nannfeldt’s tribe Lachneae (fam.
Hyaloscyphaceae; Nannfeldt 1932) to the family level and
designated Lachnum Retz. as the type genus. Han et al.
(2014) showed that Lachneae formed a monophyletic lineage within Hyaloscyphaceae sensu lato, justifying the
existence of Lachnaceae. Jaklitsch et al. (2016a) and
Wijayawardene et al. (2018a) recognize 16 genera in this
family. Phylogenetic relationships of selected lachnoid taxa
were studied previously by Cantrell and Hanlin (1997) and
Hosoya et al. (2010). We introduce a new monospecific
genus Velebitea in the family Lachnaceae based on
microscopic and macroscopic features.
Velebitea I. Kušan, Matočec & Jadan, gen. nov.
MycoBank number: MB827753; Facesoffungi number:
FoF05713
Etymology: Named after the mountain, Velebit, on
which it was collected.
Sexual morph Ascomata apothecial, comparatively
robust and medium sized, superficial, stipitate, solitary or
gregarious, deeply cupulate when young, becoming shallowly cupulate to ± plate-shaped. Hymenium creamy
whitish, yellowish with age, margin upright, covered with
long whitish hairs, as well as whole excipulum and stipe
throughout the base, excipulum paler than hymenium, stipe
centrally attached, tapering towards the base, base brownish. Subhymenium equally thick or thicker than medullary
or ectal excipulum, composed of hyaline textura epidermoidea-intricata, in living state clearly discerned from the
medulla. Marginal texture of hyaline textura porrectaprismatica, often with abundant yellow resinous inclusions,
beset with ± tidily organized cylindrical marginal hairs
which are always clearly separated from excipular ones by
a small hairless area just at the point of hymenial base
level, individual hairs ± straight, multi-celled, markedly
shorter than excipular flank hairs, apical cell cylindricobtuse to subclavate, walls thin and hyaline, cells without
refractive content when in living state, only basal 1–2 cells
with firmly attached KOH resistant granules, not stainable
in cotton blue, with additional abundant loosely attached
KOH soluble pale yellowish resinous granules, no crystals.
Medullary excipulum composed of hyaline densely woven
textura intricata, producing small hyaline crystalloid particles with age, sometimes having abundant yellow resinous inclusions, intercellular spaces overall slightly
gelified. Ectal excipulum composed of single layer of textura prismatica with cells running ± parallel to the surface, walls somewhat thickened, especially in some
outermost cells, basal areas regularly with highly refractive
golden yellow resinous accumulations; surface beset with
very untidily intricately organized flexuous excipular hairs,
multi-celled, longer than marginal hairs, walls thin, with
apical localized thickenings in some hairs, only basal 1–2
cells with firmly attached KOH resistant granules, regularly
supplemented by KOH soluble loosely attached pale yellowish granules, surface irregularly embedded in abundant
gel plaques clearly stained lilac in brilliant cresyl blue, in
cotton blue walls not cyanophilic, yellow resinous accumulations dissolved. Overall texture and hairs in Lugol’s
solution without any amyloid reactions, all yellow resinous
accumulations in texture are rapidly soluble in KOH. Paraphyses cylindrical, apically obtuse, straight, not branching in the upper part, apical and often subapical cells
contain hyaline non-refractive vacuoles. Asci 8-spored,
elongated cylindrical-deltoid, apex conical obtuse, protruding above paraphyses at full maturity, arising from
simple septate ascogenous cells, in Lugol’s solution apical
apparatus moderately euamyloid of Calycina-type. Ascospores elongated fusoid, ± straight to bent, bilaterally
symmetrical, always 1-celled in full maturity, smooth,
hyaline, poles tapered to sub-obtuse, eguttulate or with
several minute guttules, uninucleate, the remaining sporoplasm regularly occupied by several conspicuous non-refractive vacuoles, when freshly ejected without sheath; in
Lugol’s solution cytoplasm without glycogen accumulations. Asexual morph Undetermined.
Type species: Velebitea chrysotexta I. Kušan, Matočec
& Jadan
Notes: A data matrix for alignment was constructed to
determine the phylogenetic position of Velebitea chrysotexta within Lachnaceae and also to test phylogenetic
proximity to Hyaloscyphaceae (Amicodisca virella (P.
Karst.) Huhtinen), and Tetracladium spp. Phylogenetic
analysis (Fig. 67) included the ITS and LSU sequences
generated from the holotype of Velebitea chrysotexta and
other related sequence data, retrieved from GenBank as
well as a newly sequenced collection of Neodasyscypha
cerina (Pers.) Spooner (CNF 2/10442; ITS = MH886408,
LSU = MH886412) which was collected from fallen
decorticated branch of Fagus sylvatica L. in Croatia.
Sequences of Hymenoscyphus fructigenus (Bull.) Gray
were used as an outgroup taxon. Maximum likelihood
analysis of the concatenated ITS and LSU alignment was
performed by MEGA7 (Kumar et al. 2016), including 1361
total characters in the final dataset. The phylogeny based
on concatenated analysis of ITS and LSU nests the genus
Velebitea in the family Lachnaceae as a separate lineage
having comparatively basal position not belonging to any
of tested genera (Fig. 67).
Megablast search of NCBIs GenBank nucleotide database using the ITS sequence of Velebitea chrysotexta (CNF
2/10072, GenBank no. MH886407) shows that the closest
hits belong to Lachnellula spp. with the similarities
between 89% and 91% similarities. Lachnellula P. Karst. is
a genus in Lachnaceae, strictly confined to coniferous hosts
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Fungal Diversity
(Dharne 1965). On the other hand, the closest hits using the
LSU sequence (GenBank no. MH886411) are Tetracladium furcatum Descals (GenBank no. EU883428; similarity = 907/1009 (90%), gaps = 45/1009 (4%)) and T.
maxilliforme (Rostr.) Ingold (GenBank no. EU883429 and
EU883430; both with similarity = 903/1008 (90%),
gaps = 43/1008 (4%)). Tetracladium De Wild. is an
aquatic hyphomycete genus (Letourneau et al. 2010) classified as Helotiales, incertae sedis (Wijayawardene et al.
2017a, 2018a), but phylogenetic affinity is close to Helotiaceae, and Hyaloscyphaceae (Wang et al. 2015). Certain
similarity exists between Velebitea and Tetracladium in
their ecological preference to a colder climate; the sexual
morph of Velebitea lives in subalpine forests with beech
(Fagus) while some members of the genus Tetracladium
were recorded in cold snow-covered soil in a glacier zone
(Kuhnert et al. 2012).
Together with the genus Dasyscyphella Tranzschel,
Velebitea is different from all other genera in the family
Lachnaceae by partially granulated hyaline hairs.
Dasyscyphella is clearly polyphyletic (Hosoya et al. 2010;
this study) and therefore in need of taxonomic rearrangement. Since the type species of the genus Dasyscyphella,
D. cassandrae Tranzschel is not present in public DNA
sequence databases, we included all other available species
from this genus in our phylogenetic analysis because species currently ascribed to Dasyscyphella display highest
similarity to our material based on non-molecular data. The
polyphyly of the genus Dasyscyphella could be in line with
wide diversity of certain non-molecular features that may
play important role in future species distinguishing at
generic level, such as existence of hair crystals, resinous
exudates and its microchemical properties, hair wall
granulation degree, hamathecial features, details in ecology, apothecial development and anatomy, despite of high
similarity among many of species in terms of ‘‘standard’’
microscopical characters (e.g. spore and ascus shape/measurements). Many of these characters already efficiently
delimitate some monophyletic lachnacean genera, viz.
Capitotricha (Raitv.) Baral, Brunnipila Baral and
Fig. 65 Phylogram generated from maximum likelihood (RAxML)
analysis based on combined LSU and SSU sequence dataset of
representative families in Lecanoromycetes. Tree is rooted with
Icmadophila ericetorum (AFTOL-ID 4846) and Siphula ceratites
(P110). Maximum likelihood bootstrap (black) and maximum parsimony bootstrap (blue) values [ 65% are given above the nodes. The
scale bar indicates 0.05 changes. New isolate is in blue. Ex-type
strains are indicated in bold
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Fungal Diversity
Fig. 66 Micropeltis phetchaburiensis (MFLU 18-1408, holotype).
a–c Appearance of thyriothecia on host surface. d Thyriothecium
viewed in squash mount. e Vertical section through thyriothecium. f,
g Asci. h–k Ascospores. Scale bars c = 100 lm, d, e = 50 lm, f, g,
k = 20 lm, h–j = 10 lm
Lachnellula. Dasyscyphella cassandrae, the type species of
the genus, differs strikingly from Velebitea chrysotexta by
very elongate flexuous-cylindric, septate ascospores,
overall absence of KOH soluble resinous lumps and,
together with all other known species currently ascribed to
the genus Dasyscyphella in having only the apical (rarely
also subapical) smooth hair cell. Whereas, hairs in V.
chrysotexta are smooth except in basal 1–2 cells. Hairs in
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D. cassandrae (as well as in many other species) are longer
at the margin than on excipular flanks, quite opposite in V.
chrysotexta. A number of species currently ascribed to a
genus Dasyscyphella have hairs bearing conspicuous calcium oxalate crystals and/or crystal druses (such as D.
nivea positioned well aside from V. chrysotexta in our
phylogenetic analysis) that are completely lacking in V.
chrysotexta. Nearly all species currently ascribed in the
genus Dasycyphella have either lanceolate or cylindricalpointed paraphyses (as in D. cassandrae). There are however some species of the genus Dasyscyphella that
according to Raitviir (2002), also Dennis (1949), produce
resinous pigments comparable to Velebitea chrysotexta:
Dasyscyphella claviculata (Velen.) Baral & Svrček, D.
crystallina (Fuckel) Raitv., D. rubi Raitv., D. conicola
(Rehm) Raitv. & Arendh., D. nivea (R. Hedw.) Raitv., D.
tamajonica (Raitv. & R. Galán) Raitv. (all producing calcium oxalate crystals, mostly having different ecology), D.
mughonicola (Svrček) Raitv. & Arendh. (whose asci arising from croziers, coniferous substrate), D. patuloides
Raitv. & R. Galán (producing hyaline resin, asci arising
from croziers, having phyllophilous ecology) and D. sulphuricolor (Peck) J.H. Haines (producing brownish resin
that reacts vinaceous with KOH). Much work is still to be
done to uncover true phylogenetic affinities of a number of
species currently accommodated in Dasyscyphella which
might belong to severally phylogenetic lineages representing separate genera.
Velebitea chrysotexta I. Kušan, Matočec & Jadan, sp. nov.
MycoBank number: MB827754; Facesoffungi number:
FoF05714, Figs. 68, 69, 70
Etymology: Refers to its yellowish pigment inclusions in
the apothecial texture
Holotype: CNF 2/10072
Sexual morph Ascomata apothecial, superficial, stipitate, solitary or gregarious, deeply cupulate when young,
becoming shallowly cupulate to ± plate shaped,
2.5–3.2 mm high, 1.1–3 mm diam. Hymenium 80–105 lm
thick, creamy whitish in primordial stage, becoming
creamy to pale ochre yellow, yellowing with age, margin
upright, covered with whitish hairs, as well as whole
excipulum and stipe throughout the base, excipulum paler
than hymenium, completely covered with whitish hairs,
stipe 1.5–2.2 9 0.6–1 mm, centrally attached, tapering
towards the base, hairy throughout, base brownish. Subhymenium 32–47 lm thick, composed of hyaline textura
epidermoidea-intricata, cells 3.4–5.1 lm wide, while in
living state clearly discerned from the medulla. Marginal
texture forming very thin layer, composed of hyaline textura porrecta-prismatica, often containing abundant yellow
resinous accumulations, beset with ± tidily organized
marginal hairs which are always clearly separated from
123
excipular ones by small hairless area just at the point of
hymenial base level, individual hairs ± straight but
bending slightly towards the hymenial rim, projecting
towards the marginal surface perpendicularly, 5–7-celled,
66–109 lm long, 3.7–5.6 lm wide at the base, 2.2–3.8 lm
in the middle and 2.3–3.7 lm in the apical part, apical cell
cylindric-obtuse to subclavate, individual cells prismatic,
containing low refractive globules, walls hyaline and
refractive, 0.2–0.5 lm thick, may be slightly thickened in
dead state, cells without refractive content when still alive,
wall of the basal 1–2 cells firmly granulate, additionally
beset with loosely attached pale yellowish granules, no
crystals; in brilliant cresyl blue numerous non-refractive
hyaline vacuoles stained deep lilac-violet to violet after
longer exposure, cytoplasm in dead apical cells turquoise
blue, basal wall granules grey cyan to violet grey, walls
unstained; after adding 2.5% KOH hair granules persistent,
subhyaline golden yellow accumulations instantly dissolved, for few moments giving localized yellow solution
area; in cotton blue hair walls not cyanophilic, apical cells
cytoplasm pale blue, wall granules pale cyan, yellow
resinous accumulations dissolved. Medullary excipulum
30–44 lm thick, composed of hyaline densely woven textura intricata intertwining with ± vertical cells, cells thinwalled, 2.2–3.7 lm wide, running predominantly parallel
to the excipulum surface, producing hyaline crystalloid
particles with age, sometimes containing abundant yellow
resinous accumulations; in brilliant cresyl blue intercellular
space slightly rosy-lilac (gelified); in cotton blue walls not
cyanophilic. Ectal excipulum 38–52 lm thick, composed
of textura prismatica, cells 13.6–46.7 9 4.5–10.9 lm,
running ± parallel to the surface, walls somewhat thickened, especially in some outermost cells, basal area with
highly refractive golden yellow resinous accumulations; surface ornamented with flexuous excipular hairs
which are very untidily intricately organized, 4–11-celled,
92–156 lm long, 2.5–4.1 lm wide at the base, 2.2–3.1 lm
in the middle part and 2.4–3.7 lm at the apex, individual
cells prismatic, walls highly refractive, sometimes with
localized apical thickenings, in dead state more pronounced, 0.7–0.8 lm thick, basal 1–2 cells firmly granulate, with additional loosely attached pale yellowish
granules, no crystals; in brilliant cresyl blue textural cells’
cytoplasm lilac, thickest walls greenish cyan, slightly
gelified, surface with revealed abundant, lilac irregular gel
plaques; in cotton blue walls not cyanophilic, cytoplasm
bluish, yellow resinous accumulations dissolved. Stipe
excipulum of hyaline celled, wavy textura prismatica, cells
17.4–34.2 9 5.6–9.3 lm, walls thickened in the outermost
cells, giving rise to hyaline hairs as those on ectal excipulum, 90–136 lm long, 3.8–4.4 lm wide at the base,
2.7–3.5 lm in the middle and 1.8–2.8 lm at the apical part,
containing rich golden-yellow partly crystalloid
Fungal Diversity
accumulations. Overall texture and hairs in Lugol’s solution without any amyloid reactions, while resinous matter
is rapidly soluble by KOH. Paraphyses cylindrical, apically
obtuse, straight, not branching in the upper part, apical cell
19.3–35.7 9 2.1–3.2 lm, apical and often subapical cells
contain hyaline non-refractive vacuoles in a living state
(not vacuolar bodies), dead cells with highly refractive
lemon yellow content; in Lugol’s solution, brilliant cresyl
blue and cotton blue unstained. Asci 72.6–95.1 9
5.5–7.2 lm, elongated cylindrical deltoid, apex conical
obtuse, protruding above paraphyses tips up to 12 lm at
full maturity, pars sporifera 23.5–38.6 lm, 8-spored,
arising from simple septate ascogenous cells, sometimes
with lateral protuberance; in Lugol’s solution apical
apparatus moderately euamyloid, of Calycina-type. Ascospores (8.2–)8.4–10.8–14.2(–15.2) 9 (2.1–)2.2–2.6–2.8
lm, Q = (3.2–)3.3–5.4–5.7(–6) (n = 125), elongated
fusoid, ± straight to slightly bent or sickle shaped, rarely
subsigmoid, bilaterally symmetrical, poles tapered to subobtuse, 1-celled, smooth, hyaline, overmatured 1-septate,
eguttulate or with several minute guttules, 0.2–0.6 lm
diam., uninucleate, nucleus centrally positioned,
1.3–1.4 lm in diam., the remaining sporoplasm regularly
occupied by several conspicuous non-refractive vacuoles;
3–4 ascospores at uppermost positions in living mature asci
markedly shorter, mostly 2-seriate, when freshly ejected
without sheath; in Lugol’s solution cytoplasm partly yellowish, without glycogen accumulations, in brilliant cresyl
blue wall unstained, no sheath revealed, after longer
exposure internal small metachromatic corpuscles regularly formed. Ascospores in polysporic test cultures under
axenic conditions obtained by shooting asci on PDA
readily germinated nearly equally at both poles during 24 h
at 24 °C. Asexual morph Undetermined.
Habitat and phenology: Saprobic on very rotten decorticated fragments and large branches of Fagus sylvatica
Fig. 67 Maximum likelihood
phylogenetic tree based on a
concatenated ITS and LSU
sequence dataset. Sequences
recovered during this study are
shown in blue. The tree is
rooted to Hymenoscyphus
fructigenus. Maximum
likelihood bootstrap support
values greater than 50% and
Bayesian posterior probabilities
greater than 0.95 BYPP are
shown at the nodes. The bar
length indicates the number of
nucleotide substitutions per site
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Fungal Diversity
Fig. 68 Velebitea chrysotexta. a Fructification growth in situ.
b Apothecia, side view, resinous plaques visible. c Ascomata top
view. d Living mature asci protruding above paraphyses tips.
e Paraphyses. f Simple septate ascogenous cells. g Freshly ejected
living mature ascospores. h Overmature ascospores. i Freshly ejected
living mature ascospores. j Germinated ascospores on PDA after
24 h. k Amyloid reaction in asci. l Marginal hairs tips. m Marginal
apothecial area largely beset by resinous matter. d–i, m in water
mount, k, l in Lugol’s solution, j on PDA. c–h, l CNF 2/10072
(holotype), i, j, m CNF 2/10661, a, b, k CNF 2/10736. Photo by N.
Matočec and I. Kušan. Scale bars a, c = 2 mm, b = 1 mm, d–i, k,
l = 10 lm, j, m = 50 lm
(Fagaceae) hidden in the litter, in subalpine forests, fruit
bodies appear in May and June.
Known distribution: The species is known so far only
from Mt. Velebit, Croatia.
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Fungal Diversity
Fig. 69 Velebitea chrysotexta. a Apothecial section beset with
resinous matter. b Excipular flank with hairs. c Excipular hair base.
d Marginal texture with hairless area. e Marginal hairs. f Marginal
hair tips in bright field (left) and phase contrast (right). g Excipular
texture with thick subhymenial layer. h Medulla with crystalloid
particles. i Large resinous accumulations in ectal excipulum.
j Excipular hairs, apical wall thickenings present in some hairs,
phase contrast (left), bright field (right). k Excipular hairs in dead
state. All in water mount. b, d, f, j, k CNF 2/10072 (holotype), a, c, g,
i CNF 2/10661, e, h CNF 2/10736. Photo by N. Matočec and I. Kušan.
Scale bars a = 50 lm, b, d, e, g–k = 20 lm, c, f = 10 lm
Material examined: CROATIA, Lika-Senj County,
Sjeverni Velebit National Park, northern part of Mt.
Velebit, Jurekovac area, 1050 m E-SE from Jurekovački
kuk peak (1525 m), 44°450 1600 N, 15°000 5700 E, 1340 m
a.s.l., forest of Abies alba Mill., Fagus sylvatica L. and
Picea abies (L.) H. Karst., on decorticated rotten Fagus
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Fungal Diversity
Fig. 70 Velebitea chrysotexta. a Marginal hairs with resinous lumps.
b Stipe vestiture, side view. c Stipe texture and vestiture in
section. d Stipe hairs. e Living mature ascospore. f Marginal hairs
with rapidly floculated surface matter. g Marginal hairs after longer
exposure. h Excipular hairs in living and dead state. i Excipular tissue
displaying gelified areas (lilac and rosy). j Living excipular hairs.
k Medullary excipulum. l Apothecial section and vestiture, dark field.
m Apothecial vestiture, phase contrast. n Marginal hairs. o Hymenial
elements. p Excipular tissue, resinous lumps dissolved. q Excipular
hairs. a–d in water mount, e–k in brilliant cresyl blue, l–o in cotton
blue, p, q in KOH. b–q CNF 2/10072 (holotype), a CNF 2/10661.
Photo by N. Matočec and I. Kušan. Scale bars b, c, l, m = 50 lm, a,
d, f–k, n, p, q = 20 lm, o = 10 lm, e = 5 lm
sylvatica stump base together with Chlorociboria aeruginosa (Oeder) Seaver ex C.S. Ramamurthi et al. and
Hyaloscypha vitreola (P. Karst.) Boud., 27 May 2017, I.
Kušan and N. Matočec (CNF 2/10072, holotype); ibid. 2
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June 2018 (CNF 2/10661); Lika-Senj County, Paklenica
National Park, southern part of Mt. Velebit, Velika Ruja in
Javornik area, 1160 m NE-E-NE from Badanj peak
(1638 m), 44°220 6000 N, 15°270 2200 E, 1365 m a.s.l., forest of
Fagus sylvatica, on fallen decorticated branch of Fagus
sylvatica, 25 June 2018, N. Matočec and I. Kušan (CNF
2/10736).
GenBank
numbers:
ITS = MH886407,
LSU =
MH886411 (CNF 2/10072, holotype).
Class Pezizomycetes O.E. Erikss. & Winka
We follow the latest treatment and updated accounts of
Pezizomycetes in Jaklitsch et al. (2016a), Ekanayaka et al.
(2018) and Wijayawardene et al. (2018a).
Pezizales J. Schröt.
Pezizaceae Dumort.
The family Pezizaceae was introduced by Dumortier
(1829) with Peziza Fr. as the type genus. A systematic
overview of the family was recently given by Jaklitsch
et al. (2016a) who recognized 32 genera in this family.
Ekanayaka et al. (2018) reviewed the families in the class
Pezizomycetes and outlined 45 genera names belonging to
Pezizaceae, including also some taxa synonymized by
other authors.
Sarcopeziza Loizides et al.
A new genus in the Pezizaceae, Sarcopeziza Loizides
et al. was recently erected (Agnello et al. 2018) to
accommodate a single species, Peziza sicula Inzenga
(Inzenga 1869). This species was previously typified and
treated by Agnello et al. (2013) and Agnello et al. (2015).
Since the original description of Inzenga (1869) and the
modern presentations by Agnello et al. (2013) and Agnello
et al. (2018) are not fully adequate and accurate for further
advancement in taxonomy of the group, a more detailed redescription of microscopical features of Sarcopeziza sicula
is provided here, based on careful cyto- and histochemical
analyses.
Sarcopeziza sicula (Inzenga) Agnello, Loizides & P.
Alvarado, Ascomycete.org 10(4): 179 (2018)
MycoBank number: MB827574; Facesoffungi number:
FoF05715, Figs. 71, 72
Basionym: Peziza sicula Inzenga, Funghi Siciliani,
Centuria Seconda: 29 (1869)
= Sarcosphaera sicula (Inzenga) Pat., Bull. Soc. Hist.
nat. Autun 17: 154 (1904)
Sexual morph Ascomata apothecial, hypogeous to
semi-hypogeous at first, becoming epigeous during development, firstly globose with pre-defined narrowly circular
apical opening, 2–5 cm in diam., expanding when ripe,
cracked into lobes, becoming star-shaped, solitary or gregarious, reaching 4.5–10 cm in diam., base substipitate to
stipitate deeply buried in the substrate, hymenial surface
vinaceous-purplish to livid vinaceous, matte, margin
tapered in section, entire in youth, smooth, excipulum
concolorous or somewhat darker than hymenium, gibbous, ± smooth to finely pruinose; flesh of homogenous
soft waxy consistence, on cut not exuding milk,
2.5–4.2 mm thick, of the same colour as external surface
with more pinkish tinge, odourless and flavourless. Hymenium 360–400 lm thick, arranged as a regular palisade.
Subhymenium reduced and faintly discerned from the upper
medulla only by scattered ascogenous hyphae, other cells
indistinct from those in upper medulla though slightly
smaller, continuing upwards as moniliform lower paraphysal cells. Upper medulla 950–1200 lm thick, composed of textura angularis richly intertwined with elements
of short-celled textura intricata, nearly ± isodiametric
cells gradually more frequent and larger towards lower
medulla, 13.5–55 lm diam., elongated cells 8.5–88 9
5.3–16.5 lm, walls hyaline, thin, many cells contain minute refractive crystalloid particles in water mount, but
without globular content, highly refractive globules though
present in cotton blue, 0.8–5.2 lm diam., but crystalloid
matter absent; lower zone with many cells having thickened yellowish and highly refractive septal rings. Lower
medulla 500–900 lm thick, generally as central textura
intricata, but cell bundles in lower half mostly oriented ± vertically as textura fasciculata, while on the
border with upper medulla or in central areas hyphae may
be oriented predominantly ± horizontally, hyphae in upper
1/3 of the whole layer often without strict orientation, cells
cylindrical-hyphoid, 6.2–14.8 lm wide, without any type
of refractive contents, walls subhyaline, some ± cylindrical cells densely set with rich accumulations of goldenyellow and highly refractive minute granules that may
consolidate to form irregular cytoplasmic patches of isabelline, extracellular crystalloid content very sparse. Ectal excipulum compact area 220–375 lm thick, composed
of textura angularis to textura prismatica, cells
11.4–28.5 9 6.2–24.5 lm, some cells somewhat elongated, more elongated cells dominant in upper zone and
then of ± vertical orientation, walls thickened, subhyaline
to yellowish, 0.7–0.9 lm thick in cortical cells, inner cells
with thin, hyaline walls, containing minute to medium
sized refractive crystalloid particles in water mount, but
without globular content, highly refractive globules though
present in cotton blue, 0.8–6.5 lm diam., but crystalloid
matter absent; surface with occasional low pustules, ornamented chiefly by individualized, mostly scattered,
(1–)2–6-celled terminal outgrowths, 21.4–125 9 9.5–14.7 lm,
individual cells sub-cuboid to cylindrical, 9.2–40 9
9.1–14.7 lm, walls subhyaline, thin-walled; cortical cells
beset with yellowish to isabelline-ochre highly refractive
minute to larger crystalloid matter, cytoplasm of some
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Fungal Diversity
cortical cells pale isabelline. In KOH refractive cellular
content in upper medulla coalesced, lower medullar/ectoexcipular cells without refractive content, crystals not
dissolved. Overall texture in Melzer’s reagent without dextrinoid reactions, refractive cellular granules coalesced to
highly refractive large guttules; in congo red all cell
walls ± stained; in cotton blue cell walls cyanophobic in
entire excipulum, but some ectal excipular cells with bluish
cytoplasm, medullar cell walls not cyanophilic. Paraphyses
cylindrical, inconspicuous, apically obtuse to clavate,
straight to apically bent, not branching in the upper part,
apical cell 2.8–4.8 lm diam., middle and lower cells
cylindric, moniliform cells exist near the margin, wall
hyaline, thin; in Lugol’s solution unstained. Asci
355–446 9 14–17.3 lm, cylindrical, apex subtruncate with
comparatively weakly pronounced functional operculum,
8-spored, base pleurorhynchous, with two ± tightly set
septa, arising from non-repetitive croziers at widely variable
levels, operculum 5.2–5.8 lm in diam., 0.7–0.9 lm thick,
strictly apical, flat-lenticular with only slight indentation ring
best visible under oil immersion in cotton blue, periascal
mucus very thin without thick apical accumulation, filmlike, evenly thick throughout, lateral wall 3-layered, subhyaline to yellowish, 0.7–0.9 lm thick; in Lugol’s solution
periascal mucus in the broad upper area diffusely and
moderately euamyloid, reaction gradually decreasing in
intensity downwards, basally inamyloid, same in Melzer’s
reagent; in congo red periascal mucus weakly pronounced,
very thin, outermost wall layer strongly grey-red, median
layer less stained, innermost layer unstained, operculum
differentially stained red while thin operculum ring area
remained unstained. Ascospores (13.2–)13.4–15.1–16.1
(–16.7) 9 (8.1–)8.3–8.9–10,
Q = 1.48–1.57–1.76(–1.77)
(n = 50, in cotton blue), hyaline, ellipsoid, sometimes
broadly oblong, radially symmetrical, with rounded poles,
1-celled, seemingly smooth under immersion lens when in
water mount, many spores (especially when outside of the
asci) slightly roughened in cotton blue but markings are not
cyanophilic, measuring below 0.2 lm high and up to 0.3 lm
diam., mature spores initially with multiguttulate pattern,
lipid bodies, when still not coalesced 0.3–1.5 lm diam.,
lipid bodies in dead spores coalesced in various degree up to
maximal, uni-guttulate stage (visible only in water mount),
uninucleate (visible in spores having lipids in multi-guttulate
pattern), wall 3-layered, 0.7–0.8 lm thick; in Lugol’s solution without glycogene accumulations; in KOH wall
stable (not loosening); in cotton blue perispore weakly
cyanophilic, wall not loosened, de Bary bubbles present in
some mature spores, lipid bodies completely masked,
sporoplasm unstained; in acetocarmine nucleus not differentially stained, only submature spores with weakly stained
nucleus, perispore not loosened. Asexual morph
Undetermined.
123
Material examined: ITALY, Province of Brindisi, San
Pancrazio Salentino, San Antonio alla Macchia,
40°250 0000 N, 17°500 0000 E, 59 m a.s.l., on bare and mossy
soil in a sunny Pinus halepensis Mill. forest with Eryngium
campestre L. and Poaceae, 7 February 2008, A. Delle
Donne and C. Agnello (MCVE 25877, epitype).
Habitat and phenology: On sandy or calcareous soils in
frame of dry Mediterranean thickets (mostly dominated by
Cistus spp.) and grasslands and parks, sometimes in
vicinity of thermophilic Pinus spp., Pyrus spp. and Olea
europea in thermo-Mediterranean bioclimatic belt, fruit
bodies appear between January and April.
Known distribution: Cyprus, Greece, Israel, Italy, Spain
and Tunisia.
GenBank numbers ITS = MH886405, LSU = MH886409
(MCVE 25877, epitype).
Notes: A data matrix for alignment was constructed to
show a phylogenetic position of Sarcopeziza sicula within
Pezizaceae, with a special emphasis on the genus Peziza.
Phylogenetic analysis (Fig. 73) included the ITS and LSU
sequences generated from the epitype collection of Sarcopeziza sicula and other related sequences retrieved from
GenBank as well as a newly sequenced collection of
Adelphella babingtonii (Berk. & Broome) Pfister, Matočec
& I. Kušan (CNF 2/9430; ITS = MH886406, LSU =
MH886410). Ascobolus crenulatus P. Karst. was used as an
outgroup taxon. The maximum likelihood analysis of a
combined ITS and LSU sequence dataset was conducted in
MEGA 7 (Kumar et al. 2016) with a total of 1462 positions
in the final dataset.
The phylogeny based on concatenate analysis of ITS and
LSU nested the genus Sarcopeziza in the family Pezizaceae, in a species group along with Eremiomyces Trappe &
Kagan-Zur, Hapsidomyces venezuelensis J.C. Krug & Jeng
and Peziza phyllogena Cooke, next to the Terfezia-Tirmania, P. depressa, P. saniosa and Peziza ostracoderma
clades (Fig. 73). Eremiomyces echinulatus (Trappe &
Marasas) Trappe & Kagan-Zur and E. magnisporus G.
Moreno et al. cluster with high support with Sarcopeziza
sicula showing a high level of genetic similarity. Sarcopeziza sicula demonstrates a distant relationship to
Peziza vesiculosa Bull. and Sarcosphaera coronaria
(Jacq.) J. Schröt. which are type species of the genera
where it was previously combined (Inzenga 1869;
Patouillard 1904), suggesting taxonomic affinity outside of
those genera.
Besides molecular evidence, Sarcopeziza sicula is different from Peziza sensu stricto [a core species group
gathered around the type species, P. vesiculosa, cf. Hansen
et al. (2001, 2002, 2005)], primarily by faintly visible, filmlike periascal mucus that is revealed by moderately diffusely amyloid reaction in iodine mounts, gradually
decreasing in strength downwards. Whereas, all species
Fungal Diversity
Fig. 71 Sarcopeziza sicula. a Ascomata and part of hymenium after
Inzenga (1869), scanned from Venturella (2005). b Dried ascomata.
c Ascus amyloidity. d Opercular fine structure. e Ascus bases with
ascogenous cells. f Ascospore ornamentation. g Ascus bases with
ascogenous cells. h Ascospores. i Mature ascospores in multiguttulate state displaying single nucleus. j Mature ascospores during
coalescence of lipid bodies. k Completely coalesced lipid body
matter. l Ascospores with de Bary bubble present in some spores. c in
Lugol’s solution, e, g, i–k in water mount, d, f, h, l in cotton blue. All
from MCVE 25877 (epitype). Photo by N. Matočec and I. Kušan.
Scale bars b = 1 cm, c–l = 10 lm
from Peziza sensu stricto have opercular (apical) thick
accumulation of heavily amyloid mucus giving strong ringlike amyloid reaction confined to the ascal tops. Agnello
et al. (2018) define asci as diffusely amyloid along the
entire length of the asci and strongly amyloid at the apex
(type I, cf. Hansen et al. 2001). Judging to both the authors’
microphotograph and own observations, the type of amyloidity belongs to type III instead of type I. The fine
structure of the ascus wall lacks strong apical indentation
ring and lentiform operculum in Sarcopeziza sicula, characters that are constantly present in all studied species in
Peziza core species group as well as in ascobolacean representatives and species of Peziza succosa and P. succosella clade (pers. data, cf. Samuelson 1978).
Moreover, Sarcopeziza sicula is also different by ontogenetically and cytochemically comparatively stable multiguttulate lipid configuration in fully developed ascospores
and violet-purplish in flesh. Contrary to Agnello et al.
(2013, 2018), the spores in this study are recognized as
multi-guttulate when fully mature and in living state whose
lipids coalesce in dead state towards uni-guttulate or more
rarely bi-guttulate pattern. The process of lipid body coalescence in dead spores is regularly observed throughout
Ascomycota (Baral 1992). The microphotograph of ripe
asci in Agnello et al. (2018, Fig. 2f), is clearly displaying
multi-guttulate pattern of living and fully mature ascospores, while Fig. 2c, d in Agnello et al. (2013) resemble
123
Fungal Diversity
Fig. 72 Sarcopeziza sicula. a Excipular structure (bright field).
b Excipular structure (dark field). c Excipular structure, upper half
with hymenium (dark field). d Excipular structure, lower half (dark
field). e Subhymenium with minute crystalloid particles. f Upper
medulla with minute crystalloid particles. g Upper medulla with lipid
like globules. h Ectal excipulum with minute crystalloid particles.
i Cortical cells with yellow cytoplasm and hyaline hyphoid
outgrowths. j Subhymenium and upper medulla. k Upper medulla
with largest cells. l Lower medulla. m Lower medulla with fasciculate
texture. n Ectal excipulum. a, b, e, f, h in water mount, c, d, g, i–n in
cotton blue. All from MCVE 25877 (epitype). Photo by N. Matočec
and I. Kušan. Scale bars a–d = 200 lm, e–i = 20 lm, j–n = 50 lm
dead ascospores containing subsequently coalesced lipid
bodies.
Furthermore, the authors presumed that ascospore
ornamentation is not constant character in this species. It is
quite possible that ornamentation in this species is normally formed after spore ejection and extraascal dormancy.
Both available ascomata from the epitype produced large
number of dormant ascospores whose fine ornamentation is
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Fungal Diversity
clearly visible in lactic acid cotton blue under oilimmersion.
From the genus Sarcosphaera Auersw. (its type and the
only species S. coronaria), Sarcopeziza sicula is different
by the absence of paraphysal internal content, different
original lipid body configuration in mature ascospores
(multi-guttulate vs. bi-guttulate), intensively pigmented
flesh and the excipular surface, substipitate to stipitate
apothecia, and lack of discernable odour (S. coronaria has
somewhat chlorine scent).
Agnello et al. (2018) describe the subhymenium as
150–180 lm thick layer but we detected it as very reduced
and unclearly discerned tissue from the upper medulla
using high contrasting techniques on a number of entire,
thin sections (ranging from hymenium up to ectal excipular
layer) from the epitype material.
The closest pezizacean relatives of Sarcopeziza sicula
revealed in our phylogenetic analysis are species of the
genus Eremiomyces, Hapsidomyces venezuelensis and
Peziza phyllogena (a member of the genus Peziza sensu
lato). All so far produced DNA based phylogenies (along
with the present one), however, clearly showed polyphyly
in the genus Peziza, where numerous species attributed to
the other genera (especially of hypogeous representatives)
build up monophyletic clades along with certain members
ascribed to a genus Peziza. Therefore, all species putatively
ascribed to a genus Peziza should be carefully restudied
using integrative taxonomic approach. Sarcopeziza sicula
is readily recognizable from all other currently known
genera in the family Pezizaceae by specific combination of
macroscopical features (semi-hypogeous sarcosphaeroid
apothecia with unique pigmentation), ecology (subpsammophilic and/or xeric habitats in thermo-Mediterranean
zone) and microscopy, principally cytochemistry and fine
structure of the asci and multi-guttulate lipid body configuration of the ascospores, characters that were proved to
be informative for differentiation in Pezizaceae in a previous study (Pfister et al. 2009).
There is a number of similarities in excipular structures
of semi-hypogeous species Sarcopeziza sicula and
peridial/glebal microscopic structures of the closest relative, hypogeous species Eremiomyces magnisporus (cf.
Alvarado et al. 2011): (1) the same texture in peridia of E.
magnisporus and cortical layer of ectal excipulum in S.
sicula; (2) cytoplasmic pigments in peridial cells of E.
magnisporus and cortical layer cells of S. sicula; (3) same
texture in glebal sterile veins in E. magnisporus and
medullary excipulum in S. sicula; and (4) lipid-like globules in glebal sterile vein cells in E. magnisporus and
medullary excipular cells in S. sicula.
The other hypogeous species Eremiomyces echinulatus
which is a type species of the genus Eremiomyces, and E.
innocentii Ant. Rodr. & Bordallo as well as
phylogenetically close Kalaharituber pfeilii (Hennings)
Trappe & Kagan-Zur (cf. Ferdman et al. 2005 and Crous
et al. 2017) also possess very similar structures in peridial
and glebal cells and texture. Very similar excipular texture
exists in Peziza phyllogena too (pers. data) and in wider
apothecial marginal area of Hapsidomyces venezuelensis
(cf. Krug and Jeng 1984), two epigeous phylogenetically
closest relatives. Nearly the same fine opercular structure in
cotton blue mount is found in both Sarcopeziza sicula and
Peziza phyllogena (pers. data). On the other hand, genera
Eremiomyces and Kalaharituber Trappe & Kagan-Zur
differ from Sarcopeziza sicula by their entirely hypogeous
living strategy developing permanently closed stereothecia
with solid gleba and producing perfectly globose coarsely
aculeate ascospores.
The closest apothecial representatives have different
living strategy also viz. fimicolous in Hapsidomyces
venezuelensis and phyllophilous in Peziza phyllogena
versus terricolous-subpsammophilous in Sarcopeziza sicula. Ascal amyloidity is also different: thin but wider and
moderately strong apical ring-like reactive zone exist in
Peziza phyllogena extending downwards with abruptly
weakened intensity (pers. data) while Hapsidomyces
venezuelensis has weakly amyloid reaction over entire
ascus length (Hansen et al. 2001). Hapsidomyces
venezuelensis has ascospores very much resembling those
of Eremiomyces (though spinulose-reticulate). While,
Peziza phyllogena has ellipsoid ascospores (as in Sarcopeziza sicula), though rather coarsely sculptured by
cyanophilic verrucae and higher tuberculae around the
polar areas (pers. data).
Class Sordariomycetes O.E. Erikss. & Winka
The classification of the families in Sordariomycetes
follow Maharachchikumbura et al. (2016), Hongsanan
et al. (2017) and Wijayawardene et al. (2017a, 2018a).
Subclass Diaporthomycetidae Senan. et al.
Atractosporales H. Zhang et al.
Conlariaceae H. Zhang et al.
Conlariaceae was introduced by Zhang et al. (2017a) in
the order Atractosporales to accommodate the freshwater
genus Conlarium, with two species, C. aquaticum W. Dong
et al. and C. duplumascosporum F. Liu & L. Cai. We
follow the latest treatment in Zhang et al. (2017a) and the
new species, C. thailandense is introduced based on morphological characteristics coupled with phylogenetic analysis of combined LSU, SSU and ITS sequence data
(Fig. 74).
Conlarium F. Liu & L. Cai
Conlarium was introduced by Liu et al. (2012a) from a
freshwater habitat in China to accommodate C.
123
Fungal Diversity
Fig. 73 Maximum likelihood phylogenetic tree based on a concatenated ITS and LSU sequence dataset. Sequences recovered during this
study are shown in blue and ex-type strains are in bold. The tree is
rooted to Ascobolus crenulatus. Bootstrap values greater than 50% are
indicated at the nodes. The bar length indicates the number of
nucleotide substitutions per site
duplumascosporum with sexual and asexual morphs (Liu
et al. 2012a). Recently, Zhang et al. (2017a) introduced the
second freshwater taxon C. aquaticum from Thailand.
Holotype: MFLU 17-1711
Saprobic on dead wood in terrestrial habitat. Sexual
morph Undetermined. Asexual morph Hyphomycetous.
Colonies sporodochial, broadly punctiform, gregarious,
raised, dark brown to black. Mycelium mostly immersed on
natural substratum, comprising branched, pale brown to
hyaline, smooth and thin-walled hyphae. Conidiophores
absent or reduced to conidiogenous cells. Conidiogenous
cells monoblastic, holoblastic, integrated, determinate,
Conlarium thailandense X.D. Yu, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555288; Facesoffungi
number: FoF04830, Fig. 75
Etymology: Named after the country where it was collected, Thailand
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Fungal Diversity
cylindrical, hyaline, smooth, up to 5.5 lm long. Conidia
25–45 9 17–33 lm (
x = 35.9 9 26.9 lm, n = 20), acrogenous, solitary, dry, mostly irregular, subglobose to ellipsoidal, brown to dark brown, clathrate, muriform, 4–8transversely septate, 4–6-longitudinally septate, slightly
constricted at the septa, smooth and thin-walled, with a
small, sub-rounded, dark brown ornamentation on the surface of each cell. Conidial secession schizolytic.
Culture characteristics: Colonies on PDA, irregular in
shape, reaching 10–20 mm in 6 weeks at room temperature, black from above and below, umbonate, rough, dense,
edge undulate.
Material examined: THAILAND, Chiang Rai Province,
Mae Fah Luang University, on dead wood, 17 July 2017, S.
Boonmee, DP6 (MFLU 17-1711, holotype), ex-type living
culture, MFLUCC 17-2349.
GenBank
numbers:
ITS = MH624129,
LSU =
MH624127, SSU = MH624128.
Notes: Conlarium thailandense is similar to the asexual
morph of C. duplumascosporum and C. aquaticum. They
all have monoblastic, holoblastic conidiogenous cells and
mostly irregular, brown, clathrate, muriform conidia (Liu
et al. 2012a). However, they can be easily distinguished by
the number of septa (0–2-transversely septate, 0–1-longitudinally septate in C. duplumascosporum, 6–12-transversely septate, 4–10-longitudinally septate in C.
aquaticum and 4–8-transversely septate, 4–6-longitudinally
septate in C. thailandense) and conidial size (15.5–35 9
11–26.5 lm in C. duplumascosporum, 45–70 9 20–57 lm
in C. aquaticum and 25–45 9 17–33 lm in C. thailandense) (Liu et al. 2012a; Zhang et al. 2017a). Apart from
number of septa and conidial size, a number of air-bubbles
were also observed on mature conidia of C. thailandense,
which have not been reported in C. aquaticum and C.
duplumascosporum. Phylogenetic analyses of a combined
LSU, SSU and ITS sequence alignment based on maximum
likelihood and Bayesian inference show that C. thailandense is sister to C. aquaticum. A comparison of ITS
pairwise indicates that C. thailandense differs from C.
aquaticum in 13 base positions and is therefore novel following the guidelines of Jeewon and Hyde (2016).
Diaporthales Nannf.
Cytosporaceae Fr.
The family Cytosporaceae was introduced by Fries
(1825) and recognized a family in Diaporthales, which
comprises phytopathogens and saprobes (Wehmeyer 1975;
Barr 1978; Eriksson 2001; Castlebury et al. 2002).
Maharachchikumbura et al. (2015, 2016) listed 13 genera
under Cytosporaceae. Senanayake et al. (2017) accepted
five genera, which belong to Cytosporaceae based on
morphological characteristics viz. Cytospora Ehrenb.,
Pachytrype Berl. ex M.E. Barr et al., Paravalsa Ananthap.,
Xenotypa Petr. and Waydora B. Sutton. Wijayawardene
et al. (2018a) listed only genera Cytospora and Waydora in
Cytosporaceae.
Cytospora Ehrenb.
Cytospora was introduced by Ehrenberg (1818) as type
genus of the family Cytosporaceae in Diaporthales
(Wehmeyer 1975; Barr 1978; Eriksson 2001; Castlebury
et al. 2002; Norphanphoun et al. 2017). The genus was
introduced as important pathogens, which cause cankers
and dieback disease on branches of a wide range of hosts
worldwide (Adams et al. 2005, 2006; Hyde et al.
2017, 2018b; Norphanphoun et al. 2017, 2018). There are
630 epithets for Cytospora (Index Fungorum 2019) with an
estimated 110 species in Kirk et al. (2008). Norphanphoun
et al. (2017) provided a comprehensive account of the
morphology and molecular background of the genus.
Several new taxa have recently been introduced (Hyde
et al. 2016, 2017, 2018b; Norphanphoun et al. 2017, 2018;
Senanayake et al. 2017; Tibpromma et al. 2017; Zhu et al.
2018). In this study, a novel species of Cytospora is
introduced from Ulmus pumila L. based on molecular data
coupled with morphological characteristics (Fig. 76).
Cytospora ulmicola Norphanphoun, Bulgakov, T.C. Wen
& K.D. Hyde, sp. nov.
Index Fungorum number: IF555487; Facesoffungi
number: FoF05184, Fig. 77
Etymology: The specific epithet ‘‘ulmicola’’ refers to the
host plant genus Ulmus, on which the fungus was first
collected.
Holotype: MFLU 17-2080
Associated with twigs and branches of Ulmus pumila L.
(Ulmaceae). Sexual morph Undetermined. Asexual
morph
Conidiostromata
1000–1700 9 500–800 lm
diam., semi-immersed in host tissue, solitary, erumpent,
scattered, discoid, circular to ovoid, with multi-loculate,
pycnidial, embedded in stromatic tissue, with ostiole. Ostioles 200–500 lm long, with an ostiolar neck. Conidiomata
walls comprising a few layers of pseudoparenchymatous
cells of textura angularis, with inner layer thin, pale brown,
outer layer brown to dark brown. Conidiophores unbranched
or occasionally branched at the base, formed from the inner
layer of the pycnidial wall. Conidiogenous cells (6–)9–
14 9 1.3–1.8(–2.8) lm (
x = 11 9 2 lm, n = 15), blastic,
enteroblastic, flask-shaped, phialidic, hyaline, and smoothwalled. Conidia (7–)8–9.5 9 1.5–1.7(–1.9) lm (
x = 8.5 9 1.6
lm, n = 30), unicellular, hyaline, oblong to allantoid,
smooth-walled.
Culture characteristics: Conidia germinating on PDA
within 12 h. Germ tubes produced from all sides. Colonies
on PDA reaching 22–27 mm diam. after 2 weeks at room
temperature, colonies irregular in shape, medium dense,
flat to slightly raised, surface slightly rough, with small
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Fungal Diversity
Fig. 74 Maximum likelihood (ML) majority rule consensus tree for
the combined LSU, SSU and ITS sequence alignment for Atractosporales. The RAxML bootstrap support values (ML) greater than
70% and Bayesian posterior probabilities (BYPP) greater than 0.95
123
BYPP are given at the nodes (ML/BYPP). The ex-type strains are in
bold and the new isolates are in blue. The tree is rooted to
Lentomitella cirrhosa (ICMP 15131)
Fungal Diversity
Fig. 75 Conlarium thailandense (MFLU 17-1711, holotype). a, b
Colonies on the substratum. c, d Conidiogenous cells. e, f Conidia.
g Conidia with the air-bubbles. h Fragile conidium. i Colonies on
PDA (from front). j Colonies on PDA (from reverse). Scale bars
a = 100 lm, c, d, f–h = 10 lm, e = 20 lm
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Fungal Diversity
granular, to velvety, edge undulate with well-defined
margin, from above pale orange to light brown at the
margin, amber near the margin, with, zonate, pale yellowish to dandelion and dark amber at the centre; from
below pale yellowish at the margin, with dark amber at the
centre; not producing pigmentation in agar.
Material examined: RUSSIA, Rostov region, Shakhty
City, street trees, on dead branch of Ulmus pumila L.
(Ulmaceae), 27 May 2017, T.S. Bulgakov, T-1778 (MFLU
17-2080, holotype; PDD, isotype), ex-type living culture,
MFLUCC 18-1227.
GenBank
numbers:
ACT = MH940216,
ITS =
MH940220, LSU = MH940218, TUB2 = MH986792.
Notes: Cytospora ulmicola (MFLU 17-2080) was found
on a dead branch (with signs of necrosis) of Ulmus pumila
in European Russia. Cytospora ulmicola resembles Cytospora species in having an ostiolar neck with multi-loculate conidiostromata and unicellular, oblong to allantoid
conidia. Phylogenetic analysis based on a combined ITS,
LSU and ACT sequence dataset indicates that C. ulmicola
clusters with C. cotini Norph. et al. (MFLUCC 14-1050),
C. ampulliformis Norph. et al. (MFLUCC 16-0583), C.
gelida Norph. et al. (MFLUCC 16-0634), and C. ceratosperma (Tode) G.C. Adams & Rossman (MFLUCC
16-0625) (Fig. 76). However, C. ulmicola can be distinguished from related species based on molecular data. A
comparison of nucleotide polymorphisms of ITS and ACT
shows that C. ulmicola differs from C. cotini with one
polymorphism of ITS; from C. ampulliformis with one
polymorphism of ITS and ten polymorphisms of ACT;
from C. gelida with four polymorphisms of ITS and 13
polymorphisms of ACT; and from C. ceratosperma with
five polymorphisms of ITS and eight polymorphisms of
ACT. Therefore, following the guidelines of Jeewon and
Hyde (2016) we introduce it as a new species.
Melanconiellaceae Senan. et al.
Melanconiellaceae was introduced by Senanayake et al.
(2017) including phytopathogenic and saprobic species to
accommodate Greeneria Scribn. & Viala, Melanconiella
Sacc. and Microascospora Senan. & K.D. Hyde in the
order Diaporthales (Senanayake et al. 2018; Braun et al.
2018). We introduce a new monotypic genus Septomelanconiella to the family Melanconiellaceae to accommodate
a single species collected on rose apple in Thailand.
Septomelanconiella Samarak. & K.D. Hyde, gen. nov.
Index Fungorum number: IF555301; Facesoffungi
number: FoF04849
Etymology: The generic epithet ‘‘Septomelanconiella’’
reflects the septation of the conidia and parallel morphology to Melanconiella.
Saprobic on Syzygium samarangense in terrestrial
habitats. Sexual morph Undetermined. Asexual morph
123
Fig. 76 Phylogram generated from maximum likelihood analysis c
based on a combined ITS, LSU, and ACT sequence dataset. The tree
is rooted to Diaporthe eres (AFTOL-ID 935). Maximum likelihood
bootstrap values C 50% are given at the nodes. Newly generated
sequence is indicated in blue. Ex-type strains are indicated in bold
Coelomycetous. Conidiomata pycnidial, immersed in the
host, partially erumpent at maturity, mostly solitary or
confluent, subglobose to irregular, to flattened and collabent, light brown. Conidiomata walls comprising 2–3
layers of hyaline cells, of textura angularis at the base,
with light brown, thin outer layer. Conidiophores mostly
reduced to conidiogenous cells, few with conidiophore and
cylindrical conidiogenous cells. Conidiogenous cells
enteroblastic, phialidic, integrated to discrete, cylindrical,
determinate, hyaline, and finely roughened. Conidia
cylindrical to clavate, one guttulate, becoming two guttules, hyaline, 1-septate when immature; mature conidia
cylindrical to clavate, straight or slightly curved, brown,
1-euseptate, more often with 6 unequal lumina, guttulate,
dark brown at the base.
Type species: Septomelanconiella thailandica Samarak.
& K.D. Hyde
Notes: Septomelanconiella thailandica constitutes an
independent lineage basal to Melanconiella syzygii Crous
& M.J. Wingf. Despite average phylogenetic support,
Septomelanconiella is distinguished from other species as
it is characterised by 1-euseptate and luminate conidia.
Septomelanconiella is similar to Melanconiella in having
finely verrucose, brown, mature conidia.
Septomelanconiella thailandica can be phylogenetically
distingushed from the other genera in Melanconiellaceae
as circumscribed by Senanayake et al. (2017). Melanconiella syzygii was isolated from a prominent leaf spot of
Syzygium and is characterized by 2–3 layers of peridium,
1–2-septate conidiophores and hyaline to light brown
aseptate conidia (Crous et al. 2016a). Multigene phylogenetic analyses also reveal that Septomelanconiella thailandica is separated from Melanconiella syzygii.
Wijayawardene et al. (2016) described Gloeocoryneum and
Stegonosperiopsis and referred them to Ascomycota, genera incertae sedis. They have similar brown, septate conidia as in Septomelanconiella thailandica. However,
Gloeocoryneum differs in having 2–5 conidial septa, while
Stegonosperiopsis is characterized by cylindrical conidiophores. Based on morphology and phylogeny in this study,
Septomelanconiella thailandica is introduced as a new
genus and species (Figs. 78, 79).
Septomelanconiella thailandica Samarak. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF555302; Facesoffungi
number: FoF04850, Fig. 79
Fungal Diversity
123
Fungal Diversity
Fig. 77 Cytospora ulmicola (MFLU 17-2080, holotype). a Herbarium
label and specimens. b, d, e Appearance of conidiostromata on
branches of Ulmus pumila. c, f Colonies on PDA (c = from above,
f = from below). g Transverse sections through conidiostroma to
show an arrangement of the locules. h Longitudinal section through
conidiostroma. i Ostiolar neck. j Conidiostroma wall. k Conidiogenous cells. l Conidia. Scale bars b = 1000 lm, e = 100 lm, g,
h = 200 lm, i = 50 lm, j = 20 lm, k, l = 10 lm
Etymology: Name based on the country from which this
species was collected, Thailand.
Holotype: MFLU 18-0793
Saprobic on recently dead twigs of Syzygium samarangense. Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata 360–500 lm diam., 130–
200 lm high, pycnidial, immersed, partially erumpent at
maturity, solitary or confluent, subglobose to irregular, to
flattened and collabent, light brown. Conidiomata walls
28–37.5 lm wide, comprising 2–3 layers of hyaline cells,
of textura angularis at the base, with light brown, thin
outer layer. Conidiophores mostly reduced to conidiogenous cells, few with conidiophore and cylindrical conidiogenous cells. Conidiogenous cells 10.3–15.9 9 4.4–
7.3 lm (
x = 13.2 9 5.9 lm, n = 20), enteroblastic, phialidic, integrated or discrete, cylindrical, determinate, hyaline, finely roughened. Conidia when immature, cylindrical
to clavate, one guttulate, becoming two guttules, hyaline,
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Fungal Diversity
one septate; mature conidia 37–52 9 15–23 lm
(
x = 43.8 9 19.1 lm, n = 40), cylindrical to clavate,
straight or slightly curved, brown, 1-euseptate, with 6
unequal lumina, guttulate, dark brown at base with opening
1.8–3.6 lm diam. (
x = 2.9 lm, n = 30).
Culture characteristics: Conidia germinating on PDA
within 24 h, germ tubes produced from central part, often
three hyphae. Colonies on PDA reaching 37 mm diam. in
2 weeks at 25 °C, hairy, white, superficial, rough surface,
irregular edge, reverse yellowish brown.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Nang Lae, on dead twigs of Syzygium
samarangense (Blume) Merr. & L.M. Perry (Myrtaceae),
25 January 2018, M.C. Samarakoon, SAMC091 (MFLU
18-0793, holotype; KUN-HKAS 102320, isotype), ex-type
living culture, MFLUCC 18-0518, ICMP.
GenBank numbers: ITS = MH727706, LSU = MH727705,
RPB2 = MH752072.
Pseudoplagiostomataceae Cheew. et al.
The monotypic family Pseudoplagiostomataceae was
introduced by Cheewangkoon et al. (2010) to accommodate a cryptosporiopsis-like fungus isolated from Eucalyptus. Pseudoplagiostomataceae resembles Gnomoniaceae G.
Winter based on morphological characters of its sexual
morph, such as solitary, immersed, lacking stromatic ascomata with lateral beaks, asci with a distinct apical ring and
1-septate ascospores (Sogonov et al. 2008; Senanayake et al.
2017). Cheewangkoon et al. (2010) showed phylogenetically
that Pseudoplagiostomataceae is closer to families with
well-developed stromatic tissue such as Diaporthaceae
Höhn. ex Wehm. and Pseudovalsaceae M.E. Barr, or families with stromatic and non-stromatic tissues such as Valsaceae Tul. & C. Tul. and Sydowiellaceae Lar.N. Vassiljeva.
However, in our phylogenetic analyses it forms a fullysupported lineage, sister to Apoharknessiaceae Senan. et al.
(Fig. 80).
Pseudoplagiostoma Cheew. et al.
We follow the latest treatment and updated accounts of
Pseudoplagiostoma in Cheewangkoon et al. (2010),
Suwannarach et al. (2016) and Du et al. (2017). Based on
phylogenetic analyses of a combined ITS, LSU, TUB2 and
TEF1-a dataset coupled with morphological characteristics, a novel species, P. mangiferae is introduced.
Pseudoplagiostoma mangiferae Dayarathne, Phookamsak
& K.D. Hyde, sp. nov.
Index Fungorum number: IF555434; Facesoffungi
number: FoF05111, Fig. 81
Etymology: The specific epithet ‘‘mangiferae’’ refers to
the host genus Mangifera, of which the species was first
collected.
Holotype: KUN-HKAS 102244
Associated with the leaf blight symptom on Mangifera
sp. having amphigenous, subcircular to irregular, medium
brown with blackish brown, reverse medium brown, surrounded by a purple-brown margin, which is dark brown in
reverse. Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata 70–140 lm high, 90–
150 lm diam. (
x = 109 9 125 lm, n = 10), medium to
dark brown, pycnidial with pale yellow drops of exuding
conidia; subglobose, subcuticular to epidermal, with central rupture, breaking through plant tissue. Conidiomata
walls 5–10 lm wide, thin-walled, composed of 2–4 layers
of yellowish brown, pseudoparenchymatous cells, of textura angularis, intermixed with the host cells at the base
and side. Paraphyses 0.8–2 lm wide, branched, septate,
hyaline. Conidiophores absent. Conidiogenous cells 5–
11 9 3.2–12.6 lm (
x = 9 9 6 lm, n = 30), proliferating
enteroblastic, appearing as phialides with periclinal thickening and collarette, or with percurrent proliferation in the
apical part; discrete, arising from the inner cell layer,
hyaline, smooth, cylindrical to ampulliform, wider at the base,
straight. Conidia 18–24 9 11–14 lm (
x = 22 9 13 lm,
n = 50), hyaline, ellipsoidal, guttulate, smooth, thick-walled,
aseptate, straight or slightly curved, frequently slightly narrow at the middle, with obtuse apex; base tapering to flat
protruding scar.
Culture characteristics: Colonies on PDA reaching
80–85 mm diam. after 4 weeks at 20–25 °C, sparse to
medium sparse, circular, flat, surface slightly rough with
tufts of hyphae, edge entire, woolly to cottony, radiating
with sparse mycelia in the middle part; from above, pinkwhite to cream, from below, pale yellowish; not producing
pigmentation in agar.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Jinghong, Nabanhe, associated with leaf
blight symtom on living leaf of Mangifera sp. (Anacardiaceae), 21 November 2015, R. Phookamsak, XB010
(KUN-HKAS 102244, holotype), ex-type living culture,
KUMCC 18-0179.
GenBank
numbers:
ITS = MK084824,
LSU =
MK084825, TEF1-a = MK084822, TUB2 = MK084823.
Notes: Morphological characteristics clearly distinguish
Pseudoplagiostoma mangiferae from other Pseudoplagiostoma species. The conidiogenous cells of P. mangiferae (5–11 9 3.2–12.6 lm) are wider than those of P.
eucalypti (6–15 9 2–6 lm) (Cheewangkoon et al. 2010).
Pseudoplagiostoma variabile is distinguished from P.
mangiferae by its subglobose to bean-shaped, variable
conidia shape (Cheewangkoon et al. 2010). The conidia of
P. corymbiae (14–19 9 7–10 lm), P. eucalypti (14–22 9
6–11 lm), P. oldie (11–20 9 6–11 lm) and P. variabile
(6.5–19 9 6.5–10.5 lm) are shorter than those of P.
mangiferae (18.8–24 9 11.3–14.2 lm) while P. dipterocarpi
(14–36 9 7–11 lm)
has
longer
conidia
123
Fungal Diversity
Fig. 78 Phylogram generated from maximum likelihood (RAxML)
based on ITS, LSU and RPB2 partial sequence data analyses of 23
taxa and Pseudoplagiostoma oldii (CBS 124808) as an outgroup
taxon. Bootstrap values for maximum parsimony (green) and
maximum likelihood (blue) C 50% and Bayesian posterior probabilities (black) C 0.90 are given at the nodes. The newly generated
sequence is in blue. The ex-type strains are in bold. The scale bar
represents the expected number of nucleotide substitutions per site
(Cheewangkoon et al. 2010; Crous et al. 2012; Suwannarach et al. 2016). Furthermore, P. oldie is distinguished
from P. dipterocarpi by its conidia turning brown when
mature (Cheewangkoon et al. 2010). According to our
multigene analyses, P. mangiferae shows a close phylogenetic relationship with P. dipterocarpi (Fig. 80). However, they form well-separated lineages with high support
(94% ML, 98% MP and 0.97 BYPP). A comparison of ITS
nucleotide bases shows that P. mangiferae differs from P.
dipterocarpi in 67/512 bp (13.1%). We therefore, introduce P. mangiferae as a new species following the guidelines of Jeewon and Hyde (2016).
The genus Coniella was introduced by von Höhnel
(1918) and is typified by Coniella pulchella Höhn. (= C.
fragariae (Oudem.) B. Sutton). This genus comprises
endophytes, saprobes and plant pathogens (Van Niekerk
et al. 2004; Alvarez et al. 2016; Chethana et al. 2017). A
new host record collected on Prunus armeniaca L. from
Russia is reported for C. vitis, which has been previously
reported as a plant pathogen from Vitis vinifera L. (Vitaceae; Chethana et al. 2017) (Fig. 82).
Schizoparmaceae Rossman
Schizoparmaceae was introduced by Rossman et al.
(2007) and is typified by Schizoparme Shear. The family
also includes Pilidiella Petr. & Syd., the asexual morph of
Schizoparme and the closely related Coniella Höhn
(Rossman et al. 2007). In a recent revision, these three
genera were synonymized under the old name, Coniella
(Alvarez et al. 2016).
Coniella Höhn.
123
Coniella vitis Chethana, J.Y. Yan, X.H. Li & K.D. Hyde,
Pl. Dis. 101: 2129 (2017).
Facesoffungi number: FoF02722, Fig. 83
Holotype: CHINA, Yanqing, Beijing, on white rot-infected berries of Vitis vinifera (Vitaceae), 13 May 2015, X–
H Li, JZB3700001 (MFLU 16-2677), ex-type living culture, MFLUCC 16-1399.
Saprobic or pathogenic on branches and twigs of Prunus
armeniaca L. (Rosaceae). Sexual morph Undetermined.
Asexual morph Coelomycetous. Conidiomata 200–
370 lm diam. (
x = 304.2 lm, n = 20), pycnidial, solitary,
submerged in PDA, globose to slightly depressed globose,
with verruculose wall, initially hyaline becoming dark
brown to black at maturity, with a central ostiole.
Fungal Diversity
Fig. 79 Septomelanconiella thailandica (MFLU 18-0793, holotype).
a Host. b Conidiomata on the substrate. c Vertical section of
conidioma. d–h Conidiophores, conidiogenous cells and developing
conidia. i–l Conidia. m Germinated conidium. n Culture on PDA
from above after 16 days. o Culture on PDA from below after
16 days. Scale bars b = 1000 lm, c = 100 lm, m = 50 lm, d–
l = 20 lm
Conidiophores
5–9 9 2–4 lm
(
x = 7.8 9 3.3 lm,
n = 10), formed on a dense, cushion-like aggregation of
hyaline cells, subcylindrical, hyaline, smooth, simple or
branched below, mostly reduced to conidiogenous cells.
123
Fungal Diversity
Fig. 80 Phylogenetic tree generated by maximum likelihood analysis
of a combined ITS, LSU, TUB2 and TEF1-a sequence dataset of
Pseudoplagiostomataceae and related families. Related sequences
were obtained from GenBank. Thirty one strains are included in the
analyses and the tree is rooted with Phaeoacremonium novaezealandiae (CBS 110156) and P. hungaricum (CBS 123036). Tree
topology of the ML analysis was similar to the MP and BI. The best
scoring RAxML tree with a final likelihood value of -15510.220382 is
presented. The matrix had 1138 distinct alignment patterns, with
50.26% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.226739, C = 0.283205, G = 0.257579,
T = 0.232477; substitution rates AC = 1.368247, AG = 2.869863,
AT = 1.349096, CG = 0.967468, CT = 5.138587, GT = 1.000000;
gamma distribution shape parameter a = 0.391573. RAxML (black)
and maximum parsimony (blue) bootstrap support values C 60% are
shown respectively above the nodes. Bayesian posterior probabilities C 0.95 BYPP indicated in green. The scale bar indicates 0.06
changes. The ex-type strains are in bold and new isolates in blue bold
Conidiogenous cells 9.5–16 9 2.5–4 lm (
x = 13.2 9 3.3
lm, n = 10), phialidic, percurrently proliferating, hyaline,
simple, slender, smooth with a prominent periclinal thickening. Conidia 8.5–12 9 3.5–6.5 lm (
x = 10.2 9 4.9 lm,
n = 40), l:w ratio 2, inequilateral, hyaline when immature
becoming pale brown, aseptate, straight to slightly curved,
narrowly ellipsoidal, often somewhat flattened on one side,
both sides gradually tapering towards the subobtusely rounded
apex, subtruncate base, smooth-walled, and multi-guttulate
with one or two prominent guttules.
Culture characteristics: Colonies on PDA, reaching
8 cm diam. after 5 days at 28 °C, effuse, flat, mostly
immersed mycelium, aerial mycelium mostly sparse,
crenulated edges with concentric rings, white on surface
and buff in reverse. Conidia in mass, hyaline.
Material examined: RUSSIA, Rostov Region, Shakhty
City, on dead twigs (with signs of necrosis) of Prunus
armeniaca L. (Rosaceae), 1 March 2016, T.S. Bulgakov,
T1243 (MFLU 16-1537), living culture, MFLUCC 18-0093.
GenBank numbers: ITS = MH569466, LSU = MH569461,
HIS3 = MH645901, TEF1-a = MH645902.
Known hosts and distribution: Vitis vinifera (China) and
Prunus armeniaca (Russia) (Chethana et al. 2017; Farr and
Rossman 2018).
Notes: Coniella vitis has been reported from Vitis vinifera as causing grape white rot in China (Chethana et al.
2017; Jayawardena et al. 2018). Based on our phylogenetic
analysis of a combined ITS, LSU, HIS3 and TEF1-a
sequence dataset of Coniella species (Fig. 82), our strain
MFLUCC 18-0093 clusters together with the ex-type strain
of Coniella vitis (MFLUCC 16-1399) with high bootstrap
and Bayesian probabilities (100% MP and 1.00 BYPP).
When comparing our strain with the type specimen of C.
vitis (MFLUCC 16-1399), they are similar in morphology.
However, our strain has slightly larger conidiomata, and
123
Fungal Diversity
Fig. 81 Pseudoplagiostoma mangiferae (KUN-HKAS 102244, holotype). a Leaf blight symptom on living leaf of Mangifera sp.
b Conidiomata on host surface. c, d Section through conidiomata. e, f
Conidiomata walls at the side and the base respectively. g Paraphyses.
h–k Conidiogenous cells. l–n Conidia. Scale bars c–e = 50 lm, f, g,
l = 20 lm, h–k, m, n = 10 lm
123
Fungal Diversity
larger conidiogenous cells compared to the type strain
(Chethana et al. 2017).
Diaporthomycetidae, families incertae sedis
Distoseptisporaceae K.D. Hyde & McKenzie
Distoseptisporaceae was introduced by Su et al. (2016)
and classified in Diaporthomycetidae, families incertae
sedis by Wijayawardene et al. (2018a). The family consists
of one hyphomycetous genus, Distoseptispora which is
typified by D. fluminicola McKenzie et al. Seventeen
Distoseptispora species have been introduced through
previous studies, out of which eleven species are from
freshwater, and five species are from terrestrial habitats
(Hyde et al. 2016; Su et al. 2016; Xia et al. 2017; Luo et al.
2018; Tibpromma et al. 2018; Yang et al. 2018c) (Fig. 84).
Distoseptispora thysanolaenae Goonas., Dayarathne,
Phookamsak & K.D. Hyde, sp. nov.
Index Fungorum number: IF555408; Facesoffungi
number: FoF05011, Fig. 85
Etymology: Refers to the host from which the species
was isolated
Holotype: KUN-HKAS 102247
Saprobic on dead culms of Thysanolaena maxima.
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Colonies effuse, brown to dark brown,
hairy or fluffy, arising from subiculum, with tufts. Mycelium partly superficial, composed of branched, septate,
smooth, brown hyphae. Conidiophores 30–80 9 3.5–
5.5 lm (
x = 52 9 4.5 lm, n = 30), macronematous,
mononematous, light to dark brown, 2–8-septate, smooth,
usually flexuous or sometimes straight, unbranched,
cylindrical, rounded at the apex. Conidiogenous cells
monoblastic, integrated, terminal, determinate, hyaline to
light brown, cylindrical. Conidia 21.5–80 9 6.5–12.8 lm
(
x = 53 9 9.5 lm, n = 30), acrogenous, solitary, narrow
and elongated obclavate, slightly curved, 8–14-distoseptate, thick-walled, light to dark brown, paler at the apex,
tapering towards a rounded, flat apex, truncate with flat
base, with conspicuous spore attachment loci, guttulate,
smooth-walled.
Culture characteristics: Colonies on PDA reaching 43–
45 mm diam. after 4 weeks at 20–25 °C, colonies dense,
circular, flat, surface slightly rough, rugose with edge
entire, floccose to velvety; colony from above, greenish
grey at the margin, brown-grey at the centre, from below
dark grey at the margin, black at the centre; not producing
pigmentation in PDA. Sporulation on PDA after two
months. Conidiophores 10–30(–47) 9 3–5 lm (
x = 21.4 9 4.6
lm, n = 25), macronematous, micronematous, mononematous, light to dark brown, 1–3-septate, smooth, usually
flexuous or sometimes straight, unbranched, cylindrical,
rounded at the apex. Conidiogenous cells monoblastic,
123
Fig. 82 Phylogenetic tree generated by maximum parsimony analysis c
of combined ITS, LSU, HIS3 and TEF1-a sequence dataset of
Coniella species. Related sequences were obtained from GenBank.
Forty four strains are included in the analyses, which comprise 2881
characters including gaps. Single gene analyses were carried out and
compared with each species, to compare the topology of the tree and
clade stability. The tree is rooted with Melanconiella sp. (CBS
110385). The maximum parsimonious dataset consisted of 2109
constant, 558 parsimony-informative and 214 parsimony-uninformative characters. The parsimony analysis of the data matrix resulted in
the maximum of two equally most parsimonious trees with a length of
2551 steps (CI = 0.509, RI = 0.708, RC = 0.360, HI = 0.491) in the
first tree. Maximum parsimony bootstrap support values C 70% are
shown near the nodes. Bayesian posterior probabilities C 0.95 BYPP
indicated as thickened black branches. The scale bar indicates 10
changes. The ex-type strains are in bold
integrated, terminal, determinate, hyaline to light brown,
cylindrical. Conidia (20–)30–70(–243) 9 5–8(–11) lm
(
x = 53 9 8.4 lm, n = 30), acrogenous, varied in shape,
elongate cylindrical to obclavate, lanceolate, rostrate,
6–22(–50)-distoseptate, light brown to dark brown, tapering
towards a rounded, sometimes bulbous apex, truncate at the
base, with conspicuous spore attachment loci, smoothwalled.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Mengla County, Xishuangbanna Tropical
Botanical Garden (XTBG), on dead culms of Thysanolaena
maxima (Roxb.) Kuntze (Poaceae), 22 April 2017, R.
Phookamsak, IS004 (KUN-HKAS 102247, holotype), extype living culture, KUMCC 18-0182 (IS004A), KUMCC
18-0183 (IS004B).
GenBank
numbers:
ITS = MK045851,
LSU =
MK064091, TEF1-a = MK086031.
Notes: Distoseptispora thysanolaenae is the sixth species in this genus to be introduced from a terrestrial habitat.
The previously introduced species are D. martini (J.L.
Crane & Dumont) J.W. Xia & X.G. Zhang, D. tectonae
Doilom & K.D. Hyde, D. tectonigena Doilom & K.D.
Hyde, D. thailandica Tibpromma & K.D. Hyde and D.
xishuangbannaensis Tibpromma & K.D. Hyde (Hyde et al.
2016; Xia et al. 2017; Luo et al. 2018; Tibpromma et al.
2018). Distoseptispora thysanolaenae can be distinguished
from these species by its 2–8-septate conidiophore that are
30–80 9 3.5–5.5 lm and narrow elongated, obclavate,
light to dark brown conidia that are 21.5–80 9
6.5–12.8 lm, 8–14-distoseptate and smooth-walled. Distoseptispora martini has longer conidiophores (50–110 9
3.5–4.5 lm), shorter and wider conidia (15–20 9
11–16 lm) that are ellipsoid, oblate or subglobose (Xia
et al. 2017).
Distoseptispora tectonae and D. tectonigena differ in
their conidiophore dimensions (up to 40 9 4–6 lm, and up
to 110 9 5–11 lm, respectively) and conidial dimensions
Fungal Diversity
123
Fungal Diversity
Fig. 83 Coniella vitis (MFLUCC 18-0093). a Host surface from
which the saprobe was isolated. b Immature white pycnidia on PDA.
c Mature black pycnidia with hyaline spore mass on PDA. d Conidiogenous cells. e Periclinal thickening at the apex of the
conidiogenous cells. f–i Different shapes of conidia. j Upper-view
(right) and the reverse view (left) of the colony on PDA. Scale bars
a = 2 mm, c = 500 lm, d, e = 10 lm, f–i = 5 lm
((90–)130–140(–170) 9 13–14 lm, and 148–225(–360) 9
11–12 lm, respectively) and number of septa. Distoseptispora thailandica and D. xishuangbannaensis have
larger conidia (130–230 9 13.5–17 lm and 160–305 9
8–15 lm, respectively) and more septa (35–52-distoseptate, and up to 40-distoseptate, respectively) (Tibpromma
et al. 2018).
Phylogenetic analyses based on a combined ITS, LSU
and TEF1-a sequence dataset show that our taxon clusters
with other Distoseptispora species (Fig. 84). The species is
sister to D. guttulata J. Yang & K.D. Hyde (MFLUCC
16-0183). A comparison of ITS and TEF1-a nucleotide
bases shows that D. thysanolaenae differs from D. guttulata with 59 nucleotide bases of ITS and 64 nucleotide
bases of TEF1-a. Distoseptispora thysanolaenae shares
similar morphological features with D. guttulata but can be
distinguished from the latter in having shorter conidia
(21.5–80 lm versus 75–130 lm; Yang et al. 2018c).
Therefore, following the guidelines of Jeewon and Hyde
(2016) we introduce it as a new species.
Diaporthomycetidae, genera incertae sedis
123
Proliferophorum G.N. Wang, H. Zhang, K.D. Hyde &
Senan., gen. nov.
Index Fungorum number: IF555401; Facesoffungi
number: FoF04847
Etymology: The generic epithet ‘‘Proliferophorum’’
refers to the proliferation of conidiophores.
Saprobic on decaying, submerged wood in freshwater
habitat. Sexual morph Undetermined. Asexual morph
Hyphomycetous. Colonies irregular, hairy, black, gregarious. Mycelium mostly immersed in substratum, consisting
of branched, septate, subhyaline to pale brown, smooth
hyphae. Conidiophores macronematous, mononematous,
caespitose, cylindrical, unbranched, erect, straight and
curved at the upper part, dark brown, light brown at the
apex, 8–10-septate, not constricted at the septa, smooth,
sometimes percurrently proliferating 1–2 times at broken
ends, with few upper cells, guttulate. Conidiogenous cells
holoblastic, polyblastic, terminal, sympodial, pale brown or
subhyaline, with minute, truncate conidiogenous loci.
Fungal Diversity
Fig. 84 Phylogenetic tree generated by maximum likelihood analysis
of a combined ITS, LSU and TEF1-a sequence dataset of Distoseptisporaceae and other related families in Diaporthomycetidae. Related
sequences were obtained from GenBank. Fifty one strains are
included in the analyses, which comprised 2409 characters including
gaps. Single gene analyses were carried out and compared with each
species, to compare the topology of the tree and clade stability. The
tree is rooted with Sordaria fimicola (SMH 4106, FGSC 2918). Tree
topology of the ML analysis was similar to the MP and BI. The best
scoring RAxML tree with a final likelihood value of -17019.410141 is
presented. The matrix had 1186 distinct alignment patterns, with
38.62% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.237054, C = 0.262859, G = 0.290202,
T = 0.209884; substitution rates AC = 1.172995, AG = 2.180726,
AT = 1.594744, CG = 1.095103, CT = 6.416481, GT = 1.000000;
gamma distribution shape parameter a = 0.352856. The maximum
parsimonious dataset consisted of 1183 constant characters, 751
parsimony-informative and 475 parsimony-uninformative characters.
The parsimony analysis of the data matrix resulted in the maximum of
two equally most parsimonious trees with a length of 2985 steps
(CI = 0.590, RI = 0.726, RC = 0.428, HI = 0.410) in the first tree.
RAxML (black) and maximum parsimony (blue) bootstrap support
values C 60% are shown respectively above the nodes. Bayesian
posterior probabilities C 0.95 BYPP indicated in green. The scale bar
indicates 0.1 changes. The ex-type strains are in bold and a new
isolate is in blue
Conidia fusiform to cylindrical, aseptate when young, 2–3septate when mature, rarely up to 4-septate, slightly constricted at the septa, dark brown at central cells, pale brown
at end cells, guttulate at some stage, dry, smooth.
Type species: Proliferophorum thailandicum G.N.
Wang, H. Zhang, K.D. Hyde & Senan.
Notes: Proliferophorum is introduced as a monotypic
genus in the subclass Diaporthomycetidae to accommodate
a hyphomycetous species forming mononematous, caespitose conidiophores, sometimes percurrently proliferating
1–2 times at broken ends of conidiogenous cells and fusiform to cylindrical conidia. The genus was found on wood,
123
Fungal Diversity
123
Fungal Diversity
b Fig. 85 Distoseptispora thysanolaenae (KUN-HKAS 102247, holo-
type). On host substrate: a Appearance of colonies on host surface. b,
d, e Conidiophores. c, f Conidiophores with attached conidia. g–
i Variable shapes of conidia. In vitro: j Sporulation of conidia on PDA
after 4 weeks. k–m Conidiophores with attached conidia. n–s Variable
shapes of conidia. Scale bars a = 500 lm, b–i, k–r = 20 lm,
s = 50 lm
similar to the genus Minimelanolocus R.F. Castañeda &
Heredia, which has conidiophores that are generally conspicuous, mononematous, solitary or fasciculate, septate,
erect, straight or flexuous, smooth or verrucose, and brown
to dark brown with a melanised base (Castañeda-Ruiz et al.
2003). The conidia of Minimelanolocus are generally,
cylindrical, naviculate, clavate, obclavate and pale brown
to dark brown (Liu et al. 2015b). However, Proliferophorum differs from Minimelanolocus in its conidiophores that
have a few upper guttulate cells and conidia having obvious droplets. In addition, LSU sequence data analysis
shows the genus does not group with species in
Chaetothyriales. Phylogenetic analyses (Fig. 86) of taxa
within Diaporthomycetidae indicate that Proliferophorum
forms a single lineage, between Phomatosporales Senan.
et al. and Amplistromatales M.J. D’souza et al. The sporothrix-like asexual morph of Phomatosporales was reported from culture by Rappaz (1992).
Amplistromatales comprises the families Amplistromataceae Huhndorf et al. and Catabotrydaceae Petr. ex M.E.
Barr and their asexual morphs have been reported as
acrodontium-like for Amplistroma Huhndorf et al. (Maharachchikumbura et al. 2015). Proliferophorum and
acrodontium-like asexual morphs are distinct.
Proliferophorum thailandicum G.N. Wang, H. Zhang,
K.D. Hyde & Senan., sp. nov.
Index Fungorum number: IF555400; Facesoffungi
number: FoF04848, Fig. 87
Etymology: In reference to Thailand, where the species
was collected.
Holotype: MFLU 17-1054
Saprobic on decaying, submerged wood in freshwater
habitat. Sexual morph Undetermined. Asexual morph
Colonies irregular, hairy, black, gregarious. Mycelium
mostly immersed in substratum, consisting of branched,
septate, subhyaline to pale brown, smooth hyphae. Conidiophores macronematous, mononematous, caespitose,
cylindrical, unbranched, erect, straight and curved at the
upper part, dark brown, light brown at the apex, 8–10septate, not constricted at the septa, 50–100 9 4–7 lm
(
x = 82 9 5.5 lm, n = 10), smooth, sometimes percurrently proliferating 1–2 times at broken ends, with few
upper cells guttulate. Conidiogenous cells holoblastic,
polyblastic, terminal, sympodial, pale brown or subhyaline,
up to 20–30 lm long, with minute, truncate conidiogenous
loci. Conidia fusiform to cylindrical, aseptate when young,
2–3-septate when mature, rarely up to 4-septate, slightly
constricted at the septa, dark brown at central cells, pale
brown at end cells, guttulate at some stage, dry, smooth,
15–25 9 3–7 lm (
x = 23 9 5 lm, n = 20).
Culture characteristics: Colonies on PDA reaching
10 mm diam. within 15 days at 25 °C, colony circular,
medium dense slightly raised to umbonate, surface slightly
rough with edge entire, floccose to velvety, colony from
above and below black; not producing pigmentation in
agar.
Material examined: THAILAND, Chiang Rai Province,
Longkhot Subdistrict, saprobic on decaying submerged
wood in a stream, 1 September 2017, G.N Wang, 4.14.1
(MFLU 17-1054, holotype), ex-type living culture,
MFLUCC 17-0293.
GenBank
numbers:
ITS = MK028344,
LSU =
MK028343, SSU = MK028345.
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Glomerellales Chadef. ex Réblová et al.
Plectosphaerellaceae W. Gams et al.
Plectosphaerellaceae was introduced by Zare et al.
(2007) to accommodate the genera Acrostalagmus Corda,
Gibellulopsis Bat. & Maia, Musicillium Zare & W. Gams,
Plectosphaerella and Verticillium Nees, with Plectosphaerella as the type genus and P. cucumeris Kleb.
being the type species. Eleven genera are accepted in this
family viz. Acrostalagmus, Brunneomyces A. Giraldo
et al., Chordomyces Bilanenko et al., Gibellulopsis, Lectera
P.F. Cannon, Longitudinalis Tibpromma & K.D. Hyde,
Musicillium, Plectosphaerella, Sodiomyces A.A. GrumGrzhim. et al., Stachylidium Link and Verticillium (Wijayawardene et al. 2018a).
Plectosphaerella Kleb.
We follow the latest treatment and updated accounts of
Plectosphaerella in Zare et al. (2007), Carlucci et al.
(2012), Hyde et al. (2017) and Su et al. (2017), with the
updated phylogenetic analyses based on a combined ITS,
LSU and TEF1-a which were retrieved from Hyde et al.
(2017) and Su et al. (2017). A novel species, Plectosphaerella kunmingensis is introduced based on its
holomorphic characteristics and phylogenetic affinities in
Plectosphaerellaceae (Fig. 88)
Plectosphaerella kunmingensis Phookamsak, J.F. Li &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556180; Facesoffungi
number: FoF05716, Fig. 89
123
Fungal Diversity
123
Fungal Diversity
b Fig. 86 Phylogram inferred from maximum likelihood analysis of a
combined LSU, SSU and ITS sequence dataset using a GTRGAMMA
model of evolution. Maximum likelihood bootstrap support greater
than 80% and Bayesian posterior probability greater than 0.80 BYPP
are indicated at the nodes. Newly introduced strain is in blue bold and
type strains are in bold. The tree is rooted to Leotia lubrica (AFTOLID 1 = OSC 100001). The combined LSU, SSU and ITS sequence
dataset comprised 67 strains (including the new strain and outgroup
taxon) and manually adjusted dataset totally comprised 2228 characters including gaps. The best scoring RAxML tree was selected to
represent the relationships among taxa, in which a final likelihood
value of - 22073.200898
Etymology: The specific epithet ‘‘kunmingensis’’ refers
to Kunming City, Yunnan, China where the species was
first collected.
Holotype: KUN-HKAS 102246
Colonies forming on PDA. Mycelium composed of 1–
3 lm wide, septate, branched, smooth and thin-walled,
hyaline hyphae, partly immersed on PDA. Sexual morph
forming black, obpyriform ascomata after 8 weeks at 20–
25 °C. Ascomata 100–185 lm high, 80–110 lm diam.,
perithecial, superficial or immersed in culture colonies,
scattered, solitary to gregarious, subglobose to ovoid, or
obpyriform, uni-loculate, glabrous, ostiole at centre, with a
minute papilla. Peridium 7–18 lm wide, thin-walled, of
equal thickness, composed of 1–3 layers, of dark brown
pseudoparenchymatous cells, arranged in a textura angularis.
Asci (42–)45–55(–62) 9 10–13 lm (
x = 51.9 9 11.6 lm,
n = 15), 8-spored, unitunicate, cylindric-obclavate to
obpyriform, subsessile, thick-walled at apex, with J-, subapical ring. Ascospores (8–)10–13(–15) 9 3–5 lm
(
x = 11.9 9 4.3 lm, n = 50), overlapping 1–2-seriate, hyaline, ellipsoidal to subfusoid, with round ends, 1-septate,
smooth-walled with small guttules, becoming finely echinulate when stained by Melzer’s reagent. Asexual morph
Hyphomycetous, Conidiophores 15–50(–58) 9 (1.5–)3–
5 lm (
x = 38.5 9 4.2 lm, n = 20), macronematous or
micronematous, produced from prostrate hyphae, lacking
hyphal coils, sometimes branched, hyaline, yellowish brown
at the base, thick-walled, smooth, septate, branched, straight
or flexuous. Conidiogenous cells phialidic, determinate,
integrated to discrete, hyaline, subcylindrical to ampulliform, smooth-walled, with periclinal wall thickening, with
minute
collarette.
Conidia
7–10 9 4–7 lm
(
x = 8.7 9 5.4 lm, n = 50), pleurogenous or acropleurogenous, formed in slimy heads at the apex of the phialides,
hyaline, subglobose to ellipsoidal, aseptate, smooth, thinwalled, with granular contents. Chlamydospores not
produced.
Culture characteristics: Colonies on PDA reaching
40–48 mm diam. after 3 weeks at 20–25 °C, dense, irregular in shape, flat to slightly raised, slightly rough at
surface, with mycelia radiating outwards, sometimes
forming sectors of different folds, fimbriate at edge,
mucoid to floccose; from above, white yellowish to cream;
from below, pale yellowish; not producing pigmentation in
agar medium.
Material examined: CHINA, Yunnan Province, Kunming City, Kunming Institute of Botany, colonies forming
on WA as a contaminated fungus, 5 July 2017, R.
Phookamsak, KIB042 (dried culture herbarium: KUNHKAS 102246, holotype), ex-type living culture, KUMCC
18-0181.
GenBank
numbers:
ITS = MH254296,
LSU =
MH254298, RPB2 = MH254297, TEF1-a = MH254295.
Notes: Phylogenetic analyses of a combined ITS, LSU
and TEF1-a sequence dataset (Fig. 88) show the new
species, Plectosphaerella kunmingensis forming a distinct
lineage at the basal clade of Plectosphaerella in Plectosphaerellaceae with moderate support (76% ML and
0.96 BYPP). Plectosphaerella kunmingensis can be distinguished from other Plectosphaerella species in the lack
of hyphal coils, as well as having partly coloured conidiophores. The sexual morph of P. kunmingensis differs
from P. cucumerina in having smaller asci (P. kunmingensis, (42–)45–55(–62) 9 10–13 lm versus 50–80 9
6–9 lm, P. cucumerina; Carlucci et al. 2012) and wider
ascospores (P. kunmingensis, (8–)10–13(–15) 9 3–5 lm
versus (9–)10.5–14(–15) 9 2.5–3(–4) lm, P. cucumerina;
Carlucci et al. 2012) (Fig. 90).
Another species, Plectosphaerella sinensis was also
found from China isolated from a healthy stem of Cucumis
melo L. (Cucurbitaceae) (Su et al. 2017). Whereas, our
isolate was found as a contaminated fungus on WA.
Plectosphaerella kunmingensis has subglobose to ellipsoidal, aseptate conidia and lacks chlamydospores, while P.
sinensis has ellipsoidal, 0–1-septate conidia with a slightly
apiculate base, and forming intercalary or terminal, irregular, thick-walled chlamydospores (Su et al. 2017). In the
BLASTn search on NCBI GenBank, the closest matches of
ITS sequence of P. kunmingensis are P. nepalense (CBS
971.72) and P. cucumerina (XSD-75) with 95% similarities
for both strains. In addition, a comparison of ITS nucleotide bases between P. kunmingensis and P. niemeijerarum
(CBS 143233) shows that they differ in 42 base positions
(8.08%/520 bp). Based on a molecular analyses coupled
with morphological characteristics, we therefore, propose
P. kunmingensis as a new species.
Hypocreales Lindau
Cordycipitaceae Kreisel ex G.H. Sung et al.
Cordycipitaceae was validated by Sung et al. (2007) to
accommodate the type of Cordyceps Fr., C. militaris (L.)
Fr. and most of Cordyceps species forming brightly
coloured, fleshy stromata. Species of Cordycipitaceae have
123
Fungal Diversity
Fig. 87 Proliferophorum thailandicum (MFLU 17-1054, holotype).
a Colonies on submerged wood. b–h Conidiophores. i–l Conidia.
m Germinated conidium. n Colony on PDA (from above). o Colony
123
on PDA (from below). Scale bars a = 200 lm, b–g = 20 lm,
h = 10 lm, i–l = 10 lm
Fungal Diversity
been reported as obligate saprotrophs, parasites and symbionts with insects and fungi or grasses, rushes or sedges
(Sung et al. 2007; Schardl et al. 2013; Kepler et al.
2013, 2017; Li et al. 2016; Tibpromma et al. 2017; Huang
et al. 2018). Eighteen genera are listed in this family
(Wijayawardene et al. 2018a). We follow the latest treatment and updated accounts of Cordycipitaceae in Kepler
et al. (2017).
Leptobacillium Zare & W. Gams
The genus Leptobacillium was introduced by Zare and
Gams (2016) to accommodate saprotrophic and fungicolous phialidic hyphomycetes and Leptobacillium leptobactrum (W. Gams) Zare & W. Gams is the type species.
We follow the latest treatment in Zare and Gams (2016).
Leptobacillium leptobactrum var. calidius is reported for
the first time in India.
Leptobacillium leptobactrum var. calidius Zare & W.
Gams, Mycol Progr 15: 1003 (2016)
Facesoffungi number: FoF04832, Fig. 91
Holotype: GHANA, Atewa. CBS H-22406, a dried
culture of CBS 748.73 (= IHEM 3708), from living lepidopteran larva collected by H.C. Evans.
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978)
Saprobic on phylloplane. Sexual morph Undetermined.
Asexual morph Conidiophores 36–173.5 9 1.2–2.8 lm
(
x = 97 9 1.5 lm, n = 30), cylindrical, mainly solitary or
in groups (2–3), simple to branched, smooth-walled, hyaline.
Conidiogenous cells 3.5–66.5 9 0.8–2 lm (
x = 29.5 9
1.4 lm, n = 30), phialidic, cylindrical, septate, slightly
tapering towards apex, 1–3 arising from single node of
condiophores. Conidia 2.5–6.2 9 0.8–1.8 lm (
x = 4 9 1.2
lm, n = 30), produced in long chains (about 20 in a chain)
and in gloeosporic masses, cylindrical, bacilliform, fusiform,
smooth-walled, aseptate, hyaline. Chlamydospores absent.
Culture characteristics: Colonies on PDA reaching
15–28 mm diam. in 10 days, at 25 °C, white (4A1), margin
smooth, entire, flat to slightly raised, puffy, sometimes
produces exudates. Reverse yellowish white (4A2) to
crème (4A3), wrinkled.
Known hosts and distribution: Hemileia vastatrix Berk.
& Broome on Coffee (Brazil), technical equipment
(France, Netherlands), living lepidopteran larva (Ghana),
decaying wood (Poland), cyst of Heterodera glycines
Ichinohe (USA) (Zare and Gams 2016).
Material examined: INDIA, Maharashtra, Pune, on
phylloplane of Colocasia esculenta (L.) Schott (Araceae),
10 November 2017, S.K. Singh, living culture, NFCCI
4235, voucher specimen, AMH 10000.
GenBank numbers: ITS = MG786580, LSU = MG786581.
Notes: Phylogenetic analysis based on a combined ITS
and LSU sequence dataset of 24 taxa shows that the Indian
taxon (NFCCI 4235) clusters with type of Leptobacillium
leptobactrum var. calidius Zare & W. Gams (CBS 748.73)
(Fig. 90). Mega blast search of ITS sequence shows 99%
similarity (505/508) with L. leptobactrum var. calidius
(CBS 748.73) and same similarity (99% similarity,
592/594) for LSU analysis. In this isolate conidia are
produced in long chains and in gloeosporic masses. Though
slight variations in conidial length and width were
observed, our isolate is similar to L. leptobactrum var.
calidius (CBS 748.73) in overall morphological characteristics. In addition, our isolate optimally grows at
25–26 °C, the same as reported for L. leptobactrum var.
calidius (CBS 748.73) with negligible growth at 30 °C.
Optimum temperature for growth of L. leptobactrum var.
leptobactrum is 18–21 °C (Zare and Gams 2016). Leptobacillium leptobactrum var. calidius and L. leptobactrum
var. leptobactrum were treated as a synonym of L. leptobactrum (Index Fungorum 2019). However, L. leptobactrum var. calidius (CBS 748.73) forms a distinct lineage
with the type strain of L. leptobactrum (CBS 771.69).
Hence, based on similarity in morphological characteristics, temperature requirements, and phylogenetic analysis
this Indian isolate is identified as L. leptobactrum var.
calidius (Zare and Gams 2016). To our knowledge, this is
the first report of this genus, species and variety isolated
from phylloplane of a different host, Colocasia esculenta
from India.
Hypocreaceae De Not.
Hypocreaceae was introduced by De Notaris (1844) and
is typified by Hypocrea Fr. Species of Hypocreaceae are
characterized by their brightly coloured, fleshy perithecial
stromata (Rogerson 1970). The family comprises 18 genera
and more than 450 species (Kirk et al. 2008; Wijayawardene et al. 2018a).
Trichoderma Pers.
Trichoderma is frequently isolated from various habitats
including soil, decaying wood, and vegetable matter (Samuels 2006) and is known to play an important ecological
role as biocontrol agents of plant diseases, producers of
bioactive compounds, and pathogens of animals and
mushrooms (Schuster and Schmoll 2010). Based on
molecular data, more than 250 species are currently
accepted in this genus (Bissett et al. 2015). Three novel
species are introduced based upon their phylogenetic
affinities clarified by maximum likelihood and Bayesian
inference analyses of a combined TEF1-a and RPB2
sequence dataset (Fig. 92).
123
Fungal Diversity
Fig. 88 Phylogenetic tree generated from maximum likelihood analysis based on a combined LSU, ITS and TEF1-a sequence dataset of
the genera in Plectosphaerellaceae. Bootstrap support value of
maximum likelihood (left) equal to or greater than 70% and Bayesian
posterior probability equal to or greater than 0.95 BYPP are indicated
above or below the nodes. Type strains are in black bold. The new
species are in blue bold. The tree is rooted with Reticulascus clavatus
(CBS 125296)
Trichoderma koreanum S-Y. Oh, M.S. Park & Y.W. Lim,
sp. nov.
MycoBank number: MB824661; Facesoffungi number:
FoF04459, Fig. 93
Etymology: The specific epithet ‘‘koreanum’’ refers to
the type locality.
Holotype: SFC20131005-S066.
123
Fungal Diversity
Fig. 89 Plectosphaerella kunmingensis (KUN-HKAS 102246, holotype). a, b Culture characteristics on PDA (a = from above, b = from
below). c Ascomata forming on PDA after 2 months. d Squash mount
of the ascoma. e Section through ascoma. e, f Section through
peridium. g–j Ascospores. k Ascospores stained in Melzer’s reagent.
l Asci. m–p Conidiophores attached with conidia. q Conidia. Scale
bars d, e = 50 lm, l = 20 lm, f, m–p, q = 10 lm, g–k = 5 lm
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1963).
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Conidiophores branched at an acute or
right angle to the main axis, mostly unpaired. Phialides
123
Fungal Diversity
Fig. 90 Phylogram generated
from maximum parsimony
based on a combined ITS and
LSU sequence dataset of the
genus Leptobacillium and
related genera in
Cordycipitaceae. Bootstrap
support value of maximum
likelihood (left) and maximum
parsimony (right) equal to or
greater than 70% are indicated
above the nodes. Type strains
are in bold. The new strain is in
blue bold. The tree is rooted
with Gibellulopsis piscis (CBS
892.70)
8.6–13.4 9 2.4–3.7 lm, l/w 2.5–4.9, 1.7–2.6 lm wide at
the base (n = 30), typically formed in whorls of 3–5, subulate or lageniform. Conidia smooth, subglobose to ellipsoid,
3.1–4.4 9 2.6–3.3 lm, l/w 1.1–1.4(-1.5) (n = 30).
Culture characteristics: On CMD after 72 h colony
radius 17–22 mm at 15 °C, 29–35 mm at 20 °C,
44–49 mm at 25 °C, 20–29 mm at 30 °C; colonies on
CMD fill a 90 mm diam. Petri dish within 5 days at 25 °C.
Colony hyaline, radial, indistinctly zonate, aerial hyphae
spreading uniformly throughout the colony. Conidiation
starting after 4 days, light green (M. 28A5) or deep green
(M. 27E8) conidia forming in pustules, appearing around
the margin of the colony. No chlamydospores observed. No
distinct odour. Agar not pigmented. On PDA after 72 h
colony radius 12–17 mm at 15 °C, 21–30 mm at 20 °C,
35–44 mm at 25 °C, 21–24 mm at 30 °C; colonies on PDA
fill a 90 mm diam. Petri dish within 5–6 days at 25 °C.
Colony radial, mycelium dense, not finely zonate, aerial
hyphae abundant, more abundant in distance from the
inoculum. Conidiation starting after 3 days, greyish yellow
(M. 2C5) or greyish green (M. 29C5) conidia formed
abundant on aerial hyphae. On SNA after 72 h colony
radius 11–14 mm at 15 °C, 17–24 mm at 20 °C,
33–36 mm at 25 °C, 14–22 mm at 30 °C; colonies on SNA
fill a 90 mm diam. Petri dish within 6 days at 25 °C.
Colony hyaline, radial, mycelium loose, indistinctly
123
zonate, aerial hyphae more abundant in distance from the
inoculum. Conidiation starting after 3 days, greyish yellow
(M. 2B4) or greyish green (M. 29C5) conidia formed
abundant on aerial hyphae.
Material examined: REPUBLIC OF KOREA,
Gyeongsangbuk Province, Uljin, Jinjosan Mountain, elev.
170 m, isolated from root of Pinus densiflora Siebold &
Zucc. (Pinaceae) under a fairy ring of Tricholoma matsutake (S. Ito & S. Imai) Singer, October 2013, S.Y. Oh,
PF066 (SFC20131005-S066, holotype), ex-type living
culture KACC 48487; ibid., Gangwon Province, Hongcheon, Gongjaksan Mountain, elev. 300 m, isolated from
root of P. densiflora under a fairy ring of T. matsutake,
September 2013, S-Y. Oh, SFC20130926-S004.
GenBank numbers: ITS = MH050352, RPB2 =
MH025988, TEF1-a = MH025979.
Notes: Morphologically and phylogenetically (Figs. 92,
93), Trichoderma koreanum is closely related to T.
tomentosum Bissett, T. ceraceum P. Chaverri & Samuels,
and T. linzhiense K. Chen & W.Y. Zhuang. However, T.
tomentosum has shorter phialides (4.5–5 9 3–3.2 lm) and
narrower conidia (3.2–3.5 9 2.2–2.5 lm). Trichoderma
linzhiense grows faster at 25 °C (51–60 mm on CMD,
62–63 mm on PDA, and 50–51 mm on SNA after 3 days).
Trichoderma ceraceum grows slower at 20 °C (15–17 mm
Fungal Diversity
Fig. 91 Leptobacillium
leptobactrum var. calidius
(NFCCI 4235). a Colony
morphology on PDA (front
view). b Colony morphology on
PDA (reverse view). c Conidia
produced in long chains from
phialides. d Conidiophores
bearing phialides producing
conidia in chain. e Enlarged
view of conidia. f Conidia
produced in gloeosporic mass at
the tip of phialides. Scale bars
10 lm
on PDA and 6–14 mm on SNA) and has shorter phialides
(6.5–7.7 9 3–3.5 lm).
Trichoderma pinicola S-Y. Oh, M.S. Park & Y.W. Lim,
sp. nov.
MycoBank number: MB824662; Facesoffungi number:
FoF04460, Fig. 94
Etymology: The specific epithet ‘‘pinicola’’ refers to the
genus name of Pinus densiflora, the source of type strain
Holotype: SFC20130926-S233.
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1963).
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Conidiophores branched at an acute to
123
Fungal Diversity
Fig. 92 Phylogenetic tree for Trichoderma from a maximum likelihood analysis based on the combined TEF1-a and RPB2 alignments.
Branch support values are given as ML bootstrap values (C 70% ML)
and Bayesian posterior probabilities (C 0.95 BYPP). The scale bar
indicates the number of nucleotide substitutions per site. New species
names are in blue. The letter ‘‘T’’ indicates ex-type strains
the
main
axis,
mostly
paired.
Phialides
7.8–13.3 9 (2.5 -)2.6–4.1(-4.5) lm, l/w 1.9–4.4(-4.5),
1.8–3.3 lm wide at the base (n = 30), typically formed in
whorls of 3 or rarely solitary, subulate or lageniform.
Conidia 3.5–4.9(-5) 9 2.8–3.5(-3.6) lm, l/w 1.1–1.6
(n = 30), smooth, subglobose to ellipsoid. Chlamydospores
rare, globose, 5.2–10(-10.1) 9 5.2–10 lm, l/w 1–1.02
(n = 10). No distinct odour. Agar not pigmented.
Culture characteristics: On CMD after 72 h colony
radius 14–18 mm at 15 °C, 23–34 mm at 20 °C,
39–50 mm at 25 °C, 34–45 mm at 30 °C; colonies on
CMD fill a 90 mm diam. Petri dish at 4 days at 25 °C.
Colony hyaline, radial, not zonate, mycelium common,
aerial hyphae inconspicuous. Conidiation starting after
4 days, greyish green (M. 28B5) or deep green (M. 28E8)
conidia formed scarcely on aerial hyphae around the margin of the colony. On PDA after 72 h colony radius
12–15 mm at 15 °C, 18–28 mm at 20 °C, 30–38 mm at
25 °C, 21–36 mm at 30 °C; colonies on PDA fill a 90 mm
diam. Petri dish within 6 days at 25 °C. Colony hyaline,
radial, with wavy margin, mycelium common, aerial
hyphae more abundant in colony centre. Conidiation
starting after 9 days, greyish green (M. 27C6) or pastel
green (M. 29A4) conidia formed on aerial hyphae, more
abundant around the margin of the colony. On SNA after
72 h colony radius 9–12 mm at 15 °C, 19–24 mm at
20 °C, 30–43 mm at 25 °C, 19–33 mm at 30 °C; colonies
on SNA fill a 90 mm diam. Petri dish within 5–6 days at
25 °C. Colony hyaline, radial, mycelium loose, spreading
uniformly throughout the colony. Conidiation starting after
123
Fungal Diversity
Fig. 93 Trichoderma koreanum (SFC20131005-S066, holotype). a Colony on CMD after 72 h. b Colony on PDA after 72 h. c Colony on SNA
after 72 h. d–g Conidiophores. h Conidia. Scale bars d–h = 20 lm
3–4 days, greyish green (M. 29C5) or deep green (M.
27E8) conidia formed abundant on aerial hyphae.
Material examined: REPUBLIC OF KOREA, Gangwon
Province, Hongcheon, Gongjaksan Mountain, elev. 300 m,
isolated from root of Pinus densiflora under a fairy ring of
Tricholoma matsutake, September 2013, S-Y. Oh, PF233
(SFC20130926-S233, holotype), ex-type living culture
KACC 48486; ibid., Hongcheon, Gongjaksan Mountain,
elev. 300 m, isolated from root of P. densiflora under a
fairy ring of T. matsutake, September 2013, S-Y. Oh,
SFC20130926-S111.
GenBank numbers: ITS = MH050354, RPB2 =
MH025993, TEF1-a = MH025981.
Notes: Trichoderma pinicola is morphologically similar
to T. hirsutum K. Chen & W.Y. Zhuang, the latter has
longer phialides (11.4–18.3 9 2.5–3.1 lm) and lacks
chlamydospores in CMD. Phylogenetically (Fig. 92), T.
pinicola is closely related to T. simplex K. Chen & W.Y.
Zhuang. However, T. simplex has more abundant aerial
hyphae and conidia formation, and faster growth at 25 °C
(49–56 mm on CMD and 52–56 mm on PDA after 3 days).
In addition, T. pinicola showed wavy margin on PDA.
Trichoderma rugulosum M.S. Park, S-Y. Oh & Y.W. Lim,
sp. nov.
MycoBank number: MB824663; Facesoffungi number:
FoF04461, Fig. 95
Etymology: The specific epithet refers to the wrinkle in
colony reverse on PDA.
Holotype: SFC20180301-001
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1963).
Sexual morph Undetermined. Asexual morph
Hyphomycetous. Conidiophores symmetry, branches
mostly paired. Phialides typically formed in whorls of 3–4,
ampulliform to lageniform, 5.9–10.3(-10.5) 9 2.1–3.6(-3.9)
lm, l/w 1.9–3.7(-3.8), (1.2 -)1.3–1.9 lm wide at the base
(n = 30). Conidia green, smooth, globose, subglobose or
broadly ellipsoidal, (2.5–)2.6–3.2 9 (2.3–)2.3–2.9 lm, l/w
1–1.2(-1.3) (n = 30).
Culture characteristics: On CMD after 72 h colony
radius 10–14 mm at 15 °C, 24–29 mm at 20 °C,
62–69 mm at 25 °C, 69–75 mm at 30 °C; colonies on
CMD fill a 90 mm diam. Petri dish within 4 days at 25 °C.
Colony hyaline, radial, mycelium loose, indistinctly
zonate, aerial hyphae more abundant in distance from the
inoculum. Conidiation starting after 3 days, deep green (M.
27E8) or greyish green (M. 27c4) conidia forming in
pustules. No chlamydospores observed. No distinct odour.
Agar not pigmented. On PDA after 72 h colony radius
9–11 mm at 15 °C, 13–21 mm at 20 °C, 38–43 mm at
25 °C, 22–42 mm at 30 °C, colonies on PDA fill a 90 mm
diam. Petri dish within 6 days at 25 °C. Colony radial,
123
Fungal Diversity
Fig. 94 Trichoderma pinicola (SFC20130926-S233, holotype). a Colony on CMD after 72 h. b Colony on PDA after 72 h. c Colony on SNA
after 72 h. d–f Conidiophores. g Chlamydospores. h Conidia. Scale bars d–h = 20 lm
mycelium dense, zonate, aerial hyphae abundant, spreading
uniformly throughout the colony, forming wrinkle in distance from the inoculum in reverse colony, at 30 °C colony
radial, mycelium dense, with wavy margin. Conidiation
starting after 3 days, effuse in aerial hyphae or in densely
disposed granules, more abundant in 4–5 concentric rings,
yellowish green (M. 30B8). No chlamydospores observed.
No distinct odour. Olive yellow (M. 3C8) pigment diffusing into the agar. On SNA after 72 h colony radius 3–6 mm
at 15 °C, 7–11 mm at 20 °C, 14–29 mm at 25 °C,
10–21 mm at 30 °C; colonies on SNA a 90 mm diam. the
Petri dish within 8–9 days at 25 °C. Colony hyaline, radial,
mycelium loose, not finely zonate, aerial hyphae common.
Conidiation starting after 3 days, deep green (M. 27E8) or
yellowish green (M. 30B8) conidia forming in pustules,
pustules forming in 2–3 concentric rings.
Material examined: REPUBLIC OF KOREA, Jeju
Province, Chuja-do Island, isolated from Chondria crassicaulis Harvey, September 2017. M.S. Park, F181
(SFC20180301-001, holotype), ex-type living culture
KACC 48485; ibid., Chuja-do, isolated from Sargassum
thunbergii (Mertens ex Roth) Kuntze, September 2017,
M.S. Park, SFC20180301-002.
GenBank numbers: ITS = MH050353, RPB2 =
MH025986, TEF1-a = MH025984.
123
Notes: Trichoderma rugulosum is isolated from Chondria crassicaulis and Sargassum thunbergii (seaweeds) in
South Korea. Morphologically and phylogenetically, T.
rugulosum is closely related to species in the T. harzianum
complex. However, these species have faster growth at
30 °C on SNA (32–70 mm on SNA after 3 days). T.
rugulosum shows a wrinkle in colony reverse on PDA at
25 °C and wavy margin on PDA at 30 °C, features not seen
in members of the T. harzianum complex.
Hypocreales, genera incertae sedis
Emericellopsis Beyma
Emericellopsis Beyma (1940) was introduced with E.
terricola Beyma as the type species. Emericellopsis species
have been mostly isolated as saprobes from marine environments, soda lakes, and terrestrial habitats (GrumGrzhimaylo et al. 2013). We describe a novel species from
Korea based on phylogenetic analysis of a combined ITS
and TUB2 sequence dataset (Fig. 96).
Emericellopsis koreana Hyang B. Lee, S.J. Jeon & T.T.T.
Nguyen, sp. nov.
Index Fungorum number: IF554458; Facesoffungi
number: FoF05732, Fig. 97
Etymology: Named after the country where it was collected, Korea.
Fungal Diversity
Fig. 95 Trichoderma rugulosum (SFC20180301-001, holotype). a Colony on CMD after 72 h. b Colony on PDA after 72 h. c Colony on SNA
after 72 h. d Colony on PDA after 72 h from reverse. e–g Conidiophores. h Conidia. Scale bars e–h = 20 lm
Holotype: CNUFC-MOG1-1
Sexual morph Undetermined. Asexual morph
Hyphomycetous observed from MEA, acremonium-like.
Mycelia consisting of hyaline, smooth-walled, septate
hyphae, single or in bundles. Conidiophores mostly simple
and orthotropic. Conidiogenous cells (15.5–)31.5–40(–59)
lm long, tapering from 2(–2.5) lm at the base to 0.5 lm at
the apex. Conidia ellipsoid or oblong-ellipsoid, smoothsurfaced, 3–4(–5) 9 1.5–2(–2.5) lm (n = 50), hyaline,
adhering in slimy heads. No chlamydospores observed.
Culture characteristics: The isolate grows over a wide
range of temperatures with varying growth rates. The
average growth rates of CNUFC-MOG1-1 on MEA, CYA,
and PDA medium at 25 °C were 27.5, 17, and 15.5 mm
after 7 days, respectively. Optimal growth was observed
around 25 °C, slow growth was observed below 10 °C, and
no growth at 40 °C.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, garden of the Chonnam National University
located in Gwangju (35°10’20.2’’N 126°530 57.200 E), from
gut of a mosquito larva, 15 September 2016, H.B. Lee,
CNUFC-MOG1-1 (holotype), ex-type living culture,
JMRC:SF:013604.
GenBank numbers: ITS = MH173304, TUB2 =
MH243035
(CNUFC-MOG1-1);
ITS = MH173305,
TUB2 = MH243036 (CNUFC-MOG1-2).
Notes: Emericellopsis koreana, which forms a subclade
with E. donezkii Beliakova, E. humicola (Cain) Cain ex
Grosklags & Swift, E. persica Papizadeh et al. and
Emericellopsis sp., differs by having smaller conidia. In the
phylogenetic tree based on a combined ITS and TUB2
sequence dataset (Fig. 96), the strains CNUFC-MOG1-1
and CNUFC-MOG1-2 form a separate branch from other
related species of Emericellopsis and is considered to be a
new species.
Subclass Savoryellomycetidae Hongsanan et al.
Savoryellales Boonyuen et al.
Savoryellaceae Jaklitsch & Réblová
Savoryellaceae was introduced by Jaklitsch (2015) and
is typified by Savoryella E.B.G. Jones & R.A. Eaton. The
family is characterized by immersed to superficial, globose
to pyriform, coriaceous, papillate ascomata, clavate to
cylindrical, unitunicate asci, with an inamyloid apical ring,
paraphyses, ellipsoidal to fusiform, versicolorous, septate
ascospores, with or without polar mucilaginous pads or
appendages and dematiaceous, hyphomycetous asexual
morphs (Jaklitsch 2015). Most species of Savoryellaceae
are aquatic occurring on submerged wood in freshwater,
marine and brackish habitats worldwide (Jaklitsch 2015;
Xia et al. 2017). The generic type, Savoryella was re-circumscribed by Maharachchikumbura et al. (2016) and the
123
Fungal Diversity
higher level classification of the families in Savoryellomycetidae was revised by Hongsanan et al. (2017) based
on phylogenetic and molecular clock evidence. Six genera
are accepted in this family (Wijayawardene et al.
2017a, 2018a).
Canalisporium Nawawi & Kuthub.
The genus Canalisporium was introduced by Nawawi
and Kuthubutheen (1989) to accommodate muriform
asexual morph species, with conidia that are flattened
dorsiventrally, comprising a single layer of regularly
arranged cells, which are supported by a small, thin-walled,
cuneiform, pale basal cell. There are 12 species in this
genus (Zhao et al. 2013). Five species lack molecular data,
viz. C. kenyense Goh et al., C. microsporum G.Z. Zhao, C.
nigrum G.Z. Zhao, C. panamense A. Ferrer & Shearer and
C. variabile Goh & K.D. Hyde (Goh et al. 1998; Sri-Indrasutdhi et al. 2010; Zhao et al. 2013). In this study,
Canalisporium kenyense was collected from submerged
decaying wood in a freshwater stream in Thailand. The
taxon is re-circumscribed and illustrated as the reference
specimen (Figs. 98, 99).
Canalisporium kenyense Goh, W.H. Ho & K.D. Hyde,
Can J Bot 76: 148 (1998)
Facesoffungi number: FoF04845, Fig. 99
Holotype: KENYA, Mt. Kenya, Castle Forest, on rotten
wood, 25 Jan. 1984, P.M. Kirk, 1593a, IMI 285428a.
Saprobic on decaying wood submerged in a freshwater
stream. Sexual morph Undetermined. Asexual morph
Hyphomycetous, dictyosporous. Conidiophores macronematous, mononematous, unbranched, septate, up to
25 9 3–6 lm, hyaline to pale brown, smooth-walled.
Conidiogenous cells holoblastic, monoblastic, integrated,
terminal, determinate, hyaline to pale brown, smoothwalled. Conidia acrogenous, solitary, subglobose, flattened,
muriform, smooth, pale brown to dark brown,
40–50 9 25–30 9 13–20 lm with 2 straight columns of
vertical septa and 4–6 rows of transverse septa, slightly
constricted at the septa, apical rows of cells darker than the
basal rows, dark and thickly banded at the septa, canals in
the septa obscured by dark pigmentation, apex comprising
a single cell, the number of cells per conidium varies from
13 to 20, base comprising a single cell, cuneiform, sometimes swollen, thin-walled, pale brown, or comprising three
thin-walled, pale brown, small cells in a row.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Nang Lae Nai Village, on submerged
decaying wood in a freshwater stream, 31 December 2016,
Y.Z. Lu, CR01 (MFLU 17-1086, reference specimen
designated here); ibid., KUN-HKAS 97477.
GenBank
numbers:
ITS = MH701998,
LSU =
MH701999, TEF1-a = MH708885.
123
Known hosts and distribution: On decaying wood
(Kenya), submerged wood (Hong Kong), decorticated
branches of dead tree (China), decaying branches of
unidentified plant (China) (Goh et al. 1998; Zhuang 2001;
Zhao et al. 2013; Farr and Rossman 2018).
Notes: Our new collection resembles Canalisporium
kenyense in morphological characters of the conidiophores,
conidiogenous cells and conidia, and their measurements
align with those of the holotype (IMI 285428a) (Goh et al.
1998) and other specimens of C. kenyense (Zhao et al.
2013). Therefore, we identify our new collection as C.
kenyense which is the first report from Thailand. Phylogenetically, C. kenyense forms a distinct lineage within the
genus Canalisporium with strong support (96% ML and
1.00 BYPP; Fig. 98). Based on morphological characters
and phylogenetic support, we designate our collection
MFLU 17-1086 as a reference specimen.
Subclass Sordariomycetidae O.E. Erikss. & Winka
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae Réblová et al.
Chaetosphaeriaceae is a widespread, species-rich family, which was invalidly introduced by Locquin (1984) to
accommodate Chaetosphaeria Tul. & C. Tul., Loramyces
W. Weston, Niesslia Auersw., Rhagadostoma Körb. and
Zignoëlla Sacc., (Réblová et al. 1999; Hyde et al. 2017).
Réblová et al. (1999) re-described and validated
Chaetosphaeriaceae and accepted Ascocodinaea Samuels
et al., Chaetosphaeria, Melanochaeta E. Müll. et al., Melanopsammella Höhn., Porosphaerella E. Müll. &
Samuels, Porosphaerellopsis Samuels & E. Müll. and
Striatosphaeria Samuels & E. Müll. Currently, 38 genera
are accepted in Chaetosphaeriaceae (Hyde et al. 2017;
Tibpromma et al. 2018; Wijayawardene et al. 2018a). We
introduce Thozetella lithocarpi sp. nov. from Lithocarpus
sp. and record Macaranga tanarius as a new host record for
Chaetosphaeria panamensis based on both morphological
and phylogenetic analyses (Fig. 100).
Chaetosphaeria Tul. & C. Tul.
The saprobic genus, Chaetosphaeria was introduced by
Tulasne and Tulasne (1863) based on the type species, C.
innumera Berk. & Broome ex Tul. & C. Tul., and currently
comprise 165 epithets (Index Fungorum 2019). Chaetosphaeria is placed in Chaetosphaeriaceae (Chaetosphaeriales) based on molecular data (Réblová et al. 1999;
Huhndorf et al. 2004; Maharachchikumbura et al. 2015),
although it was previously placed in Lasiosphaeriaceae by
Barr (1990). Chaetosphaeria species have a diverse distribution having been recorded from both temperate and
tropical countries (i.e. Canada, China, Europe, Hong Kong,
New Zealand, Thailand) (Hyde et al. 1999, 2017; Réblová
2000; Perera et al. 2016; Wijayawardene et al. 2017a). Host-
Fungal Diversity
Fig. 96 Phylogenetic tree of Emericellopsis koreana (CNUFCMOG1-1 and CNUFC-MOG1-2) and related species based on
maximum likelihood analysis of a combined ITS and TUB2 sequence
dataset. Sequences of Bionectria spp. were used as outgroup taxa.
Bootstrap values (C 50%) from 1000 replications are indicated at the
nodes. New taxa are shown in blue and ex-type strains in bold
specificity of the taxa in this group has not yet been proven
given that they have been recorded from various plant
families (i.e. Arecaceae, Euphorbiaceae, Pinaceae) (Hyde
et al. 1999; Perera et al. 2016; Farr and Rossman 2018).
globosa, inner layer comprising several layers of flattened,
brown pseudoparenchymatous cells, arranged in textura
prismatica. Paraphyses 3–4 lm wide at the base, tapering
towards the apex, numerous, septate, arising from the basal
cavity, embedded in a hyaline gelatinous matrix. Asci 125–
150 9 9–12 lm (
x = 137.5 9 11.2 lm, n = 10), 8-spored,
unitunicate, arising from the basal hymenium, cylindrical,
rounded at the apex, with a J-, apical ring. Ascospores 50–
60 9 3–4 lm (
x = 56.4 9 3.5 lm, n = 20), fasciculate,
hyaline, filiform, straight to gently curved, with rounded
ends, slightly tapering to base, 7-septate, smooth-walled,
lacking a gelatinous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
8 mm diam. after 2 weeks at 25–30 °C, colonies medium
dense, circular, surface slightly rough with edge fimbriate,
effuse, velvety, margin, slightly irregular; from above, light
brown to yellowish at the margin, light brown to grey at the
centre, from below white to yellowish at the middle, light
brown at the margin, grey to whitish at the centre, white to
light purple at the middle; mycelium cream to whitish with
tufting; not producing pigmentation in PDA.
Chaetosphaeria panamensis Huhndorf & F.A. Fernández,
Fungal divers 19:33 (2005)
Facesoffungi number: FoF02657, Fig. 101
Holotype: PANAMA, Barro Colorado Island National
Monument, Shannon trail, 50 to 150 m, [9.1667, -79.8333],
on decorticated wood, 23 August 1997, SMH, FAF,
SMH3596 (in F).
Saprobic on decaying wood of Macaranga tanarius.
Sexual morph Ascomata 200–250 lm diam., 180–230 lm
high, black, scattered, solitary, sparse, semi-immersed in
the host at the base of ascomata, becoming superficial,
globose, coriaceous, setose, ostiolate, papillate. Setae
scattered over entire ascomata, dark brown, stiff, pointed,
50–60 lm long. Papilla central, short, brown, with periphyses. Peridium 25–35 lm thick, composed of two cell
layers, outer layer comprising 4–5 layers of thick, dark
brown cells, arranged in textura angularis to textura
123
Fungal Diversity
Fig. 97 Emericellopsis koreana (CNUFC-MOG1-1, holotype). a, b
Colonies on malt extract agar (MEA) (a: obverse view, b: reverse
view). c Conidiophores with branched form in the basal region (LM).
d, g Unbranched conidiophores, conidial heads (LM). e, f Conidia on
slimy head and single conidia (LM). g–i Curved or straight
conidiophore and ellipsoid or oblong-ellipsoid conidia (SEM). Scale
bars c–g = 20 lm, h = 4 lm, i = 5 lm
Material examined: TAIWAN, Chiayi, Shihnong Forest
Area, decaying wood of Macaranga tanarius (L.) Müll.Arg
(Euphorbiaceae), 25 June 2017, D.S. Tennakoon, DTW027
(MFLU18-0087), living culture, FU30910.
Known hosts and distribution: Decorticated wood
(Panama), Pinus sp. (Thailand), Macaranga tanarius
(Taiwan) (Huhndorf and Fernández 2005; Perera et al.
2016).
GenBank numbers: ITS = MH974685, LSU = MH974686.
Notes: Chaetosphaeria panamensis was introduced by
Huhndorf and Fernández (2005), and was collected from
decorticated wood in Panama. In this study, C. panamensis
is reported on another host, Macaranga tanarius from
Taiwan. Phylogenetic analyses based on a combined LSU,
ITS and TUB2 sequence dataset show that our strain
(FU30910) clusters with the ex-type strain of C. panamensis (SMH 3596) and another representative strain
(MFLUCC 15-1011) with high support (100% ML and
1.00 BYPP; Fig. 100). Based on morphological characters
(superficial, globose ascomata with setae, cylindrical asci
and filiform, slightly curved, 7-septate, hyaline ascospores)
and molecular data of ITS region, our isolate is identical to
the type (SMH 3596; Huhndorf and Fernández 2005) and
hence, the new isolate is identified as C. panamensis. This
123
Fungal Diversity
Fig. 98 Phylogram generated from maximum likelihood analysis of a
combined LSU and ITS sequence dataset of species in Canalisporium
Nawawi & Kuthub. and Savoryella E.B.G. Jones & R.A. Eaton
(Savoryellaceae). Twenty-three strains are included in the combined
sequence dataset, consisting of 1768 total characters with gaps (655
characters for ITS and 1113 for LSU). Pleurotheciella rivularia
Réblová et al. (CBS 125238) is the outgroup taxon. The best scoring
RAxML tree with a final likelihood value of -7016.458694 is
presented. RAxML bootstrap support values equal to or greater than
75% and Bayesian posterior probabilities equal to or higher than 0.95
BYPP are given above the nodes (ML/BYPP). Hyphen (‘‘–’’)
indicates a value lower than 75% for RAxML and a posterior
probabilities lower than 0.95 for Bayesian inference analysis. Newly
generated sequences are in blue. Ex-type strains are in bold
is the first record of C. panamensis on decaying wood of
Macaranga tanarius from Taiwan.
mass, with a white spore mass at the apex. Microawns 24–
54 lm long, 3–3.5 lm wide, visible as small hairs on the
sporodochial mass, aseptate, variously-shaped, sigmoid or
sickle-shaped, apex straight, hyaline, smooth-walled, thickwalled. Conidiophores 21–30 9 1.5–3.5 lm, macronematous, packed in a bundle, usually ramose. Conidia 20–
35 9 2–3 lm (
x = 26 9 2.7 lm, n = 25), falcate, inequilateral, truncate at both ends, hyaline, smooth-walled, with
a single filiform setula at each end, 6.9–8.7 lm long.
Culture characteristics: Colonies on MEA reaching
35 mm diam. after 3 weeks, margins effuse, grey to dark
brown, flat, lacking aerial mycelium, reverse dark brown to
black.
Material examined: THAILAND, Chiang Mai Province,
on a dried seed of Lithocarpus sp. (Fagaceae; 19 species of
Thozetella Kuntze
We follow the latest treatment and updated accounts of
Thozetella in Tibpromma et al. (2018).
Thozetella lithocarpi R.H. Perera & K.D. Hyde, sp. nov.
Index Fungorum number: IF555300; Facesoffungi
number: FoF04923, Fig. 102
Etymology: Referring to the host genus, Lithocarpus.
Holotype: MFLU 16-1068
Saprobic on Lithocarpus fruits. Sexual morph Undetermined. Asexual morph Hyphomycetous. Colonies
effuse, superficial, sessile sporodochial, greenish white.
Sporodochia subulate, scattered, sessile, of greenish white
123
Fungal Diversity
Fig. 99 Canalisporium kenyense (MFLU 17-1086, reference specimen). a Colony on substrate. b–e Conidia with attached conidiogenous cells.
f–j Conidia. Scale bars a = 200 lm, b = 50 lm, c–j = 20 lm
Lithocarpus have been reported from Chiang Mai Province, according to BGO plant database), 22 December
2015, R.H. Perera, S-7 (MFLU 16-1068, holotype), extype living culture, MFLUCC 16-0194.
GenBank
numbers:
ITS = MH810433,
LSU = MH810432.
Notes: Thozetella lithocarpi (MFLUCC 16-0194) shows
a close relationship with T. pandanicola Tibpromma &
K.D. Hyde, T. pinicola S.Y. Yeung et al. and T. nivea
(Berk.) Kuntze (Fig. 100). However, T. lithocarpi can be
123
distinguished from T. pandanicola and T. pinicola in
having larger conidia and presence of microawns (Jeewon
et al. 2009; Tibpromma et al. 2018). Thozetella lithocarpi
produces microawns with a straight apex, while they are
undulating to geniculate in T. nivea (Barbosa et al. 2011).
A comparison of nucleotides in ITS gene region shows that
T. lithocarpi differs from T. pandanicola and T. pinicola by
9 nucleotides (1.9%) and T. nivea by 18 nucleotides
(3.9%). Our new species is also compared with Thozetella
species that lack molecular data. Thozetella lithocarpi
Fungal Diversity
Fig. 100 Phylogram generated from maximum likelihood analysis of
a combined LSU, ITS and TUB2 sequence dataset of Chaetosphaeriaceae. Maximum likelihood bootstrap support values greater than
70% and Bayesian posterior probabilities (BYPP) above 0.95 are
shown above the nodes. The new isolates are in blue and ex-type
strains in bold. The tree is rooted with Gelasinospora tetrasperma
(AFTOL-ID 1287) and Sordaria fimicola (CBS 508.50)
123
Fungal Diversity
Fig. 101 Chaetosphaeria panamensis (MFLU 18-0087). a Appearance of ascomata on host. b Close-up of the ascoma. c, d Section of
ascoma. e Section through ostiole. f Section of peridium.
g Paraphyses. h–j Asci. k–n Ascospores. o Colony from above.
p Colony from below. Scale bars c, d = 100 lm, e = 50 lm,
f = 30 lm, g–j = 60 lm, k–n = 20 lm
produces comparatively larger conidia (20–35 9 2–3 lm
versus 11–17 9 2–2.5 lm) and smaller microawns
(24–54 lm long, 3–3.5 lm wide versus 40–100 lm long,
2.5–4 lm wide) than T. cubensis R.F. Castañeda & G.R.W.
Arnold (Castañeda-Ruiz and Arnold 1985). Thozetella
canadensis Nag Raj produces verrucose microawns, while
T. lithocarpi produces smooth-walled microawns (Nag Raj
1976). Thozetella aculeata Prisc. Silva & Grandi, T. buxifolia Allegr. et al., T. effusa B. Sutton & G.T. Cole, T.
radicata (E.F. Morris) Piroz. & Hodges, T. serrata Whitton
et al., T. submersa F.R. Barbosa & Gusmão and T. tocklaiensis (Agnihothr.) Piroz. & Hodges also produces
smaller conidia than those of T. lithocarpi. Thozetella
ypsiloidea J.S. Monteiro et al. differs from T. lithocarpi by
its Y-shaped microawns. Microawns of T. serrata have a
serrated edge, while microawns have a straight apex in T.
lithocarpi (Monteiro et al. 2016).
123
Coniochaetales Huhndorf et al.
Coniochaetaceae Malloch & Cain
Fungal Diversity
Coniochaetaceae was established to accommodate two
genera Coniochaeta (Sacc.) Cooke and Coniochaetidium
Malloch & Cain by Malloch and Cain (1971). Species of
Coniochaetaceae can be distinguished from other families
in having ascospores with elongated germ slits. Cultures
are frequently pink or orange and have a yeast-like
appearance. They usually grow better at low temperatures.
Conidia are produced in abundance as phialospores or
rarely as aleuriospores (Malloch and Cain 1971; Huhndorf
et al. 2004; Wanasinghe et al. 2018). We followed the
latest phylogenetic analyses and the updated accounts of
Coniochaeta in Nasr et al. (2018), Samarakoon et al.
(2018) and Wanasinghe et al. (2018). A new species is
introduced based on its morphological distinctiveness
coupled with strong phylogenetic support (Fig. 103).
Coniochaeta (Sacc.) Cooke
We follow the latest treatment and updated accounts of
Coniochaeta in Samarakoon et al. (2018) and Wanasinghe
et al. (2018).
Coniochaeta simbalensis S. Rana & S.K. Singh, sp. nov.
MycoBank number: MB824288; Facesoffungi number:
FoF04831, Fig. 104
Etymology: The specific epithet ‘‘simbalensis’’ refers to
the place of collection.
Holotype: AMH-9941
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Saprobic on mushroom rhizospheric soil. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Hyphae
thin- to thick-walled, simple to branched, smooth-walled,
constricted near the septa, wall thickened, guttulate, anastomosis observed. Conidiophores 1.2–6.3 lm wide,
starting in 2–3 days from slender, thin hyaline to thick,
smooth-walled hyphae, mostly reduced to conidiogenous
cells. Phialides 1.8–60 9 1–3.7 lm (
x = 14.3 9 2.2 lm,
n = 30), variable, produced laterally from superficial
hyphae, solitary or in groups (1–3), ventricose, base narrow,
and middle swollen with elongated narrow tip (collar)
tapered collula, variable in length, all curved, reduced in size
to dentate structure like adelophialides to elongated branched. Conidia 2.2–8.8 9 1.3–3.3 lm (
x = 4.5 9 2 lm,
n = 30), 1-celled, oval to cylindrical to subglobose, produced sarcinately in gleosporic mass, rarely found in zipperlike arrangement, yeast-like cells observed frequently,
monopolar budding seen in conidia, sporulation abundant,
variable in size. Chlamydospores 5–8.6 9 3.4–6.7 lm
(
x = 6.8 9 4.9 lm, n = 30), light to dark olivaceous brown,
observed frequently, terminal to intercalary, solitary or in
branched chains, wall thickened and darkened, constricted
near the septa, variable in shape, globose, cylindrical to
pyriform, sometimes produced laterally from short and long
stalk of * 16–28 lm. Stalks septate, simple to branched,
light to olivaceous brown, smooth-walled.
Culture characteristics: Colonies on PDA reaching
39–45 mm diam. after 2 weeks at 25 °C; from above
brownish grey (9D2) with margins fading to dull red (9B3),
flat, sulcate, entire with smooth margin; from below,
brownish grey (9E2) with outer margins fading to pale
orange (6A3), sulcate. Colonies on MEA reaching to
15–20 mm diam. after 2 weeks at 25 °C; from above
brownish grey (6F2), slightly raised, sulcate, and irregular;
from below grey (6E1). Colonies on PCA reaching
29–34 mm diam. after 2 weeks at 25 °C; from above
smoke brown (4F3); from below, silver grey (4E2) with a
margin of nearly 5 mm yellowish white (4A2) in flat,
glazy, margin entire and irregular.
Material examined: INDIA, Himachal Pradesh, Kangra
Dist, Simbal (31.9754 N’’ 76.6507 E’’), Mushroom rhizospheric soil, 8 July 2017, S. Rana, (AMH 9941, holotype),
ex-type living culture, NFCCI 4236.
GenBank numbers: ITS = MG825743, LSU = MG917738.
Notes: Coniochaeta simbalensis differs from other Coniochaeta species based on the sequence analysis. On
megablast analysis, ITS sequence of C. simbalensis shows
94.92% (468/493) similarity and 25 gaps (5%) with C.
cateniformis (Perdomo et al.) Gené & Guarro (UTHSC
01-1644, type), 92.23% (487/528) similarity and 29 gaps
(5%) with C. canina (Deanna A. Sutton et al.) Deanna A.
Sutton et al. (UTHSC 11-2460, type), 93.37% (366/392)
similarity and 5 gaps (1%) with C. hoffmannii (J.F.H.
Beyma) Z.U. Khan et al. (CBS 245.38, type) and 87.36%
(491/562) similarity and 42 gaps (7%) with C. acaciae
Samarakoon et al. (MFLUCC 17-2298). Interestingly, the
sources of isolation of these species are quite distinct.
Coniochaeta cateniformis (UTHSC 01-1644) was isolated
from canine bone marrow, C. canina (UTHSC 11-2460)
from bone aspirate, canine breed German Shepard, C.
hoffmannii (CBS 245.38) from butter (Khan et al. 2013)
and C. acaciae (MFLUCC 17-2298) from dead trunk and
branches of Acacia sp. (Samarakoon et al. 2018), whereas,
C. simbalensis was isolated from rhizospheric soil of
unidentified mushroom growing in soil.
Coniochaeta simbalensis produces brownish grey (9D2)
colonies with margins fading to dull red (9B3) and sulcate
on PDA, whereas C. cateniformis and C. canina produces
orange white and non-sulcate colonies. Similarly, C. simbalensis produces brownish grey (6F2), colonies on MEA
as against orange red to yellowish red colonies produced by
C. acaciae. Phialides of highly variable length were frequently observed in vitro by C. simbalensis. Conidia are
highly variable in shape and size in C. simbalensis, ranging
from oval to cylindrical to subglobose, non-truncate, and
straight; whereas conidia are ovoidal to ellipsoidal, truncate and smaller in C. cateniformis, and ellipsoidal, straight
123
Fungal Diversity
Fig. 102 Thozetella lithocarpi (MFLU 16-1068, holotype). a Dried
seed of Lithocarpus sp. b, c Appearance of conidiomata on host
substrate. d–f Conidiophores with conidia. g–i Microawns. j–l
Conidia. m Germinating conidium. Scale
c = 500 lm, d–i = 20 lm, j–m = 10 lm
to slightly curved in C. canina (Perdomo et al. 2013; Troy
et al. 2013) and ellipsoidal in C. acaciae (Samarakoon
et al. 2018). Coniochaeta simbalensis also readily produces
chlamydospores in culture, a feature absent in C. acaciae
and C. prunicola Damm & Crous (Damm et al. 2010).
Therefore, based on phylogenetic inference (Fig. 103),
morphological and cultural distinctness C. simbalensis is
proposed as a new species.
T. labiosa S.W. Wong et al., which was collected from
submerged wood in a small stream in Australia and Hong
Kong (Wong et al. 1998). The genus is characterized by
dark brown, immersed to semi-immersed, subglobose
ascomata, with periphyses, papillate, thin-walled, pale
brown peridium, paraphyses, 8-spored, unitunicate, cylindrical asci, with an unusual J-, lip-like, refractive apical
ring, and hyaline, ellipsoidal-fusiform, aseptate ascospores,
with narrow, roughed mucilaginous sheath (Wong et al.
1998). The asexual morph of this genus is unknown (Wijayawardene et al. 2017b). Based on the general morphology of the ascus apical ring observed with light
microscopy and scanning, transmission electron microscopy, a new genus was established (Wong et al. 1998).
Wong et al. (1998) suggested to place the genus in its own
family due to a unique character of its apical ring.
Wijayawardene et al. (2018a) treated the genus in Sordariomycetes, genera incertae sedis. We collected a
Phyllachorales M.E. Barr
Phyllachoraceae Theiss. & H. Syd.
We follow the latest treatment and updated accounts of
Phyllachoraceae in Dayarathne et al. (2017) and Mardones
et al. (2017, 2018). Updated phylogenetic analysis
(Fig. 105) was retrieved from Dayarathne et al. (2017).
Tamsiniella S.W. Wong et al.
Tamsiniella was introduced as a monotypic genus by
Wong et al. (1998) to accommodate the freshwater fungus,
123
bars
b = 1 mm,
Fungal Diversity
Fig. 103 Phylogram generated from maximum likelihood analysis for
Coniochaeta simbalensis using a combined LSU and ITS sequence
dataset based on the Tamura–Nei model (Tamura and Nei 1993).
Phylogenetics analyses were conducted in MEGA7 (Kumar et al.
2016). Chaetosphaeria innumera and C. pygmaea were used as
outgroup taxa. Type strains are indicated in bold. Newly generated
sequence is indicated in blue
specimen from a small river in Yunnan, China. Based on
morphological comparison, our collection is typical of the
type of Tamsiniella. We therefore, designate our collection
as an reference specimen based on morphological characteristics and geographical distribution (Fig. 105).
diam. [type: 130–225 lm high, 180–250 lm diam.], black,
gregarious or scattered, superficial, subglobose to ellipsoidal,
uni-loculate, thin-walled, laterally ostiolate, with a mass of
spores oozing when old. Peridium 6–9 lm wide [type:
5–11 lm wide], comprising 3–4 layers of dark brown, thickwalled, compressed cells of textura angularis. Hamathecium
comprising numerous, ca 6 lm wide at the base, 2 lm diam.
at the apex [type: 4–5 lm wide], cylindrical, unbranched,
hyaline, septate, paraphyses, slightly constricted at the septa,
tapering towards the apex. Asci 80–110 9 7.5–9 lm
(
x = 93 9 8.5 lm, n = 15) [type: 80–102 9 8–10.5 lm], 8spored, unitunicate, cylindrical to cylindric-clavate with a
short, twisted pedicel, apically obtuse and inwardly concave,
slightly wider than subapical apparatus, with J-, a refractive,
lip-like apical ring, 1.9–2 9 2.9–3.1 lm. Ascospores
Tamsiniella labiosa S.W. Wong, K.D. Hyde, W.H. Ho &
S.J. Stanley, Can J Bot 76(2): 334 (1998)
Facesoffungi number: FoF05052, Fig. 106
Holotype: AUSTRALIA, North Queensland, Mount
Lewis, on submerged wood in a small stream, July 1993,
T.M. and K.D. Hyde, ML9 (HKU (M) 2276; IFRD199014).
Saprobic on decaying wood submerged in freshwater.
Sexual morph Ascomata 100–120 lm high, 130–150 lm
123
Fungal Diversity
123
Fungal Diversity
b Fig. 104 Coniochaeta simbalensis (AMH 9941, holotype). a Colony
morphology on PDA (front view). b Colony morphology on SDA
(front view). c Colony morphology on PCA (front view). d Hyphal
wall septate, thickened, guttulate, and showing anastomoses. e Terminal to intercalary chlamydospores. f Phialides with gleosporic mass
of conidia. g Adelophialide with gleosporic mass of conidia
(magnified view). h Discrete phialides and adelophialides. i Ventricose phialides in group with conidia. j Discrete phialides and
dispersed conidia. k Mass of conidia. Scale bars = 10 lm
17–20 9 3.5–5 lm (
x = 19 9 4.5 lm, n = 15) [type:
(12–)15–21 9 3.8–4.5(–5) lm], overlapping 1-seriate, hyaline, aseptate, fusiform to ellipsoidal, straight or curved,
guttulate, thin-walled, with a thin, hyaline, mucilaginous
sheath, 1–2 lm wide. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
20 mm in 12 days at 25 °C, circular, white to yellow–
brown from above, white to light yellow from below,
surface smooth with sparse mycelium, dry, edge entire.
Material examined: CHINA, Yunnan Province, Pingbian, on submerged wood in a small river, 20 September
2017, W. Dong, WF-33A (MFLU 18-1191, a reference
specimen is designated here; ibid., KUN-HKAS 101711),
living culture, MFLUCC 18-1018 = KUMCC 18-0060.
Known hosts and distribution: On submerged woods or
twigs in Australia, Brazil, China, Hong Kong (Wong et al.
1998; Barbosa et al. 2013; this study).
GenBank
numbers:
ITS = MK034865,
LSU =
MK034866, SSU = MK034867.
Notes: Our isolate shares the size range of asci, ascospores, peridium and paraphyses with the type of Tamsiniella labiosa (see description). However, it has a slightly
smaller ascomata compared to the type. Our isolate was
collected from submerged wood in Yunnan, China,
whereas the type specimen was collected from submerged
wood in North Queensland, Australia. Based on morphological comparison and geographical distribution, we
hence, designate our collection as a reference specimen of
T. labiosa.
In phylogenetic tree, Tamsiniella labiosa forms a distinct lineage in the order Phyllachorales M.E. Barr with
moderate support (77% ML and 0.99 BYPP, Fig. 105).
Phyllachorales species are distinctive as they are biotrophic on various hosts (Pearce and Hyde 1994; Dayarathne et al. 2017; Mardones et al. 2017, 2018).
Phyllachorales species are characterized by deep black
stromata of various shapes; pseudostroma inside the host
tissue and usually beneath an epidermal clypeus; perithecia
usually strongly melanized; cylindrical to clavate asci with
an inconspicuous apical ring; and globose to filiform,
mostly hyaline, 1-celled, rarely brown or septate ascospores (Parbery 1967; Cannon 1991; Dayarathne et al. 2017;
Mardones et al. 2017, 2018). However, our collection was
isolated from decaying wood submerged in freshwater,
which was the same habitat as another freshwater ascomycete genus Ascovaginospora Fallah et al. in Phyllachoraceae (Wijayawardene et al. 2018a). Therefore,
Tamsiniella is accommodated in Phyllachoraceae based on
phylogenetic analyses.
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss
Lasiosphaeriaceae Nannf.
Lasiosphaeriaceae was established by Nannfeldt (1932),
circumscribed by species with black ascomata and cylindrical asci, brown to hyaline ascospores, and typified by
Lasiosphaeria Ces. & De Not. The family was re-circumscribed by Maharachchikumbura et al. (2015, 2016) and 35
genera were accepted in the family. Based on phylogeny,
Lasiosphaeriaceae is sister to Chaetomiaceae with high
support and this family has been shown to be paraphyletic,
with many genera polyphyletic (Chang et al. 2010; Kruys
et al. 2015; Maharachchikumbura et al. 2015, 2016).
Wijayawardene et al. (2018a) listed 32 genera in
Lasiosphaeriaceae.
Zopfiella G. Winter
Zopfiella was established by Winter (1884) to introduce
Z. tabulata (Zopf) G. Winter and Z. curvata (Fuckel) G.
Winter. Phylogenetic studies reported that the genera
Zopfiella, Triangularia Boedijn, Cercophora Fuckel and
Podospora Ces. are polyphyletic (Miller and Huhndorf
2005; Cai et al. 2005, 2006a, b; Chang et al. 2010; Kruys
et al. 2015; Maharachchikumbura et al. 2015, 2016). A new
species Z. indica, is introduced within Lasiosphaeriaceae
based on evidence from morphology and DNA sequence
data (Fig. 107).
Zopfiella indica Devadatha, Jeewon & V.V. Sarma, sp.
nov.
Index Fungorum number: IF554286; Facesoffungi
number: FoF04269, Fig. 108
Etymology: Refers to country of its origin
Holotype: AMH-9907
Saprobic on the bark of intertidal mangrove wood. Sexual
morph Ascomata 320–555 lm high, 220–405 lm diam.
(
x = 455.4 9 317 lm, n = 10), perithecial, lacking stromatic tissues, superficial, pyriform, solitary to gregarious,
coriaceous, dark brown to black, enclosed with copious,
flexuous, septate, light brown to dark brown, short to long
hyphae, prevalent at the base, about 75–200 lm long, 1–
2.5 lm thick, ostiolate. Ostiole 50–130 lm long, 40–125 lm
diam. (
x = 105 9 104 lm, n = 10), black, short papillate,
straight to curved, darker than remaining part of the perithecium, filled with brown cells. Periphyses 1–2 lm wide
(
x = 1.7 lm,
n = 20).
Peridium
5–25 lm
wide
(
x = 13.2 lm, n = 10), membranous to coriaceous, pale
brown to dark brown, comprising two layers, inner stratum
123
Fungal Diversity
Fig. 105 Phylogram generated from maximum likelihood analysis
based on a combined LSU, SSU and ITS sequence dataset of taxa in
Phyllachorales and related orders. Bootstrap support values for
maximum likelihood (left) equal to or greater than 70% and the values
123
of Bayesian posterior probabilities (right) equal to or greater than 0.90
are given above the nodes. The new isolate is in blue. The tree is
rooted with Xylaria polymorpha (MUCL49884) and X. hypoxylon
(STMA07069). Ex-type strains are indicated in bold
Fungal Diversity
Fig. 106 Tamsiniella labiosa (MFLU 18-1191, reference specimen).
a–c Appearance of black ascomata superficial on host. d Vertical
section of ascoma. e Structure of peridium. f Paraphyses. g–
j Unitunicate asci. k–m Ascospores. n Germinated ascospore
o Colony on PDA (from above). p Colony on PDA (from below).
Scale bars d = 50 lm, e, j–n = 10 lm, f–i, o = 20 lm
123
Fungal Diversity
flattened with several layers of hyaline to pale brown cells of
textura prismatica, outer stratum comprising 2–3 layers of
textura angularis. Paraphyses 1–2.5 lm wide (
x = 1.9 lm,
n = 20), hyaline, filiform. Asci 115–200 9 15–45 lm
(
x = 147.1 9 26.5 lm, n = 40), 8-spored, unitunicate,
cylindrical to clavate, evanescent, short pedicellate,
20–75 lm 9 2.5–7.5 lm (
x = 46 9 5 lm, n = 40), apical
ring
indistinct.
Ascospores
23–35 9 10–20 lm
(
x = 29.9 9 15.7 lm, n = 50), partially overlapping 1–2seriate, hyaline at first turning golden yellow and olivaceous
brown to dark brown at maturity, apical cell ovoid to ellipsoidal, rugose, apical cauda, single, lash-like, attached to the
apical part of the terminal cell, 8–55 9 3–8 lm
(
x = 20 9 5 lm, n = 20), smooth-walled, collapsing, primary appendage attached to the base of the pedicel and
similar to apical cauda, 15–25 9 3–8 lm (
x = 20 9 5 lm,
n = 20), mostly collapsing. Asexual morph Undetermined.
Culture characteristics: Ascospores germinated on
SWA within 24 h, germ tubes arise from ends of the
ascospore. Colonies on MEA reaching 70–85 mm diam.
after 25 days of incubation at room temperature, initially
deep olive grey, becoming dark olive grey at maturity,
surface umbonate, margin entire, velvety, circular, reverse
dark olive grey.
Material examined: INDIA, Tamil Nadu, Tiruvarur
District, Muthupet mangroves (10.4°N 79.5°E), on the bark
of intertidal mangrove wood, 28 November 2015, B.
Devadatha, AMH-9907 (holotype), ex-type living culture,
NFCCI-4217.
GenBank
numbers:
ITS = KY863506,
LSU =
KY863507,
SSU = MF168941,
RPB2 = MF182396,
TEF1-a = MF182400, TUB2 = MF406208.
Notes: Multigene phylogenetic analyses indicate that
Zopfiella indica shares a sister relation with Z. karachiensis
(S.I. Ahmed & Asad) Guarro and Triangularia tanzaniensis R.S. Khan & J.C. Krug in a strongly supported monophyletic clade (Fig. 107). Zopfiella indica is clearly
distinguished from Z. karachiensis and Triangularia tanzaniensis based on its ascospores having ovoid to ellipsoidal, rugose head cell, apical cauda attached to apical
part of the terminal cell and primary appendage attached to
the basal pedicel.
Species belonging to Podospora are predominantly
coprophilous whereas those belonging to Zopfiella predominantly thrive in soil or on plant substrata and less on
dung. Given the existing taxonomic confusion vis-a-vis the
genera Zopfiella and Podospora, we prefer to place Z.
indica in the genus Zopfiella based on its occurrence on
decaying wood, and presence of a septum in the dark cell.
Recent phylogenetic analysis showed that Zopfiella is
polyphyletic, interspersed with species belonging to other
genera and hence is in need of revision along with other
closely related genera such as Podospora and Triangularia,
123
both at the morphological and molecular level, by including a larger representation of the species belonging to these
genera. Phylogenetic analyses carried out by Cai et al.
(2006b) also indicated that Zopfiella does not constitute a
monophyletic group. They felt that the characters of nonostiolate ascomata, and the absence of gelatinous appendages in ascospores that are considered important in
delineating Zopfiella from Podospora (Guarro et al. 1991)
are not reliable in understanding phylogenetic relationships
(Cai et al. 2006b). This is because they found non-ostiolate
Zopfiella species interspersed in different clades in the trees
suggesting multiple origins of this morphological character. Also they found that the presence or absence of
gelatinous appendages are phylogenetically less informative as different Zopfiella species grouped in different
clades that also include many species possessing elaborate
gelatinous appendages. Cai et al. (2006b) suggested that
Zopfiella should be restricted to species with a septum in
the dark cell.
There are two strains under the name Zopfiella
karachiensis (IFO32902 and CBS 657.74) in GenBank,
which were included in our phylogenetic analysis. However, these two strains formed a separated lineage in our
phylogenetic tree (Fig. 107). Zopfiella karachiensis
(IFO32902) is sister to Z. lundqvistii (NBRC30585);
whereas, Z. karachiensis (CBS 657.74) clustered with
Triangularia tanzaniensis and Z. indica (our present
strain). Taxonomic revision of genera in this family is
needed based on multigene phylogenetic analyses.
Subclass Xylariomycetidae O.E. Erikss. & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Amphisphaeriaceae G. Winter
The family Amphisphaeriaceae was introduced by
Winter (1887) to accommodate Amphisphaeria and allied
taxa. Amphisphaeriaceae is mainly characterized by
perithecial, semi-immersed to erumpent ascomata, dark
peridium, unitunicate, cylindrical asci, with J ? or J-,
apical rings, pale to dark brown, septate ascospores and
forming coelomycetous asexual morphs (Senanayake et al.
2015; Maharachchikumbura et al. 2016). Three genera are
accepted in this family, Amphisphaeria Ces & De Not.,
Griphosphaerioma Höhn., and Lepteutypa Petr. (Wijayawardene et al. 2018a). An updated evolutionary relationship of the family in Xylariomycetidae was presented
by Samarakoon et al. (2016) and Hongsanan et al. (2017).
Amphisphaeria Ces. & De Not.
Amphisphaeria was established by Cesati and De
Notaris (1863) without assigning a generic type. Petrak
(1923) proposed A. umbrina (Fr.) De Not. as the lectotype
of Amphisphaeria. Hyde et al. (1996) epitypified and
described A. umbrina. Senanayake et al. (2015) re-
Fungal Diversity
Fig. 107 Phylogram generated from maximum parsimony analysis
based on a combined LSU, TUB2, ITS and RPB2 gene regions of
Zopfiella and related taxa in Lasiosphaeriaceae. Bootstrap support
values for maximum likelihood (green), maximum parsimony (blue)
equal to or greater than 75% and the values of Bayesian posterior
probabilities (purple) equal to or greater than 0.95 BYPP are given
above each branch respectively. The new isolate is in blue. Ex-type
strains are indicated in bold. The tree is rooted with Coniochaeta
discoidea (Udagawa & Furuya) Dania Garcı́a et al. (SANK12878)
123
Fungal Diversity
123
Fungal Diversity
b Fig. 108 Zopfiella indica (AMH-9907, holotype). a, b Ascomata
superficial on the bark of intertidal mangrove wood. c Squash mount
of ascoma. d Section through peridium. e Hyaline filiform paraphyses. f–i Immature and mature asci. j–o Ascospores with apical and
basal cauda. Scale bars c = 100 lm, h–i = 50 lm. d–g, j–o = 10 lm
circumscribed Amphisphaeria and introduced A. sorbi
Senan. & K.D. Hyde with the link between the sexual and
asexual morphs. We follow the latest treatment and updated accounts of Amphisphaeriaceae in Senanayake et al.
(2015) and Maharachchikumbura et al. (2016). Based on
phylogenetic analyses of a combined LSU, SSU and ITS
sequence dataset (Fig. 109) coupled with morphological
characteristics, we therefore, introduce a new species from
intertidal branches and twigs of Suaeda monoica Lam.
(Amaranthaceae) in India.
Amphisphaeria mangrovei Devadatha & V.V. Sarma, sp.
nov.
Index Fungorum number: IF554279; Facesoffungi
number: FoF04273, Fig. 110
Etymology: Named after the fungal habitat from marine
environment, where the fungus found.
Holotype: AMH-9948
Saprobic on intertidal branches and twigs of Suaeda
monoica. Sexual morph Ascomata 150–280 lm high, 140–
250 lm diam. (
x = 192 9 211 lm, n = 10), immersed to
erumpent, globose to subglobose, gregarious to solitary,
coriaceous, brown, short papillate, ostiolate. Ostioles 45–
60 lm long, 35–45 lm diam. (
x = 50 9 40 lm, n = 10),
short, periphysate, brown, 0.5–2 lm (
x = 1.5 lm, n = 20).
Peridium equal in thickness, 10–20 lm wide (
x = 13 lm,
n = 10), both at the base and sides, comprising two layers,
inner stratum with 3–4 layers of hyaline to light brown cells
of textura angularis and outer stratum with 2–3 layers of light
brown to brown cells of textura angularis, fusing with the
host tissue. Paraphyses 1–2 lm wide (
x = 1.7 lm, n = 20),
filamentous, septate, longer than asci, embedded in a gelatinous matrix. Asci 80–130 9 5–10 lm (
x = 100 9 8 lm,
n = 30), 8-spored, unitunicate, cylindrical to obclavate,
apically rounded with a J-, apical ring, short pedicellate.
Ascospores 12–15 9 4–6 lm (
x = 13 9 5 lm, n = 50),
partly overlapping 1-seriate, light brown, one median septate,
ellipsoidal, smooth-walled, lacking a mucilaginous sheath.
Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
SWA within 24 h, germ tubes arising from both ends of the
ascospore. Colonies on MEA reaching 80–90 mm diam.
after 7 days of incubation at room temperature, cream to
white from above, yellow and white at margin from below,
granular and cottony, undulate, umbonate, irregular.
Material examined: INDIA, Tamil Nadu, Tiruvarur,
Muthupet mangroves (10.4°N 79.5°E), on intertidal
branches and twigs of Suaeda monoica, 29 October 2016,
B. Devadatha, AMH-9948 (holotype), ex-type living culture, NFCCI-4247.
GenBank
numbers:
ITS = MG844283,
LSU =
MG767311, SSU = MG844279.
Notes: Amphisphaeria mangrovei differs from other
Amphisphaeria species in having smaller ascomata, asci
and ascospore dimensions and the marine habitat. Amphisphaeria mangrovei has similar morphological characters to A. sorbi such as clypeate ascomata, asci with J-,
apical ring and single, median septate, ellipsoidal ascospores (Senanayake et al. 2015). However, A. sorbi has
larger ascomata, asci and ascospores, and slightly constricted ascospores with a thick mucilaginous sheath. Amphisphaeria umbrina and A. vibratilis (Fuckel) E. Müll.
have clypeate ascomata, asci with J ? , discoid subapical
rings and larger ascospores. Phylogenetic analyses of a
combined LSU, SSU and ITS regions reveal that A. mangrovei is sister to A. sorbi and A. umbrina with significant
support (96% ML, 90% MP and 1.00 BYPP; Fig. 109).
Based on morphological characters and molecular phylogenetic analyses, a new species A. mangrovei is introduced.
Sporocadaceae Corda.
Jaklitsch et al. (2016b) proposed the family Sporocadaceae based on morphological observations and phylogenetic analyses of a concatenated ITS-LSU sequence
dataset with asexual morph genera that are acervular coelomycetes having hyaline, pale or dark brown, septate
conidia. The type genus is Seimatosporium Corda (Jaklitsch et al. 2016b). The family Sporocadaceae hitherto
includes 22 genera (Jaklitsch et al. 2016b; Wijayawardene
et al. 2017a, 2018a). We follow the latest treatment and
updated accounts of Sporocadaceae in Jaklitsch et al.
(2016b), Maharachchikumbura et al. (2017) and Wanasinghe et al. (2018). The updated phylogenetic analyses are
derived from Maharachchikumbura et al. (2017) and
Wanasinghe et al. (2018).
Bartalinia Tassi
We follow the latest treatment and updated accounts of
Bartalinia in Jaklitsch et al. (2016b) and Wanasinghe et al.
(2018). In this study, Bartalinia kunmingensis is introduced
from Zea mays (Poaceae) in Yunnan, China based on
morphological characteristic and phylogenetic analyses of
ITS and LSU sequence data (Fig. 111).
Bartalinia kunmingensis Thiyag., Wanas., Phookamsak &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556209; Facesoffungi
number: FoF05717, Fig. 112
Etymology: The specific epithet ‘‘kunmingensis’’ refers
to Kunming City, Yunnan, China, where the holotype was
collected.
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Fungal Diversity
Holotype: KUN-HKAS 102242
Saprobic on dead leaves of Zea mays. Sexual
morph Undetermined. Asexual morph Coelomycetous.
Conidiomata 80–110 lm high, 110–140 lm diam.
(
x = 126.2 9 99 lm, n = 10), pycnidial, dark brown to
black, immersed, slightly raised, solitary to gregarious, uniloculate, globose to subglobose, glabrous, ostiolate, with a
minute papilla. Conidiomata walls 8–20 lm wide, slightly
thick-walled, of equal thickness, comprising several cell
layers of brown, pseudoparenchymatous cells of textura
angularis, paler towards the inner layers. Conidiophores
arising from the inner cavity, reduced to conidiogenous cells. Conidiogenous cells (3.6–)4–7.5 9 2–5 lm
(
x = 5.4 9 2.8 lm, n = 30), holoblastic, phialidic,
rarely with 1–2 percurrent proliferations, discrete,
hyaline, ampulliform to subcylindrical, or obclavate, aseptate, smooth-walled. Conidia (17.5–)20–25 9 3–4 lm
(
x = 22.1 9 3.9 lm, n = 30), cylindrical to subcylindrical,
straight to slightly curved, 4-septate, not constricted at the
septa, with longest cell at the second from base, bearing
appendages; basal cell 2.5–4 lm long (
x = 3.2 lm), obconic, truncate at base, hyaline, thin and smooth-walled, second
cell from the base 6.5–8 lm long (
x = 7.4 lm), pale yellowish, third cell 4–5.5 lm long (
x = 4.8 lm), pale yellowish, fourth cell 4–5.5(–6) lm long (
x = 5 lm), pale
yellowish, apical cell 2–3(–3.7) lm long (
x = 2.9 lm),
conical, hyaline and smooth-walled, forming three-branched
tubular, flexuous, 10–20 lm long apical appendages; basal
appendages 5–6 lm long, single, absent at immature state,
tubular, unbranched, centric.
Culture characteristics: Colonies on PDA reaching
38–40 mm diam. after one week at room temperature. Colony dense, irregular in shape, flat, slightly raised, surface
smooth, with edge undulate, floccose to fluffy, entire margin,
forming black stromatic after 2 months; from above, white
yellowish to cream at the margin, grey yellowish to dark
yellowish, slightly radiated outwards colony, from below,
black; not producing pigmentation on agar medium.
Material examined: CHINA, Yunnan Province, Kumming, Kunming Institute of Botany, on dead leaves of Zea
mays L. (Poaceae), 5 November 2015, D.S. Tennakoon,
COE002 (KUN-HKAS 102242, holotype), ex-type living
culture, KUMCC 18-0178.
GenBank
numbers:
ITS = MK353083,
LSU =
MK353085, SSU = MK353148, RPB2 = MK492668,
TEF1-a = MK492656.
Notes: Phylogenetic analyses of a combined ITS and
LSU sequence dataset (Fig. 111) show that Bartalinia
kunmingensis (KUMCC 18-0178) clusters with Bartalinia
species and is sister to B. robillardoides Tassi (CBS
122705, ex-epitype strain). A comparison of ITS region
shows that B. kunmingensis is not significant different from
B. robillardoides (only two differentiated nucleotide
123
bases); however, B. kunmingensis is different from B.
robillardoides in 207/890 bp (23.2%) in RPB2 region. We
therefore, identify our isolate as a new species which was
found from corn (Zea mays) in China. Bartalinia kunmingensis differs from B. robillardoides (CBS H-21728) in
having smaller conidiomata and paler yellowish conidia.
(Crous et al. 2014a).
Robillarda Sacc.
The genus Robillarda was introduced by Saccardo
(1880a) and is typified by R. sessile Sacc. This genus
contains about 38 species (Crous et al. 2015a). The asexual
morph has been reported with its unique characteristics
such as solitary or gregarious, separate, subglobose, unicellular, immersed, ostiolate, glabrous pycnidia, holoblastic, ampulliform, hyaline, conidiogenous cells, originating
on the inner wall of the pycnidium and ellipsoidal, 1-septate, smooth-walled, hyaline conidia with single branched,
apical appendage (Crous et al. 2015a; Borse et al. 2016;
Wijayawardene et al. 2016). In this study, the new species,
Robillarda mangiferae is introduced from leaf blight on
mango in Yunnan, China.
Robillarda mangiferae Thiyag., Wanas., Phookamsak &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556210; Facesoffungi
number: FoF05718, Fig. 113
Etymology: The specific epithet ‘‘mangiferae’’ is based
on the host genus Mangifera, from which the taxon was
isolated.
Holotype: KUN-HKAS 102245
Associated with a leaf blight symptom on Mangifera.
Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata 250–310 lm high, 300–340 lm diam.,
black, pycnidial, semi-immersed to erumpent, solitary, scattered, irregular in shape, uni-loculate, glabrous, minutely
ostiolate, with beak-like papilla. Conidiomata walls 8–27 lm
wide, thin-walled, of unequal thickness, slightly thick at the
sides, composed of two types of cell layers, inner layers
comprising hyaline, flattened, pseudoparenchymatous cells of
textura angularis to textura prismatica; outer layers comprising dark brown to black, thick-walled, coriaceous cells, of
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 5–10 9 3–7 lm (
x = 6.9 9
4.8 lm, n = 30), holoblastic, proliferating percurrently 1–3
times, discrete, subcylindrical to ampulliform, hyaline, aseptate, smooth-walled, arising from the inner cavity. Conidia
(7.5–)10–11(–12) 9 (2.5–)3–4(–4.5)
lm
(
x = 10.9 9
3.5 lm, n = 50), hyaline, oblong to ellipsoidal, or subfusoid,
narrower towards the basal cell, straight, (0–)1-septate, thin
and smooth-walled, apical cell developed into a branched
appendage; appendages 25–35 9 1–2.5 lm (
x = 29.3 9
1.7 lm, n = 50), dividing into 2–3 branches, straight, non-
Fungal Diversity
Fig. 109 Phylogram generated from maximum likelihood analysis
based on a combined LSU, SSU and ITS gene regions of Amphisphaeriaceae and other related taxa. Bootstrap support values for
maximum likelihood (green), maximum parsimony (blue) equal to or
greater than 75% and the values of Bayesian posterior probabilities
(purple) equal to or greater than 0.95 BYPP are given above each
branch respectively. The new isolate is in blue. Ex-type strains are in
bold. The tree is rooted with Xylaria hypoxylon (CBS 122620)
123
Fungal Diversity
Fig. 110 Amphisphaeria mangrovei (AMH-9948, holotype). a Ascomata immersed in intertidal branches and twigs of Suaeda monoica. b, c
Vertical sections of ascomata. d Peridium. e Paraphyses. f–i Immature and mature asci. j–n Ascospores. Scale bars b, c = 50 lm, d–n = 10 lm
123
Fungal Diversity
Fig. 111 Phylogram generated from maximum likelihood analysis
based on ITS and LSU sequence dataset of the representative species
in Sporocadaceae. The updated sequence dataset was derived from
Wanasinghe et al. (2018). Sixty strains are included in the sequence
analyses. Phlogicylindrium eucalyptorum (CBS 111689) and
Phlogicylindrium uniforme (CBS 131312) are used as outgroup taxa.
Bootstrap support values for ML equal to or greater than 50% are
given above the nodes. Newly generated sequences are in blue. Extype strains are indicated in bold
123
Fungal Diversity
Fig. 112 Bartalinia kunmingensis (KUN-HKAS 102242, holotype).
a, b Conidiomata on host substrate. c Vertical section of conidioma.
d Conidioma wall. e–i Conidiogenous cells with conidia. j–m
Conidia. n Germinating conidium. Scale bars a, b = 200 lm,
c = 50 lm, d–m = 10 lm, n = 20 lm
flexuous, broadly tubular, narrower towards apex, inconspicuously septate at the apex.
Culture characteristics: Colonies on PDA reaching
80–85 mm diam. after 1 week at 20–25 °C, sparse to
medium sparse, circular, flat, surface slightly rough with
white tufts hyphae, or small granular, edge entire, floccose,
forming small, black pycnidia on colony and embedded in
media agar after 3 weeks; from above, white-grey to
greenish grey, from below, white to cream at the margin,
radiated with pale brown to black concentric ring at the
middle, white-grey at the centre; not producing pigmentation in agar.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Jinghong, Nabanhe, associated with leaf
blight symptom on living leaf of Mangifera sp., 21
November 2015, R. Phookamsak, XB011 (KUN-HKAS
102245, holotype), ex-type living culture, KUMCC
18-0180.
GenBank
numbers:
ITS = MK353084,
LSU =
MK353086, SSU = MK353149.
Notes: Robillarda mangiferae resembles Robillarda
species in having oblong to subfusoid, septate conidia, with
an apical cell modified into a branched appendage. However, R. mangiferae can be distinguished from other Robillarda species by its appendage being straight, nonflexuous, broad tubular and narrower towards the apex,
with inconspicuous septa at the apex, as well as its conidiogenous cells being holoblastic and proliferating percurrently. Robillarda species have flexuous, narrow
tubular, aseptate appendages and holoblastic conidiogenous cells, proliferating sympodially or percurrently near
the apex (Crous et al. 2015a; Wijayawardene et al. 2016).
Phylogenetic analyses of a combined ITS and LSU
sequence dataset show that R. mangiferae clusters with
other Robillarda species and Ellurema indica Nag Raj &
W.B. Kendr. [current name = Hyalotiopsis Punith.,
123
Fungal Diversity
proposed by Réblová et al. (2016), and Wijayawardene
et al. (2016)] in Sporocadaceae, but the species forms a
distinct lineage at the base of this clade with moderate
support (92% ML and 1.00 BYPP; Fig. 111). Robillarda
mangiferae differs from Ellurema indica in having oblong
to ellipsoidal, or subfusoid, (0–)1-septate conidia, with
straight, non-flexuous, broadly tubular apical appendages.
The asexual morph of Ellurema indica (= Hyalotiopsis) has
cylindrical to fusiform or obclavate conidia, with more
than 1-septate and 2–3 apical bi- or tri-furcate, filiform,
flexuous appendages (Wijayawardene et al. 2016, 2017a).
Based on morphological characteristics and phylogenetic
analyses of a combined ITS and LSU sequence dataset
(Fig. 111), we hence, introduce a new species, R. mangiferae in this study.
Xylariales Nannf.
Diatrypaceae Nitschke
We follow the latest treatment and updated accounts of
Diatrypaceae in de Almeida et al. (2016), Shang et al.
(2017, 2018) and Senwanna et al. (2017). A higher level
classification with divergence time estimates for Diatrypaceae was provided by Hongsanan et al. (2017) and
placed the family in order Xylariales (Xylariomycetidae,
Sordariomycetes) and a similar scheme is followed in
Wijayawardene et al. (2018a). A novel species, Peroneutypa mangrovei is introduced in Diatrypaceae based on
analysis of a combined ITS and TUB2 sequence dataset
(Fig. 114), coupled with morphological characteristics. In
addition, the new genus Neoeutypella is introduced as a
monotypic genus to accommodate N. baoshanensis. The
new genus was collected from dead wood in Baoshan,
China. The sexual and asexual morphs are described and
illustrated.
Neoeutypella M. Raza, Q.J. Shang, Phookamsak & L. Cai,
gen. nov.
Index Fungorum number: IF555373; Facesoffungi
number: FoF04927
Etymology: The generic epithet ‘‘Neoeutypella’’ refers to
the taxon resembling Eutypella.
Saprobic on dead wood of Pinus armandii. Sexual
morph Ascostromata entostromatic, carbonaceous, visible
as black, solitary to gregarious, globose to long irregular in
shape on host surface, erumpent through host epidermis,
producing yellow pigments surrounding ascostroma. Ascomata perithecial, black, immersed to semi-immersed in
stromatic tissues, aggregated, globose or subglobose, ostiolate, papillate, slightly conspicuous, with periphyses.
Peridium thickened unequally, two-layered, outer layer
comprising 5–7 layers of thick-walled, hyaline to dark
brown cells of textura angularis, inner layer comprising
3–5 layers of thin-walled, hyaline to brown cells of textura
prismatica. Hamathecium comprising aseptate, filamentous
paraphyses, tapering toward the apex, embedded in hyaline
gelatinous matrix. Asci 8-spored, unitunicate, spindleshaped, long pedicellate, apically rounded with refractive
cytoplasmic strands, amyloid, with a J ? , subapical ring.
Ascospores overlapping 1–3-seriate, allantoid, slightly or
moderately curved, initially hyaline, becoming pale brown
at maturity, aseptate, mostly with small 1–2 guttules.
Asexual morph Hyphae branched, smooth, hyaline, septate. Conidiophores long, branched, with phialides,
mononematous, macronematous, hyaline. Conidiogenous
cells smooth-walled, hyaline, holoblastic, discrete, phialidic, doliiform, ampulliform or irregular in shape. Conidia
filiform, solitary, aseptate, smooth-walled, unbranched,
hyaline to pale yellow.
Type species: Neoeutypella baoshanensis M. Raza, Q.J.
Shang, Phookamsak & L. Cai
Notes: Neoeutypella resembles Eutypella (Nitschke)
Sacc. in forming large entostroma, 8-spored, spindleshaped asci and allantoid ascospores, with a libertella-like
asexual morph. Phylogenetic analyses of maximum likelihood, maximum parsimony and Bayesian inference based
on the combined ITS and TUB2 sequence dataset
(Fig. 114) show that Neoeutypella baoshanensis groups
with Eutypella caricae (De Not.) Berl. (strains EL51C and
GL08362). Sequences of these two strains are available in
GenBank, but no morphological description is available for
comparative studies. The two Eutypella caricae strains and
Neoeutypella form a distinct lineage from Eutypella sensu
stricto. Acero et al. (2004) mentioned that these two strains
might have been misidentified and a taxonomic revision of
these species is needed. Eutypella caricae and Neoeutypella are phylogenetically closely related to Diatrypella
banksiae Crous which produced an asexual morph (Crous
et al. 2016a). Nevertheless, Neoeutypella can be differentiated from D. banksiae in shape and size of conidia.
Neoeutypella baoshanensis (see below) has filiform conidia
[(16.5–)25–37(–40) 9 1.2–1.9 lm], whereas Diatrypella
banksiae has spindle-shaped conidia [(25–)27–30(–
35) 9 1.5(–2) lm] (Crous et al. 2016a).
Based on morphological comparison of our new taxon
and Eutypella caricae described by Saccardo (1882) and
Berlese (1902), Neoeutypella baoshanensis (99% similarity
in ITS, 90% similarity in TUB2) differs from Eutypella
caricae in having larger asci (N. baoshanensis, (45–
)52–110(–125) 9 (19–)26–37(–40) versus 35–45 9 6–7,
E. caricae) and ascospores (N. baoshanensis, (30–)35–43(–
50) 9 (8–)9–11(–12) versus 9–11 9 2.5–3, E. caricae)
(Saccardo 1882; Berlese 1902). Neoeutypella baoshanensis
has spindle-shaped asci and pale yellowish to pale brown
ascospores, whereas E. caricae has clavate asci and hyaline
ascospores (Saccardo 1882; Berlese 1902). Based on phylogenetic support coupled with morphological differences,
123
Fungal Diversity
Fig. 113 Robillarda mangiferae (KUN-HKAS 102245, holotype).
a Substrate showing disease symptoms. b, c Conidiomata on
substrate. d Section through conidioma. e Conidioma wall. f–h
Conidiogenous cells attached to conidia. i–n Conidia. Scale bars b–
d = 100 lm, e–h, i, m, n = 10 lm, j–l = 5 lm
we therefore introduce our isolate as a new species in the
new genus Neoeutypella.
angularis, inner layer comprising 3–5 layers of thin-walled,
hyaline to brown cells of textura prismatica. Hamathecium
3–7 lm wide (
x = 4.7 lm, n = 20), comprising aseptate,
filamentous paraphyses, tapering towards the apex, embedded in hyaline gelatinous matrix. Asci (60–)75–85(–90) 9
(5.5–)6.5–7.5(–8) lm (
x = 77.5 9 8 lm, n = 25), 8-spored,
unitunicate, spindle-shaped, long pedicellate, apically rounded with refractive cytoplasmic strands, amyloid, with a
J ? , subapical ring. Ascospores (8.5–)10–11.5(–13) 9
(2–)2.3–2.5(–3) lm (
x = 10.8 9 2.4 lm, n = 50), overlapping 1–3-seriate, initially hyaline, becoming pale brown at
maturity, allantoid, slightly or moderately curved, aseptate,
mostly with 1–2 small guttules. Asexual morph Hyphae
branched, smooth, hyaline, septate, 1.5–3.5 lm diam.
Conidiophores long, branched, with phialides, mononematous, macronematous, hyaline. Conidiogenous cells (12–
)14–35.5(–40) 9 4–13(–15)
lm
(
x = 25 9 8.5 lm,
n = 20), smooth-walled, hyaline, holoblastic, discrete,
phialidic, doliiform, ampulliform or irregular in shape. Conidia (16.5–)25–37(–40) 9 1.2–1.9 lm (
x = 29 9 1.5 lm,
Neoeutypella baoshanensis M. Raza, Q.J. Shang,
Phookamsak & L. Cai, sp. nov.
Index Fungorum number: IF555372; Facesoffungi
number: FoF04928, Fig. 115
Etymology: The specific epithet ‘‘baoshanensis’’ refers
to the locality Baoshan (Yunnan, China), where the holotype was collected.
Holotype: HMAS 255436
Saprobic on dead wood of Pinus armandii. Sexual morph
Ascostromata 650–1100 lm diam., entostromatic, carbonaceous, black, solitary to gregarious, globose to long irregular
in shape on host surface, erumpent through host epidermis,
producing yellow pigments surrounding ascostroma. Ascomata perithecial, 500–770 high, 450–530 diam., black,
immersed to semi-immersed in stromatic tissues, aggregated,
globose or subglobose, ostiolate, papillate, slightly conspicuous, with periphyses. Peridium 145–250 wide, thickened
unequally, two-layered, outer layer comprising 5–7 layers of
thick-walled, hyaline to dark brown cells of textura
123
Fungal Diversity
n = 50), filiform, solitary, aseptate, smooth-walled, unbranched, hyaline to pale yellow.
Culture characteristics: Colonies on PDA reaching
6.5–7 mm diam. after 1 week at 25 ± 2 °C, circular, flat,
slightly raised, surface dull with edge undulate, filamentous
at the margin; from above, white with cotton consistency;
from below, pale yellow and not producing pigment in
PDA medium. Asexual morph produced on PDA after
3 weeks and colony becomes black from below.
Material examined: CHINA, Yunnan Province, Baoshan
City, Longling County, on dead wood of Pinus armandii
Franch. (Pinaceae), October 2015, M. Raza, HMAS
255436 (holotype), ex-type living culture, LC 12111.
GenBank numbers: ITS = MH822887, TUB2 =
MH822888.
Peroneutypa Berl.
We follow the latest treatment and updated accounts of
Peroneutypa in Senwanna et al. (2017) and Shang et al.
(2018).
Peroneutypa mangrovei Devadatha & V.V. Sarma, sp.
nov.
Index Fungorum number: IF554285; Facesoffungi
number: FoF04271, Fig. 116
Etymology: Specific epithet in reference to the habitat.
Holotype: AMH-9944
Saprobic on decaying wood of Avicennia marina. Sexual morph Ascostromata absent or poorly developed
between perithecial necks, perithecia solitary to gregarious,
up to four in groups, dark brown to black, immersed
becoming raised to erumpent through the host tissue with
median necks. Ascomata 250–525 lm high, 100–330 lm
diam. (
x = 375 9 202 lm, n = 10), erumpent to
immersed, globose to subglobose, gregarious to solitary,
ostiolate, with short beaks, periphysate, brown to black.
Ostiolar canals 50–85 lm wide (
x = 67 lm, n = 5), with
moderate neck length 100–350 lm (
x = 231 lm, n = 5),
cylindrical, straight, dark brown to black. Periphyses filamentous, short, 0.5–2 lm wide (
x = 1.5 lm, n = 10).
Peridium 15–35 lm wide (
x = 22 lm, n = 10), comprising
two layers, inner stratum with many layers of hyaline cells
of textura angularis and outer stratum with 2–3 layers of
light brown to black cells of textura angularis. Hamathecium composed of numerous, 1–2 lm (
x = 1 lm, n = 20)
wide, filamentous, septate paraphyses, longer than asci,
embedded in a gelatinous matrix. Asci 14–20 9 3–4 lm
(
x = 17 9 3.5 lm, n = 20), 8-spored, unitunicate, cylindrical to clavate, short pedicellate, apically rounded to
truncate, with a J-, apical ring. Ascospores 3–5 9 1–
1.5 lm (
x = 4 9 1 lm, n = 30), overlapping 1–3-seriate,
hyaline to pale yellow, straight to allantoid, aseptate,
smooth-walled, lacking guttules, light brown in mass.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching
50–80 mm diam. after 15 days of incubation at room
temperature, initially white, becoming light grey at maturity, reverse light yellow, cottony, punctiform, flat, circular,
entire.
Material examined: INDIA, Puducherry, Thengaithittu
mangroves (11.5°N 79.5°E), on decaying wood of Avicennia marina (Forssk.) Vierh. (Acanthaceae), 12 March
2016, B. Devadatha (AMH-9944, holotype), ex-type living
culture, NFCCI-4246.
GenBank
numbers:
ITS = MG844286,
LSU =
MG844278, SSU = MG844282, TUB2 = MG844282.
Notes: Peroneutypa mangrovei shares similar morphological characters such as J-, apical ring asci, overlapping
ascospores dimensions with P. diminutispora D.A.C.
Almeida et al. (de Almeida et al. 2016). However, it differs in
having perithecia single or in groups of up to four, shorter
ascomata and ostiole, longer asci (14–20 9 3–4 lm) and its
exclusive occurrence in marine habitat. Peroneutypa diminutispora has perithecia arranged in a single layer, occurring
singly or in groups of up to seven, shorter asci
(8–14 9 3.5–5 lm) that are urn-shaped (de Almeida et al.
2016). Peroneutypa cosmosa (Speg.) Carmarán & A.I.
Romero has larger asci (18–25 9 5–7 lm) with J?, apical
rings and longer ascospores (6–8 lm) (Carmarán et al. 2006).
Peroneutypa longiasca Senwanna et al. and P. mackenziei
Q.J. Shang et al. have longer ascospores (5–7 lm) (Senwanna et al. 2017; Shang et al. 2017). This is the first report of
Peroneutypa species from a marine habitat (Jones et al.
2015). ITS and TUB2 gene phylogeny also supports P.
mangrovei as distinct forming an independent lineage, sister
to Eutypa microasca E. Grassi & Carmarán, and cluster with
Peroneutypa diminutispora and P. cosmosa (Fig. 114). A
comparison of ITS nucleotides between P. mangrovei and P.
dimunitispora (GenBank no. KM396647) results in 7.4% (47
nucleotides) and 5.5% (26 nucleotides) along with P. cosmosa (GenBank no. KF964568).
Hypoxylaceae DC.
We follow the latest treatment and updated accounts of
Hypoxylaceae in Daranagama et al. (2018) and Wendt et al.
(2018).
Hypoxylon Bull.
Hypoxylon is the largest genus in Hypoxylaceae with
over 140 accepted species classified on the basis of morphology, phylogeny and chemotaxonomy (Kuhnert et al.
2014; Wijayawardene et al. 2017a). More than 1000 epithets are listed in Index Fungorum (2019). We introduce a
new species Hypoxylon teeravasati based on morphology
and multigene analysis (Fig. 117).
Hypoxylon teeravasati Devadatha, V.V Sarma & E.B.G
Jones, sp. nov.
123
Fungal Diversity
123
Fungal Diversity
b Fig. 114 Phylogram generated from maximum likelihood, maximum
parsimony and Bayesian inference analyses based on a combined ITS
and TUB2 sequence dataset of diatrypaceous species. Bootstrap
support values for maximum likelihood (black) and maximum
parsimony (green) equal to or greater than 60% and Bayesian
posterior probabilities (blue) equal to or greater than 0.95 BYPP are
shown above the nodes. The new isolates are in blue and ex-type
strains are in bold. The tree is rooted to Lopadostoma turgidum (LT2
and CBS 133207)
Index Fungorum number: IF554278; Facesoffungi
number: FoF04272, Fig. 118
Etymology: The specific epithet ‘‘teeravasati’’ in Sanskrit, refers to the coastal environment where the fungus
thrives
Holotype: AMH-9906
Colour codes follow: A mycological colour
chart (Rayner 1970).
Saprobic on decaying wood of mangrove trees. Sexual
morph Ascostromata 310–770 9 400–655 lm diam.
(
x = 533 9 498 lm, n = 10), glomerate to hemispherical,
effuse-pulvinate, connected to each other by thick stromatal tissue at the base, with very conspicuous perithecial
mounds, surface burnt sienna (plate 7, D8), with KOH
extractable pigments eye brown (plate 7, F6), the tissue
below the perithecial layer inconspicuous. Perithecia 300–
650 lm diam. (
x = 442 lm, n = 10), immersed, sphaerical. Ostioles papillate, without apparent disk formation.
Peridium 45–100 lm (
x = 68 lm, n = 10), comprising
several layers of textura angularis and outer layer of
pseudoparenchyma. Hamathecium composed of 1–3 lm
Fig. 115 Neoeutypella
baoshanensis (HMAS 255436,
holotype). a Blackish
ascostromata surrounded by
yellow pigments on Pinus
armandii. b Vertical section of
ascostroma. c Ostiole. d, e
Peridial structure. f Paraphyses
stained with Indian ink. g Asci
attached with paraphyses
stained with Indian ink.
h Immature ascus. i–k Mature
asci. l J?, apical ring stained
with Melzer’s reagent. m–q
Ascospores. r Germinated
ascospore. s, t Culture
characteristics on PDA
(s = from above, t = from
below). u Conidiogenous cells
with conidia. v Phialides with
young developing conidia. w, x
Conidiogenous cells. y Conidia.
Scale bars b = 1000 lm, c,
d = 50 lm, e–g = 20 lm, h–k,
r, u, y = 10 lm, l–q = 5 lm
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Fungal Diversity
diam., hyaline, aseptate and filiform paraphyses, longer
than asci. Asci 65–160 9 7–13 lm (
x = 105 9 8.2 lm,
n = 20), 8-spored, unitunicate, cylindrical, pedicellate,
with apical ring bluing in Lugol’s solution, discoid, 1–
2.5 9 2–3.5 lm (
x = 1.9 9 3.1 lm, n = 20). Ascospores
9–15 9 4–7 lm (
x = 11.7 9 7.6 lm, n = 30), 1-seriate,
unicellular, ellipsoid-inequilateral, with narrowly rounded
ends, brown to dark brown, with a straight germ slit more
or less running the entire spore-length, perispore dehiscent
in 10% KOH, epispore smooth. Asexual morph
Undetermined.
Culture characteristics: Ascospores germinating on
SWA within 24 h, germ tubes arising from terminal ends of
the ascospore. Colonies on MEA at room temperature
reaching 45–60 mm diam. within 25 days, honey yellow at
centre and olive buff at margin, reverse clove brown with
yellow exudates and dawn grey at margin, honey yellow
diffusible pigments, filiform, umbonate, circular and
velvety.
Material examined: INDIA, Tamil Nadu, Tiruvarur,
Muthupet mangroves (10.4°N 79.5°E), on decaying wood
of Avicennia marina (Acanthaceae), 28 November 2015,
B. Devadatha, AMH-9906 (holotype), ex-type living culture, NFCCI-4216; ibid. on decaying branches and twigs of
Suaeda monoica Forssk. ex J.F. Gmel, PUFD4 (paratype
at Pondicherry University, Puducherry).
GenBank
numbers:
ITS = KY863509,
LSU =
MF385274, SSU = MF385273, RPB2 = MG986895,
TEF1-a = MF182401, TUB2 = MG986894.
Notes: Multigene analysis shows that Hypoxylon teeravasati shares a strongly supported sister relationship with
H. jaklitschii Sir & Kuhnert and H. lenormandii Berk. &
M.A. Curtis as a basal taxon. All three taxa constitute a
single monophyletic clade with high support (Fig. 117), but
their relationships with other Hypoxylon species are not
well-resolved. We found distinct nucleotide base pair differences between H. teeravasati and H. jaklitschii across
ITS and TUB2 genes [36 within ITS and 23 within TUB2]
which is in agreement with guidelines by Jeewon and Hyde
(2016) to establish new species. Hypoxylon teeravasati, H.
lenormandii, H. jaklitschii and H. croceum J.H. Mill. share
similar stromatal characters and overlapping asci dimensions. However, H. teeravasati can be easily distinguished
from H. lenormandii, H. jaklitschii and H. croceum in
having a burnt sienna surface, with KOH extractable pigments eye brown and larger perithecia. Hypoxylon lenormandii has surface greyish sepia, fuscous, or brown
vinaceous, dull orange brown to dark brown granules
beneath
and
between
perithecia,
with
KOH
extractable pigments hazel, fulvous, umber or ochreous
(Kuhnert et al. 2015; Liu et al. 2015a). Hypoxylon jaklitschii has a sepia to dark brick surface, pruinose, with
orange brown or dark brown granules immediately beneath
123
the surface and between perithecia, with KOH
extractable pigment umber or dark brick and smaller
ascospores (9.5–12 9 4–5.5 versus 9–15 9 4–7) (Kuhnert
et al. 2015). Hypoxylon croceum is distinguished from H.
teeravasati in having an initially sepia or fuscous surface,
becoming dark brown at maturity and abelline or hazel
KOH extractable pigments (Miller 1933). Hypoxylon croceum is the only report of Hypoxylon species from a marine
habitat (Jones et al. 2015). Hypoxylon teeravasati is
saprobic on decaying branches and twigs of Avicennia
marina and Suaeda monoica, exclusively from marine
environments. Hypoxylon lenormandii and H. jaklitschii
have been reported from terrestrial habitats.
Phylum Basidiomycota R.T. Moore
We follow the latest treatment of Basidiomycota in
Zhao et al. (2017).
Class Agaricomycetes Doweld
The classifications of the families in Agaricomycetes
herein follow Hibbett et al. (2014) and Zhao et al. (2017).
The subclasses, orders and families of Agaricomycetes are
listed in alphabetical order.
Subclass Agaricomycetidae Parmasto
Agaricales Underw.
Agaricaceae Chevall.
We follow the latest treatments and updated accounts of
Agaricaceae in Zhao et al. (2016), Zhou et al. (2016) and
Hyde et al. (2017). Three specimens were collected and
preliminary BLAST using ITS sequence data indicated that
two taxa belong to Agaricus sect. Xanthodermatei and the
third to Coprinus. Based on distinctive morphological
characteristics and phylogenetic support, two novel species
of Agaricus sect. Xanthodermatei as well as Coprinus trigonosporus sp. nov. are introduced in this study. An
updated phylogenetic tree based on maximum likelihood
and Bayesian inference analyses also confirms their
placement (Fig. 119).
Agaricus L.: Fr.
We follow the latest treatment and updated accounts of
Agaricus in Thongklang et al. (2014), Karunarathna et al.
(2016), Zhao et al. (2016) and Zhou et al. (2016). Detailed
taxonomic revision of taxa in Agaricus sect. Xanthodermatei was discussed by Chen et al. (2016), Kerrigan
(2016), Mahdizadeh et al. (2016), Zhao et al. (2016) and
Parra et al. (2018).
Agaricus memnonius M.Q. He & R.L. Zhao, sp. nov.
Fungal names: FN570535; Faceoffungi number:
FoF03940, Fig. 120
Etymology: The Latin epithet ‘‘memnonius’’ meaning
‘‘brown-black’’ refers to the colour of the pileus surface.
Fungal Diversity
Fig. 116 Peroneutypa mangrovei (AMH-9944, holotype). a Ascomata semi-immersed in the decaying wood of Avicennia marina. b–d
Longitudinal sections of ascoma. e Section through peridium.
f Paraphyses interspersed with asci. g–l Immature and mature asci.
m–q Ascospores. Scale bars b–d = 100 lm, e–q = 10 lm
123
Fungal Diversity
Fig. 117 Phylogram generated from maximum likelihood analysis
based on ITS, LSU, RPB2 and TUB2 gene regions of Hypoxylaceae
and related taxa in Xylariales. Bootstrap support values for maximum
likelihood (green), maximum parsimony (blue) equal to or greater
123
than 75% and the values of Bayesian posterior probabilities (purple)
equal to or greater than 0.95 are given above each branch
respectively. The new isolate is in blue. Ex-type strains are indicated
in bold. The tree is rooted with Xylaria hypoxylon (CBS122620)
Fungal Diversity
123
Fungal Diversity
b Fig. 118 Hypoxylon teeravasati (AMH-9906, holotype). a Ascostro-
mata on the decaying wood of Avicennia marina. b Cross section of
ascostroma. c, d Longitudinal sections of ascostroma. e Peridium.
f Filamentous paraphyses. g–k Immature and mature asci (g–i = Asci
showing apical bluing in Lugol’s solution). l Immature ascospore. m–
o, q Mature ascospores. p Mature ascospore with dehiscent perispore
in KOH. r Germinating ascospore. s KOH extractable pigments. Scale
bars c, d = 100 lm, e–r = 10 lm
Holotype: HMAS 0278359
Pileus 50 mm diam., plane or plano-concave, disc black or
black-brown, slightly depressed, margin straight, exceeding
lamellae; surface dry, with black-brown fibrillose scales
against white background, scales triangular, appressed,
extremely denser at disc, scattered towards the margin.
Lamellae up to 3 mm broad, free, crowded, pink, edge even,
intercalated with lamellulae. Annulus superous, double,
membranous, white, pendant, upper side smooth, lower side
cogwheel, white, edge light brown. Stipe 57 9 5 mm (8 mm
at base), white, hollow, cylindrical with slightly bulbous
base, surface dry, smooth, silky, with rhizomorphs. Context
fleshy, white. Odour unknown. KOH reaction: positive yellow. Schäffer’s reaction: negative. Basidia 15–19.5 9
6.5–9.3 lm, clavate, hyaline, 4-spored, smooth. Basidiospores 4.5–5.3 9 3.3–4.1 lm, (
x = 5 ± 0.2 9 3.6 ± 0.2,
Q = 1.2–1.5, Qm = 1.4 ± 0.1, n = 20), ellipsoid, smooth,
thick-walled, brown. Pleurocystidia absent. Cheilocystidia
not very conspicuous, can be single and multiseptate (generally no more than three elements), the terminal element
clavate, cylindrical, 12.1–24.8 9 6.9–13.7 lm. Pileipellis a
cutis composed of hyphae of 4.6–14.4 lm diam., smooth,
cylindrical, slightly constricted at septa, pigment intracellular, light brown or brown.
Material examined: CHINA, Sichuan Province, Miyi
County, 13 September 2015, ZRL20151118 (HMAS
0278359, holotype).
Host and habitat: Solitary on soil in forest with bamboo
around.
Distribution: Sichuan Province (China).
GenBank
numbers:
ITS = MG763128,
LSU =
MG765263, TEF1-a = MG765265.
Notes: The phylogenetic trees generated by maximum
likelihood and Bayesian inference analyses (Fig. 119) show
that Agaricus memnonius forms a distinct lineage within
Agaricus sect. Xanthodermatei in the clade Xan II and the
tree topology is similar with previous studies (Zhao et al
2016; Zhou et al. 2016; Parra et al. 2018).
Agaricus memnonius has relatively small and slender
basidiomes, which are similar to the European species A.
laskibarii L.A. Parra & Arrillaga, A. xanthodermulus Callac
& Guinb. and A. parvitigrinus Guinb. & Callac (Parra 2013).
However, A. memnonius has black-brown and triangular
scales on the pileus, while the other three species have
greyish brown and not triangular scales. Some species
123
described from China also have small and slender basidiomes, such as A. gregariomyces J.L. Zhou & R.L. Zhao and
A. karstomyces R.L. Zhao. However, A. gregariomyces has
larger and elongate basidiospores (5.6–6.3 9 3.5–4.0 lm,
Q = 1.6–1.9; Zhou et al. 2016). Agaricus karstomyces has
dot-like scales on the pileus, while A. memnonius has triangular scales. Based on phylogenetic and morphological
studies, A. memnonius is introduced as a new species and is
characterized by its distinct phylogenetic position in section
Xanthodermatei, small and slender basidiome, black-brown
and triangular scales on the pileus, and the single and multiseptate cheilocystidia.
Agaricus langensis M.Q. He & R.L. Zhao, sp. nov.
Fungal names: FN570534; Facesoffungi number:
FoF03941, Fig. 121
Etymology: The Latin epithet ‘‘langensis’’ meaning
‘‘originating from Lang’’ refers to the Lang County where
the holotype was collected.
Holotype: HMAS 0278317
Basidiomes flavescent when rubbed. Pileus 26–49 mm
diam., parabolic when young, then convex, disc slightly
subumbonate, margin straight, sometimes with appendiculate remains of veil; surface dry, with grayish brown fibrillose
scales against white background, scales appressed, covering
the whole pileus, denser at disc, scattered towards the margin.
Lamellae up to 5 mm broad, free, crowded, pink, edge even,
intercalated with lamellulae. Annulus up to 6 mm in diam.,
superous, double, membranous, white when fresh, yellowish
when dry, pendant, upper side smooth, lower side cogwheel,
white, edge light brown. Stipe 59–76 9 6–7 mm (7–10 mm
at base), white, hollow, cylindrical, surface dry, smooth or
slightly fibrillose, with rhizomorphs. Context fleshy, white.
Odour unknown. KOH reaction: positive yellow. Schäffer’s
reaction: negative. Basidia 18.5–25.3 9 6.7–8.9 lm, clavate,
hyaline,
4-spored,
smooth.
Basidiospores
6.3–8.3(–8.5) 9 3.7–5.1 lm, (
x = 7.2 ± 0.6 9 4.4 ± 0.3,
Q = 1.4–1.9, Qm = 1.6 ± 0.1, n = 20), ellipsoid, elongate,
smooth, thick-walled, brown. Pleurocystidia absent.
Cheilocystidia absent. Pileipellis a cutis composed of hyphae
of 4.8–12.1 lm diam., smooth, cylindrical, slightly constricted at septa, hyaline, light brown or brown.
Material examined: CHINA, Tibet, Lang County,
Gongga Village, Alt. 3384 m, 29°160 N, 93°110 E, S-Y. Su,
ZRL20152282 (HMAS 0278317, holotype).
Habit and habitat: Scattered on soil in forest.
Distribution: Tibet (China).
GenBank
numbers:
ITS = MG763129,
LSU =
MG765264, TEF1-a = MG765266.
Notes: Agaricus langensis (HMAS 0278317) belongs to a
clade called Xan III (Thongklang et al. 2014; Parra et al. 2018)
which is present (but not always indicated) in all previous
multigene trees of Agaricus sect. Xanthodermatei (Zhao et al.
Fungal Diversity
Fig. 119 Maximum likelihood tree of Agaricus sect. Xanthodermatei
based on LSU, TEF1-a and ITS sequences with Agaricus campestris
(LAPAG370) as the outgroup taxon. The bootstrap values and
Bayesian posterior probabilities more than 50%/0.90 (BS/BYPP) are
indicated at the nodes. The branches in bold mean the related
PP [ 0.95, ‘‘T’’ refers to sequences from type specimen
2016, Zhou et al. 2016, Parra et al. 2018) and which includes in
our tree A. flavidodiscus L.A. Parra et al., A. langensis and 32
other species (Fig. 119). The new species forms a sister lineage with Agaricus sp. (ZRL2012629) with moderate support
(75% ML and 1.00 BYPP) in our phylogenetic analyses
(Fig. 119). Agaricus parvitigrinus Guinb. & Callac resembles
A. langensis in the field, because both have small to medium
sized basidiomes, a convex pileus covered by greyish fibrillose scales. However, A. langensis has larger basidiospores
than those of A. parvitigrinus (5.8 9 3.7 lm; Parra 2013).
Agaricus menieri Bon, A. xanthodermulus Callac & Guinb.
and A. xanthodermus Genev. have similar sized basidiospores,
but have distinct cheilocystidia (Parra 2013), a feature lacking
in A. langensis. Agaricus tibetensis J.L. Zhou & R.L. Zhao
also has small to medium sized basidiomes, same sized
basidiospores as A. langensis, and no cheilocystidia (or rare in
A. tibetensis), and both species originated from Tibet (Zhou
et al. 2016). However, their ITS sequences differ at 33 base
positions, and the molecular phylogeny also indicates that
they are different species.
Based on the phylogenetic analyses and morphological
characteristics, we introduce this new species, which is
123
Fungal Diversity
Fig. 120 Agaricus memnonius
(HMAS 0278359, holotype).
a Basidiome in the field.
b Basidiospores.
c Cheilocystidia. d Pileipellis
hyphae. e Basidia. Scale bars
a = 2 cm, c–e = 10 lm,
b = 5 lm
characterized by its distinct phylogenetic position in
sect. Xanthodermatei, small to medium sized basidiomes,
relatively large basidiospores and absence of cheilocystidia.
Coprinus Pers.
Traditionally, Coprinus comprised all coprinoid species
(black spore print and mature lamellae, plicate-sulcate
pileus) and it was the type genus of the family Coprinaceae
Overeem & Weese. On the basis of earlier phylogenetic
123
studies, Redhead et al. (2001) transferred most Coprinus
species into four genera of the new family Psathyrellaceae
Vilgalys et al. Coprinus comatus (O.F. Müll.) Pers., and a
few related species remained in Coprinus, which was
transferred to family Agaricaceae (Moncalvo et al. 2002;
Vellinga 2004). Coprinus species have rather small to large
basidiomes with squamulose pileus, ring-like partial veil
and deliquescent lamellae. A phylogenetic tree is presented
in Fig. 122.
Fungal Diversity
Coprinus trigonosporus Tkalčec & Mešić, sp. nov.
MycoBank number: MB826852; Facesoffungi number:
FoF05719, Fig. 123
Etymology: The species is named after its basidiospores
that are often rounded triangular in frontal view.
Holotype: CNF 1/6594
Pileus up to 24 mm broad and 32 mm high when still
closed, 32–46 mm broad at maturity, ellipsoid to oblong at
first, later obtusely conical to subapplanate with broad subumbonate centre, often radially splitting, when young
entirely covered with a dense, white universal veil forming
imbricate, rather large scales with mostly upturned lower
edge, later veil splitting up into patches of different sizes
(from one scale to group of scales) except at the centre,
surface strongly plicate-sulcate, white at first, becoming pink
brown to vinaceous brown and soon black, margin sometimes
with few small, white, appendiculate remnants of partial veil
at maturity. Lamellae free, crowded, broad, often sinuous
when young, white at first, later turning pink to vinaceous
brown, soon becoming black, weakly deliquescent. Stipe
34–56 9 5.5–10 mm, tapering upwards or subcylindrical,
with subbulbous, obconical, buried base (up to 15 mm wide),
white, central, hollow, dry, ± tomentose when young, (sub)
glabrous at maturity, partial veil remaining as a white, narrow
ring at the top of the base. Context white. Odour and taste
fungoid. Spore print black. Basidia 30–62 9 12–24 lm,
clavate, 4-spored, thin-walled or moderately thick-walled
(up to 1 lm), first hyaline, at maturity partially (in the middle
part) or entirely with pale to dark brown parietal pigment,
surrounded by 6–12 hymenophysalides (pseudoparaphyses).
Hymenophysalides 15–42 9 6–14 lm, clavate, thin-walled,
sometimes moderately thick-walled (up to 0.8 lm), hyaline
to pale brown. Basidiospores [500/5/1] (9.3–)10.7–14.6–
18.6(–20.8) 9 (8.1–)9.6–11.8–14.3(–15.1) 9 (7.8–)8.5–10–
11.6(–12.3) lm, averages of different basidiomes
13.9–15.3 9 11.7–12.2 9 9.8–10.3 lm, Qf = 0.9–1.2–1.7,
Qs = 1.2–1.4–1.8, av. Qf = 1.1–1.3, av. Qs = 1.4–1.5,
rounded triangular, cordiform, ovoid or subglobose in frontal
view (rarely ellipsoid), with rounded to flat base and rounded
apex, ovoid to ellipsoid in side view, flattened, smooth,
moderately thick-walled (up to 1 lm), with 1.5–3.5 lm
wide, slightly to strongly eccentric, rarely central germ-pore
(sometimes ring-like protruding), medium to dark brown in
H2O and KOH, semitransparent, non-amyloid and nondextrinoid.
Pleurocystidia
absent.
Cheilocystidia
16–58 9 11–28 lm, broadly to narrowly clavate, thin-walled, hyaline. Pileipellis a cutis, composed of repent, thinwalled, 5–15 lm wide, subhyaline to brown hyphae with
intracellular pigment. Veil cells on the pileus
12–170 9 2–38 lm, cylindrical to inflated, in chains, often
constricted at the septa, occasionally branched, sometimes
with individual, simple, subglobose to finger-like excrescences, thin-walled, rarely moderately thick-walled (up to
Fig. 121 Agaricus langensis (HMAS 0278317, holotype). a Basidiomes in the laboratory. b Basidiospores. c Basidia. d Pileipellis
hyphae. Scale bars a = 2 cm, c, d = 10 lm, b = 5 lm
0.8 lm), smooth, rarely finely encrusted, hyaline. Stipitipellis a cutis of parallel, repent, thin-walled, hyaline,
1.5–10 lm wide hyphae. Clamp connections absent; pseudoclamps present on some septa in veil, pileipellis and trama.
Material examined: SAUDI ARABIA, Jizan Province,
village, 8 km S from Harub, 17°220 0300 N, 42°520 1000 E,
257 m a.s.l., on sandy soil in a courtyard, 22 September
2010, leg. M. Čerkez, CNF 1/6594 (holotype).
Habit and habitat: In group (approximately 20 basidiomes), on sandy soil with some sheep dung, among scattered herbaceous plants and bushes in a courtyard.
Distribution: So far known only from the type locality in
the Kingdom of Saudi Arabia, Jizan Province.
GenBank
numbers:
ITS = MH422561,
LSU = MH422563.
Notes: Based on a megablast search of NCBIs GenBank
nucleotide database, the closest hit using ITS sequence for
our new species is Coprinus vosoustii Pilát [GenBank no.
JF907844, similarity = 627/668(94%), Gaps = 9/668(1%)].
Phylogenetic analysis based on the ITS sequence data of
Coprinus sensu stricto shows that C. trigonosporus formed a
distinct lineage and clustered with C. vosoustii and
123
Fungal Diversity
C. sterquilinus (Fr.) Fr.. Morphologically, C. trigonosporus
is best characterised by a considerable number of cordiform
and rounded triangular basidiospores. Other species in the
genus have ellipsoid or ovoid basidiospores (van de Bogart
1976; Moreno and Heykoop 1998; Cacialli et al. 1999; Uljé
2005; Crous et al. 2016b). The rather peculiar microscopic
characteristic of C. trigonosporus is the large number of
hymenophysalides (6–12) surrounding the basidium.
Amanitaceae E.-J. Gilbert
Amanitaceae is defined as those agarics having bilateral,
divergent lamellae trama and a longitudinally acrophysalidic stipe context. The family comprises five genera,
Amanita Pers., Catatrama Franco-Mol., Limacella Earle,
Limacellopsis Zhu L. Yang et al. and Myxoderma Fayod ex
Kühner (Cui et al. 2018). In this family, Amanita is the
most species-rich genus (Bas 2000; Tulloss 2005; Yang
2005; Cui et al. 2018). More than 1000 species have been
described worldwide with ca. 600 accepted names; about
60 taxa have been reported from India (Yang 2000; Bhatt
et al. 2003; Tulloss and Yang 2016).
Amanita Pers
Amanita was recently discussed by Ariyawansa et al.
(2015a), Tulloss and Yang (2016), Tulloss et al. (2016) and
Cui et al. (2018). We follow the latest treatment and
updated accounts of Amanita in Tulloss and Yang (2016),
Tulloss et al. (2016), Tibpromma et al. (2017) and Cui et al.
(2018). Two novel species (belonging to Amanita
sect. Amanita and A. sect. Vaginatae), collected from the
north-western part of Indian Himalaya are introduced
together with their morphology and phylogenetic placements (Figs. 128, 131). In addition, A. altipes and A.
Fig. 122 Maximum likelihood
phylogenetic tree of Coprinus
trigonosporus and closely
related species based on ITS
sequence dataset and calculated
with MEGA6.0 software
(Tamura et al. 2013). The new
species is shown in blue.
Maximum likelihood bootstrap
values greater than 50% are
indicated at the nodes. The tree
is rooted with Tulostoma
kotlabae and T. niveum. The bar
indicates the number of
nucleotide substitutions per site
123
melleialba are reported from Thailand for the first time,
which was established based on morphology and DNA
sequence analyses of LSU and RPB2 regions (Fig. 124).
Amanita altipes Zhu L. Yang, M. Weiss & Oberw.,
Mycologia 96(3): 636 (2004)
Facesoffungi number: FoF04857, Fig. 125a
Holotype: China, Yunnan Province, Lijiang Country,
Laojunshan, altitude 3800 m., on soil under Abies, Betula,
Picea, Quercus and/or Salix, 14 August 2000, Zhu L. Yang
2915, KUN-HKAS 36609.
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes medium-sized. Pileus 70 mm in wide,
parabolic when young, convex to plano-convex, depressed
at centre, broadly umbonate, yellowish to yellow (3A4–7),
pale yellow toward margin, often with dark yellow to
brownish (5D6–8) over disk, viscid when moist; universal
veil mostly over disc as felty, floccose patches, 2–5 mm
wide, thick, yellowish to yellow to dirty yellow (3A2–5);
margin tuberculate-striate (0.13–0.41R), non-appendiculate. Lamellae 2–4 mm broad, free to nearly free, crowded
to plentiful, white to yellowish (2A1–3); lamellulae of 2–5
lengths, crowed, truncate; context white, hollow in centre.
Stipe 150 9 16 mm. (length includes bulb), with tapering
upward, yellowish (3A2–4), becoming whitish toward stipe
base, often covered with yellow to yellowish (3A4–6)
squamules above partial veil, white, with yellowish to
whitish squamules or fibrils under partial veil, white (1A1).
Bulb subglobose to ovate, 8–32 mm diam., white.
Universal veil on stipe base as very short volval limb on
bulb margin, floccose patches or warts near apex of bulb,
yellow to yellowish (3A6–8). Partial veil subapical,
Fungal Diversity
Fig. 123 Coprinus trigonosporus (CNF 1/6594, holotype). a Basidiomes. b Veil on the pileus. c–p Basidiospores. q, r Basidia surrounded by
hymenophysalides. s Cheilocystidia. Scale bars a = 10 mm, b = 30 lm, c–p = 5 lm, q = 20 lm, r, s = 10 lm
membranous, 18 mm below apex of stipe, surface yellowish, with a yellow edge. Odour indistinct. Lamellar
trama bilateral, divergent; mediostratum 50–60 lm wide;
filamentous hyphae 1.8–8 lm wide, branching, hyaline, of
abundant fusiform to subellipsoid, inflated cells
(60–90 9 12–33 mm); vascular hyphae not observed.
Subhymenium 30–50 lm thick; inflated cells dominating,
in 2–3 layers, ovoid to subellipsoid, 10–25 9 8–18 lm,
subtended by concatenated partially inflated hyphal
segments. Basidia 32–70 9 10–14 lm, narrowly clavate to
clavate, mostly 4-spored, occasionally 2-spored, with
sterigmata up to 7 lm long; clamps absent. Basidiospores
[60/1/1]
(7.5–)8–10(–11) 9 (7–)7.5–9.5(–10)
lm,
(Q = 1–1.1(–1.2); Q’= 1.07 ± 0.06), smooth, hyaline,
colourless, thin-walled, inamyloid, globose to subglobose,
rarely broadly ellipsoid, apiculus rather variable, sublateral,
very prominent to rather small, cylindric to truncate-conic;
contents monoguttulate or occasionally granular; white in
123
Fungal Diversity
deposit. Lamellar edge sterile; filamentous hyphae 3–7 lm
wide, hyaline, colourless or pale yellow, thin-walled;
inflated cells dominating, mostly globose to subglobose
and sometimes ovoid, 15–25 9 10–18(–24) lm, colourless, thin-walled. Pileipellis up to 155 lm thick; upper
layer (70–130 lm thick) strongly gelatinized, composed of
interwoven, thin-walled, colourless, filamentous hyphae
3–8 lm wide; lower layer (50–65 lm thick) composed of
compactly arranged, filamentous hyphae 3–8 lm wide;
vascular hyphae rare. Universal veil on pileus filamentous
hyphae 2.5–6.2 lm wide, branching, with slightly inflated
elements; inflated cells very abundant to nearly dominant,
globose to subglobose to ovoid or ellipsoid to subfusiform
(45–90 9 15–30 lm), in chains of 2–4, thin-walled,
colourless vascular hyphae occasional. Universal veil on
stipe base composed 5–6 irregularly arranged elements;
filamentous hyphae very abundant 2.2–15 lm wide; inflated cells abundant to very abundant, fusiform to long
ellipsoid, 40–80 9 10–35 lm, colourless or with intracellular pale brown pigment, thin- to slightly thick-walled,
terminal or in chains of 2–3 and then terminal, becoming
rare toward inner layer; filamentous hyphae abundant;
vascular hyphae occasional. Stipe trama longitudinally
acrophysalidic; filamentous hyphae 2.2–14 lm wide;
acrophysalides
subfusiform
to
clavate,
250–450 9 25–40 lm; vascular hyphae rare. Partial veil
filamentous hyphae 2–5 lm wide, gelatinized, branching,
hyaline, inflated cells terminal, thin-walled, ellipsoid to
long ellipsoid, 15–70 9 10–24 lm, colourless, or with
intracellular pale brown pigment, thin-walled; vascular
hyphae rare. Clamp connections absent in all parts of
basidiomes.
Material examined: THAILAND, Lampang Province,
Along road number 1252, 18.935, 99.390833, elev.1450 m,
14 June 2013, B. Thongbai, BZ201342 (MFLU 14-0065).
Host and habitat: Solitary in forests dominated by
Fagaceae.
Distribution: Known from China and Thailand.
GenBank
numbers:
LSU = MH716040,
RPB2 = MH727686.
Notes: Amanita altipes, originally described from southwestern China by Yang et al. (2004), resembles several
species in the section Amanita, such as A. elata (Massee)
Corner & Bas, A. gemmata (Fr.) Bertillon, A. orientigemmata Zhu L. Yang & Yoshim. Doi, A. russuloides (Peck)
Sacc., and A. xylinivolva Tulloss et al. Amanita altipes
shares some morphological similarities with these yellow
to yellowish species which have been discovered in either
Europe or Asia. Molecular analyses indicate that A. altipes
is a distinct species closely related to A. melleialba. The
combined gene phylogenetic analyses indicate that Thai A.
altipes sequences clustered with A. altipes sequences from
China with 100% ML support (Fig. 124).
123
Amanita flavoalba Mehmood & R.P. Bhatt, sp. nov.
MycoBank number: MB820829; Facesoffungi number:
FoF04390, Figs. 126, 127
Etymology: Referring to the yellow centre and whitish
margin of the pileus.
Holotype: CAL 1405.
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes small to medium-sized. Pileus 40–60 mm
wide, initially hemispherical then convex to plane, slightly
umbonate, yellow to vivid yellow or cadmium yellow
(2A7–8) over centre, white to yellowish white (1A2–2A2)
toward margin, viscid to sub-viscid when moist, shiny;
context 3–4 mm thick, thinning slowly toward margin,
white, unchanging when bruised or exposed; margin striate
(0.1–0.2R), non-appendiculate. Universal veil on pileus as
membranous to sub-membranous patches, 2–4 mm wide,
white or snow white (1A1), distributed irregularly over the
pileus surface. Lamellae free to narrowly adnate, leaving
decurrent line on stipe, crowded (8–10 lamellae/10 mm at
margin), white, unchanging, 3–8 mm broad. Lamellulae
truncate to subtruncate, plentiful, in several lengths. Stipe
70–95 9 5–7 mm (length includes bulb), nearly cylindric or
slightly tapering upward, slightly flaring at apex, white to
yellowish white (1A2), finely fibrillose above and below
annulus; context white, unchanging when cut or bruised,
stuffed. Partial veil medium to subapical, membranous,
white. Bulb 17–19 9 15–17 mm, subglobose to ovoid,
remnants of universal veil on top of bulb white, sub-membranous to shaggy or as cottony patches. Odour indistinct.
Taste not recorded. Spore print white. Macrochemical test no
reaction to 5% KOH on pileal surface and context. Subhymenium wst-near = 25–40 lm; wst-far = 38–55 lm thick,
with basidia arising from short inflated hyphal segments up
to 8 9 10 lm. Hymenophoral trama bilateral, divergent;
wcs = 30–45 lm, composed of long ellipsoid to subfusiform
inflated cells (up to 65 9 30 lm), filamentous hyphae
3–7 lm wide, thin-walled, colourless, hyaline. Basidia (42–
)45–55(–60) 9 (10–)10.5–11.5(–12) lm, thin-walled,
colourless; sterigmata 3–4 lm long; clamp connections not
observed at the base of basidia. Basidiospores [60/3/2](8–
)8.5–10.5(–11) 9 (7–)7.5–9(–9.5)
lm,
(L = 9.5 lm;
W = 8.3 lm; Q = (1.09–)1.11–1.17(–1.21); Q’ = 1.14),
subglobose, sometimes broadly ellipsoid, thin-walled, hyaline, smooth, non-amyloid, apiculus sublateral, up to 1.5 lm
high; contents monoguttulate. Lamellae edge sterile, with
inflated, clavate or subglobose to pyriform cells,
14–35 9 8–13 lm, dominating, thin-walled, colourless.
Pileipellis 110–130 lm thick, in two layers; upper layer
40–50 lm thick, slightly gelatinized, colourless, filamentous
hyphae 2–4 lm wide, subradially arranged to interwoven;
lower layer 70–80 thick, filamentous hyphae 2–5 lm wide,
subradially to densely arranged with yellowish brown
Fungal Diversity
Fig. 124 Phylogenetic tree of
Amanita species inferred from a
combine LSU and RPB2
sequence dataset using
maximum likelihood. Bootstrap
values (BS) C 50% are shown
above the branches. The first
records of Amanita species from
Thailand in this study are in
blue font. Voucher collection
identifiers are provided after
each species name. Type
specimens are indicated in bold
intracellular pigment, thin-walled. Universal veil on pileus
filamentous hyphae 3–7 lm wide, dominating, hyaline;
inflated cells globose to subglobose 20–50 9 26–47 lm,
broadly ellipsoid to ellipsoid or fusiform, 37–64 9
12.5–20 lm, thin-walled, colourless, hyaline. Pileus context
filamentous hyphae 5–8 lm wide, colourless, hyaline, thinwalled, inflated cells 30–90 9 10–30 lm, colourless, thinwalled; vascular hyphae not observed. Universal veil on stipe
base filamentous; hyphae 4–9 lm wide, dominating,
branching, hyaline, thin-walled; inflated cells subglobose to
ovoid 18–36 9 21–37 lm, broadly ellipsoid to subclavate
39–74 9 12.5–20 lm, abundant to very abundant. Partial
veil filamentous hyphae 2–4 lm wide, branching, thin-walled, colourless, hyaline, with inflated cells up to
25 9 13 lm. Stipe context longitudinally acrophysalidic;
acrophysalides dominating 190–280 9 26–36 lm, filamentous hyphae (3–7 lm wide). Clamp connections absent
in all tissues.
Material examined: INDIA, Uttarakhand, Bageshwar
District, Dhakuri, 2570 m, N30°04.9620 E79°55.1590 , 2
August 2016, T. Mehmood, TM 16-1249 (CAL 1405,
123
Fungal Diversity
Fig. 125 Basidiomes.
a Amanita altipes. b Amanita
melleialba. Scale bars 10 cm
holotype); ibid., Dhakuri, 3 August 2016, T. Mehmood,
TM 16-1280.
Host and habitat: On ground under Abies pindrow
(Royle ex D. Don) Royle in temperate coniferous forest
Distribution: Indian Himalaya.
GenBank numbers: LSU = KY861748 (CAL 1405),
MF695813 (TM 16-1280).
Notes: A combination of macro- and micromorphological features like inamyloid basidiospores and bulbous stipe
base place A. flavoalba in Amanita [subg. Amanita]
sect. Amanita (Yang 1997).
Amanita flavoalba is distinct from other known species
of Amanita sect. Amanita by a combination of characteristics: smaller basidiomes, yellow to vivid yellow pileus
centre and whitish to yellowish white toward pileus margin, membranous to submembranous universal veil on
pileus, subglobose basidiospores, universal veil on the
pileus composed of abundant filamentous hyphae mixed
with scattered inflated cells. In the field, A. flavoalba is
easily recognized by its yellow pileus with yellowish white
margin, easily detachable universal veil on pileus in the
form of membranous to submembranous patches and its
occurrence under Abies pindrow. The presence of subglobose basidiospores, very abundant filamentous hyphae
mixed with scattered inflated cells in universal veil on the
pileus are also quite striking.
Amanita flavoalba might be confused with A. melleialba
Zhu L. Yang et al. (originally described from China),
however, the latter possesses a honey pileus with longer
marginal striation, subconical to granular universal veil
remnants on pileus, ellipsoid to sometimes broadly ellipsoid basidiospores (7.5–9.5 9 6–7 lm, with Q = 1.41) and
123
it occurs in subtropical forests dominated by plants of
Fagaceae (Ariyawansa et al. 2015a).
Amanita flavoalba is similar to A. pseudosychnopyramis
Yang Y. Cui et al., but the latter has yellowish brown to
brownish pileus, universal veil remnants as conical to
pyramid, grey to brownish grey warts and it occurs in
subtropical forests of Fagaceae (Ariyawansa et al. 2015a).
Amanita parvipantherina Zhu L. Yang et al. and A.
subparvipantherina Zhu L. Yang et al. are easily segregated from A. flavoalba by the universal veil remnants on
pileus in the form of subconical to granular warts (Yang
et al. 2004; Ariyawansa et al. 2015a). Amanita parvipantherina also has broadly ellipsoid to ellipsoid basidiospores
(8.5–11.5 9 6.5–8.5 lm, Q = 1.38; Yang et al. 2004) and
A. subparvipantherina has broadly ellipsoid to ellipsoid
basidiospores (9–11.5 9 6.5–8 lm, with Q = 1.38; Ariyawansa et al. 2015a). Amanita flavoalba resembles A. altipes
by its yellowish pileus but A. altipes pileus has a brownish
tinge and its basidiospores are globose to subglobose
(8–10 9 7.5–9.5 lm, with Q = 1.07; Yang et al. 2004).
Amanita elata (originally described from Singapore) differs
from A. flavoalba by its pale dingy to ochraceous buff
pileus with very faint sulphur-yellow tinge, exannulate
stipe, smaller basidiospores (7–8.5 9 6.8–7.7 lm) and its
occurrence in tropical forests (Corner and Bas 1962).
Our molecular phylogenetic analysis shows closeness
(though well separated in Fig. 128) with a few taxa (in
Amanita sect. Amanita) like A. xylinivolva (originally
described from Colombia; GenBank no. FJ890036), A.
crenulata Peck (originally described from the USA; GenBank no. HQ539687) and A. breckonii Thiers & Ammirati
(originally reported from the USA; GenBank no.
KJ535440).
Fungal Diversity
Fig. 126 Amanita flavoalba (CAL 1405, holotype). a–e Fresh basidiomes in the field and base camp. f Basidiospores. g, h Basidia and element
of subhymenium. i Elements of universal veil from pileus surface. Scale bars a = 10 mm, f–i = 20 lm
Amanita xylinivolva can be easily separated by its
basidiomes without a pileal umbo, white to off-white to
dingy grey volval remnants, whitish buff lamellae, marked
to abrupt bulb and microscopically, globose to subglobose
basidiospores with comparatively lower Q value
(8–10.2 9 7.2–9.5 lm, Q0 = 1.08; Tulloss et al. 1992).
Amanita crenulata has a brownish beige to greyish pileus,
universal veil on pileus as flocculose to subpyramidal warts
and subglobose to broadly ellipsoid basidiospores (Q0 =
1.08) (Peck 1900). Amanita breckonii has ellipsoid to
elongate basidiospores (9.8–12.8 9 6.2–8.7 lm, Q0 =
1.65) and clamp connections at the base of basidia (Thiers
and Ammirati 1982). Macro- and micromorphology coupled with the LSU-based phylogenetic analysis corroborate
A. flavoalba as a novel species.
Amanita melleialba Zhu L. Yang, Qing Cai & Yang Y.
Cui, in Ariyawansa et al., Fungal Divers: https://doi.org/10.
1007/s13225-015-0346-5, [163] (2015), Fig. 125b
123
Fungal Diversity
Fig. 127 Amanita flavoalba
(CAL 1405, holotype).
a Basidiomes. b Basidia and
element of subhymenium.
c Elements of universal veil
from pileus surface.
d Basidiospores. Scale bars
a = 10 mm, b–d = 10 lm
Holotype: CHINA, Yunnan Province, Puer City,
Caiyanghe Nature Reserve, Fagaceae (elev. 1300 m), 11
July 2014, G. Wu 1339 (HKAS 83446).
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes small-sized. Pileus 30–48 mm wide, parabolic when young, planoconvex to plane at maturity,
depressed at centre, yellowish (4A3–4) to yellow (3B5–6),
becoming yellowish to whitish (3A2) toward margin, slightly
viscid when moist; universal veil mostly over disc, scarce
towards margin, as granular or small warts dirty white (5B2),
cream to yellowish (4A2–3), honey; margin tuberculate-
123
striate (0.4–0.6R), non-appendiculate. Lamellae 2–4 mm
broad, free to nearly free, crowded, white (1A1); lamellulae
of 3–4 lengths, truncate. Stipe 40–75 9 4–7 mm (length
includes bulb), slightly tapering upward, white (1A1) to dull
white ground (1A2), often covered with floccose squamules
above partial veil, white (1A1), floccose squamules to granules under partial veil, white (1A1); context stuffed to nearly
hollow in centre, thin, white (1A1). Bulb subglobose to
napiform, 6–12 mm wide, white (1A1). Universal veil on
stipe base as very short volval limb on bulb margin and
floccose squamules to granules dirty white (1A2). Partial veil
subapical, membranous, 15–30 mm below apex of stipe,
Fungal Diversity
white (1A1) to cream (1A2), with a yellow floccose edge.
Odour indistinct. Lamellar trama bilateral, divergent;
mediostratum 30–60 lm wide; filamentous hyphae
1.8–8 lm wide, branching, hyaline, with slightly inflated
elements; vascular hyphae not observed. Subhymenium
30–60 lm thick; inflated cells dominating, in 3–4 layers,
subglobose to ovoid, 10–19 9 8–17 lm, subtended by
concatenated partially inflated hyphal segments. Basidia
32–50 9 10–14 lm, narrowly clavate to clavate, mostly
4-spored, occasionally 2-spored, with sterigmata up to 5 lm
long; clamps absent. Basidiospores [60/1/2] (7.2–)7.3–
9.4(–10) 9 (5.3–)6.5–7(–7.3) lm, Q = 1.2–1.28(–1. 56);
Q’ = 1.4 ± 0.06, smooth, hyaline, colourless, thin-walled,
inamyloid, ellipsoid, sometimes broadly ellipsoid; apiculus
rather variable, sublateral, very prominent to rather small,
cylindric to truncate-conic; contents monoguttulate or
occasionally granular; white in deposit. Lamellar edge sterile; filamentous hyphae 3–7 lm wide, hyaline, colourless or
pale yellow, thin-walled; inflated cells dominating, mostly
globose to subglobose and sometimes ovoid, 9–30 9 9–18(–
24) lm, colourless, thin-walled. Pileipellis up to 155 lm
thick; upper layer (50–70 lm thick) strongly gelatinized,
composed of interwoven, thin-walled, colourless, filamentous hyphae 3–8 lm wide; lower layer (50–85 lm thick)
composed of compactly arranged, filamentous hyphae
3–8 lm wide; vascular hyphae rare. Universal veil on pileus
filamentous hyphae, 2.5–6.2 lm wide, branching, with
slightly inflated elements; inflated cells very abundant to
nearly dominant, globose to subglobose to ovoid or ellipsoid
to subfusiform (20–60 9 10–30 lm), in chains of 2–4, thinwalled, colourless; vascular hyphae occasional. Universal
veil on stipe base composed of two layers; inflated cells
abundant to very abundant, fusiform to ellipsoid,
20–40 9 10–30 lm, colourless or with intracellular pale
brown pigment, thin- to slightly thick-walled, terminal or in
chains of 2–3 and then terminal, becoming rare toward inner
layer; filamentous hyphae abundant; vascular hyphae occasional. Stipe trama longitudinally acrophysalidic; filamentous hyphae 2.2–15 lm wide; acrophysalides subfusiform to
clavate, 120–350 9 25–45 lm; vascular hyphae rare. Partial veil filamentous hyphae 2–5 lm wide, gelatinized,
branching, hyaline, inflated cells terminal, thin-walled,
ellipsoid to long ellipsoid, 15–65 9 10–28 lm, colourless,
thin-walled; vascular hyphae rare. Clamp connections absent
in all parts of basidiomes.
Material examined: THAILAND, Chiang Rai Province,
Chiang-khong District, Huay-sor Subdistrict, Nensomburn
Village, 5 August 2015, B. Thongbai, BZ201518 (MFLU
15-3316).
Habit and habitat: Gregarious in forests dominated by
Fagaceae.
Distribution: Known from China and now Thailand.
GenBank numbers: LSU = MH716041, RPB2 =
MH727687.
Notes: Amanita melleialba belongs to subg. Amanita
sect. Amanita. It was originally described from southwestern and central China. In the field, the distinguishing
morphological characteristics of A. melleialba include its
small basidiomes, universal veil mostly over disc as
granular or small warts as well as stipe base. Amanita
melleialba is very similar to A. parvipantherina, also
known from China. However, A. parvipantherina has larger basidiospores (8.5–11.5 9 6.5–8.5 lm) and is distributed in mixed forests with Pinus yunnanensis (Yang
et al. 2004; Yang 2005, 2015, Ariyawansa et al. 2015a).
Combined-gene phylogenetic analyses show that Thai A.
melleialba cluster with A. melleialba strains from China
with 100% ML support (Fig. 124).
Amanita subtropicana Mehmood & R.P. Bhatt, sp. nov.
MycoBank number: MB824531; Facesoffungi number:
FoF04391, Figs. 129, 130
Etymology: Referring to the subtropical region of
Uttarakhand, the type locality.
Holotype: CAL 1660
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes small to medium-sized. Pileus 30–65 mm
wide, olive brown (5F4–5) yellowish brown (5F6–8) to
brown (6E4) over centre, honey yellow to khaki (4D4–6)
toward margin, initially hemispherical, then convex to
plano-convex, slightly umbo, dry, shiny; context 8–12 mm
thick, white, unchanging when cut or bruised; margin
tuberculate-striate, up to 14 mm wide, non-appendiculate.
Universal veil remnants on pileus as felted, subconical to
subpyramidal warts. Lamellae free, crowded, (8–12 per
10 mm at margin), 5 mm broad, initially white (1A1),
fading yellowish white (1B2) with age. Lamellulae truncate, unevenly distributed. Stipe 90–160 9 10–20 mm,
nearly cylindrical, slightly tapering upward, with apex
slightly expended, initially white (5A1), covered with
white farinose squamules; context white, hollow,
unchanging. Partial veil absent. Universal veil remnants on
stipe base white (1B1) to yellowish white (1C1), felted
warts, mostly in complete ring, sometimes irregularly distributed. Odour indistinct. Taste not recorded. Spore print
white. Subhymenium wst-near = 25–40 lm; wst-far =
45–60 lm; basidia arising from subglobose to irregular
shaped cells, 6–16 9 4–14 lm. Hymenophoral trama
bilateral, divergent; wcs = 36–55 lm wide, with inflated
(clavate to fusiform) cells, 40–60 9 10–14 lm; filamentous, undifferentiated hyphae 3–7 lm wide, thin-walled,
colourless, hyaline. Basidia (32–)46–60(–68) 9 (12–
)13–15(–16) lm, 2- to 4-spored, thin-walled, colourless;
sterigmata 3–5 lm long; clamp connections not observed
123
Fungal Diversity
Fig. 128 Maximum likelihood phylogram of Amanita showing the
position of Amanita flavoalba. Phylogenetic analysis was conducted
in MEGA 6.0 (Tamura et al. 2013) based on LSU sequence data.
Bootstrap support values ([ 50%) obtained from maximum likelihood analysis are shown above or below the branches at nodes. New
taxon is highlighted in blue. Type specimens are indicated in bold.
Amanita morrisii Peck, A. brunneolocularis Tulloss & Franco-Mol.
and A. orsonii Ash. kumar & T.N Lakh. were used as the outgroup
taxa
at the base of basidia. Basidiospores [80/4/2] (7–)8.5–11(–
12) 9 (5.5–)6–8(–8.5) lm, (L = 10.5 lm; W = 7.5 lm;
Q = (1.19–)1.33–1.47(–1.6); Q’ = 1.41), broadly ellipsoid
to ellipsoid, non-amyloid, hyaline, thin-walled, smooth,
apiculus sublateral, up to 1.7 lm high; contents monoguttulate. Lamellar edge cells sterile with inflated cells, subglobose to pyriform, 20–48 9 18–36 lm, thin-walled,
colourless, hyaline or sometimes with yellowish brown
pigments. Pileipellis 80–140 lm thick, in two layers;
gelatinized suprapellis, 25–50 lm thick, composed of
compactly arranged filamentous, undifferentiated hyphae
2–5 lm wide, thin-walled, colourless, hyaline; ungelatinized subpellis, 55–90 lm thick, composed of filamentous, undifferentiated hyphae 3–7 lm wide, radially
arranged with yellowish brown intracellular pigments.
Pileus trama filamentous, undifferentiated hyphae 3–8 lm
123
Fungal Diversity
wide; with broadly ellipsoid to ellipsoid cells,
30–65 9 10–20 lm. Universal veil on pileus with inflated
cells globose to subglobose, 20–44 9 18–38 lm, broadly
ellipsoid to ellipsoid, 16–62 9 12–38 lm; filamentous,
undifferentiated hyphae 3–6 lm wide. Universal veil on
stipe base is similar to that of pileus surface. Stipe context
longitudinally
acrophysalidic;
acrophysalides
118–224 9 24–32 lm;
filamentous,
undifferentiated
hyphae 2–10 lm wide, thin-walled hyaline. Clamp connections absent in all tissues.
Material examined: INDIA, Tehri Garwhal, Byasi,
510 m, N30°04.000 E78°28.090 , 9 August 2015, T. Mehmood, TM 15-915 (CAL 1660, holotype); ibid., Byasi, 18
August 2017, T. Mehmood, TM 17-1574.
Habit and habitat: Solitary to scattered, on ground under
Shorea robusta C. F. Gaertn. (Dipterocarpaceae) in subtropical mixed forest.
Distribution: Indian Himalaya.
GenBank numbers: LSU = MG913204 (TM 17-1574);
MG923799 (TM 15-915).
Notes: In the field, Amanita subtropicana is distinct
from other known species of Amanita sect. Vaginatae by a
combination of macroscopic characteristics: olive brown to
yellowish brown pileus over centre, honey yellow to khaki
colour towards margin which is covered by felted, subconical, subpyramidal to warty universal veil remnants on
the pileal surface and broadly ellipsoid to ellipsoid basidiospores and putative association with Shorea robusta in
subtropical broadleaf forest. Amanita ceciliae (Berk. &
Broome) Bas, A. liquii Zhu L. Yang et al., A. griseofolia
Zhu L. Yang, and A. cinctipes Corner & Bas are species of
sect. Vaginatae which are somewhat similar to A. subtropicana on the basis of friable universal veil at the stipe
base.
Amanita ceciliae, a European species, can be easily
segregated from A. subtropicana by its yellow brown to
grey-brown or olive-brown pileus with greyish to brownish
universal veil remnants (Phillips 1990; Breitenbach and
Kränzlin 1995). Amanita liquii, originally described from
southwestern China, differs from A. subtropicana by its
dark brown to blackish pileus with dark universal veil
remnant and globose to subglobose basidiospores. Furthermore, it is associated with Abies and Picea (Yang et al.
2004). Amanita griseofolia differs by its grey to brownish
grey pileus and felted to verrucose universal veil and globose to subglobose basidiospores (10–13.5 9 9.5–13 lm;
Yang 2004). Amanita cinctipes (originally described from
Singapore) is distinct from A. subtropicana by its mouse
grey to greyish brown pileus with abundant universal veil
remnants on the base of the stipe forming 2–4 rings and
globose to subglobose basidiospores (Corner and Bas
1962).
Three Indian species, Amanita cornelii Mehmood et al.,
A. emodotrygon Mehmood et al. and A. rajendrae Mehmood et al. (GenBank no. KX528072, KX539266 and
MF170174, respectively, in Fig. 131) of sect. Vaginatae
are easily segregated from A. subtropicana by their distinct
saccate universal veil on stipe base (Das et al. 2017b;
Tibpromma et al. 2017).
Initial BLASTn search results of LSU sequence from
Indian collection (TM 17-1574) against the NCBI database
exhibited 87% similarity with 100% query coverage of
Amanita subtropicana to ‘‘Amanita sp. Aus09’’ (GenBank
no. KY349232). Phylogenetically (Fig. 131), A. subtropicana might have some closeness with Amanita sp. (Aus09),
A. madagascariensis L.P. Tang et al. and A. strobilaceovolvata (Beeli) E.-J. Gilbert. Unfortunately, no morphological description of Amanita sp. (Aus09) is available for
comparison of important morphological characteristics.
Amanita madagascariensis originally described from East
Africa can easily be separated by its dirty white or dull
white pileus covered with greyish to brownish grey universal veil remnants and occurrence under Eucalyptus
(Tang et al. 2015). Amanita strobilaceovolvata, originally
described from the Congo, has a yellow to pale yellow
pileus, a strobiloid saccate universal veil and subglobose to
broadly ellipsoid basidiospores (8–10 9 7–9 lm) (Gilbert
1940; Tang et al. 2015).
Hygrophoraceae Lotsy
The family Hygrophoraceae contains 25 genera and
over 600 species (Lodge et al. 2014). Most species in the
family prefer a humicolous habitat, except for a few that
are lignicolous, ectomycorrhizal, or on mosses (Griffith
et al. 2002).
Hygrocybe (Fr.) P. Kumm
Hygrocybe is cosmopolitan in distribution (Senthilarasu
et al. 2010). Currently, 457 species are listed as legitimate
in Index Fungorum and MycoBank (2019). The genus is
characterized by the presence of variously coloured
basidiomes with waxy lamellae, absence of veilar remnants
on the pileus margin as well as stipe, and white, inamyloid,
smooth basidiospores (Boertmann 1995; Babos et al. 2011;
Lodge et al. 2014; Hosen et al. 2016). Most species predominantly grow on soil, except for a few found on tree
trunks or on logs (Lodge et al. 2006).
Hygrocybe lucida K. Acharya & A.K. Dutta, sp. nov.
MycoBank number: MB826973; Facesoffungi number:
FoF05720, Figs. 132, 133
Etymology: Refers to the bright colouration of the
pileus.
Holotype: CUH AM123
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
123
Fungal Diversity
Fig. 129 Amanita subtropicana (CAL 1660, holotype). a, b Fresh basidiomes in the field. c Universal veil remnants on pileus surface. d, e
Basidiospores. f Basidia and element of subhymenium. g Elements of universal veil on pileus surface. Scale bars a, b = 10 mm; c–g = 20 lm
Basidiomes small to medium-sized. Pileus 6–23 mm
diam., convex to broadly convex when young, becoming
somewhat infundibuliform at maturity; surface somewhat
sticky when moist, greyish orange (5B5–6) to orange
(6A6–7) or reddish orange (7A7) with a slightly darker (7A8,
7B8) disc when young, becoming light orange (6A5) to
orange (6A6–7) with reddish orange (7A7, 7B7) to orangered (8A7, 8B7–8) centre, no colour change on bruising or
with age, smooth and glabrous, often slightly fibrillose;
margin at first entire, turning wavy at maturity. Lamellae ca.
2–3 mm broad, adnexed to broadly adnate, thick, white
(1A1), regular, often forked towards margin from half-way to
the stipe, distant, with lamellulae of 1–2 lengths; edge concolorous with the sides, smooth. Stipe 4–11 9 1.5–3 mm,
central, orange (5A6–7) to deep orange (5A8) or greyish
123
orange (5B6), cylindrical, rarely slightly broader towards the
apex, mostly glabrous, semi moist, often slightly fibrillose,
hollow. Odour and taste indistinct. Lamellar trama regular,
made up of 3.5–7.5 lm broad, cylindrical, IKI-, thin-walled,
hyaline, hyphae, individual compartments 29–76 lm long,
tubuliform elements absent. Basidia dimorphous; macrobasidia 50–53(–58) 9 5–7(–8.5) lm, narrowly clavate to
cylindric-clavate, thin-walled, with refractive oleaginous
contents, 4-spored; microbasidia 32–35(–40) 9 4–5(–7)
lm, narrowly clavate, thin-walled, with refractive contents,
4-spored. Basidioles 32–39(–58) 9 5–6(–7) lm, narrowly
clavate, thin-walled, hyaline, with refractive contents. Basidiospores vaguely dimorphous; macrospores 9–10–10.3(–
10.8) 9 (6–)6.5–7–7.2 lm, Q = 1.4–1.45–1.5, ellipsoid,
hyaline, thin-walled, inamyloid, with refractive oleaginous
Fungal Diversity
Fig. 130 Amanita subtropicana
(CAL 1660, holotype).
a Basidia and element of
subhymenium. b Elements of
universal veil from pileus
surface. c Basidiospores. Scale
bars a = 10 mm, b–d = 10 lm
contents when viewed in KOH, not constricted at the middle;
microspores
(6.4–)7–7.2–7.5(–7.9) 9 (3.9–)5–5.5–6.5(–
6.8) lm, Q = 1.1–1.3–1.8 lm, subglobose to ellipsoid,
hyaline, thin-walled, IKI-, with few refractive oleaginous
contents in KOH, not constricted at the middle. Lamellar
edge fertile. Cheilo- and pleurocystidia absent. Pileipellis an
ixotrichodermium, made up of erect, mostly unbranched to
often branched, cylindrical hyphae, (5–)7–10(–15) lm
broad, hyaline, IKI-, thin-walled, hyphal end often broader.
Pileus tramal hyphae 7–9 lm broad, hyaline, IKI-, thinwalled. Stipitipellis an ixocutis with thin-walled hyphae,
3.5–5.5 lm wide, hyaline. Stipe trama hyphae 7–8 lm
broad, interwoven, hyaline, thin-walled, IKI-. Caulocystidia
absent. Clamp connections present in all parts of the basidiome including the base of basidia.
Material examined: INDIA, West Bengal, North-24Parganas, near Basirhat, 22°390 10.000 N, 88°520 00.300 E (alt.
9 m), 9 August 2015, K. Acharya & A.K. Dutta, CUH
AM123 (holotype); ibid., North-24-Parganas, Basirhat,
22°380 36.100 N, 88°530 35.300 E (alt. 9 m), 12 August 2015, K.
Acharya & A.K. Dutta, CUH AM126.
Habit and habitat: Uncommon, solitary or scattered, on
humus-rich soil.
Distribution: India.
GenBank numbers: ITS = MH599084 (CUH AM123);
ITS = MH599083 (CUH AM126).
Notes: The distinguishing features of Hygrocybe lucida
include small basidiome size, a convex (when young) to
somewhat infundibuliform (at maturity), orange to reddish
orange pileus that does not change colour on bruising or
with age; white, adnexed to broadly adnate lamellae; an
orange or greyish orange stipe; dimorphous basidiospores
and basidia; absence of cheilo- and pleurocystidia; an
ixotrichodermium-type pileipellis; regular lamellae trama
composed of short hyphal elements (29–76 lm long); an
ixocutis-type stipitipellis; absence of caulocystidia; and
presence of clamp connections in all parts of the basidiome. This combination of features place the present species in sect. Firmae of subgen. Pseudohygrocybe (Pegler
and Fiard 1978; Pegler 1986; Cantrell and Lodge 2001;
Lodge and Ovrebo 2008). The morphological placement is
further supported by the molecular phylogenetic analysis
based on ITS sequence data (Fig. 134).
123
Fungal Diversity
Fig. 131 Maximum likelihood phylogram of Amanita showing the
position of Amanita subtropicana. Phylogenetic analysis was conducted in MEGA 6.0 (Tamura et al. 2013) based on LSU sequence
data. Bootstrap support values ([ 50%) obtained from maximum
likelihood analysis are shown above or below the branches at nodes.
The new taxon is highlighted in blue and bold on the tree. Type
specimens are in bold. Amanita caesareoides Lj.N. Vassiljeva and A.
caesarea (Scop.) Pers. were used as the outgroup taxa
Within sect. Firmae, species with more or less similar
size of the macrospores include Hygrocybe alwisii (Berk.
& Broome) Pegler, H. diversicolor (Petch) Pegler, and
H. earlei (Murrill) Pegler. Hygrocybe alwisii, previously
reported from India, and differing by a larger pileus
(45–55 mm), adnexed to almost free lamellae, a longer
stipe (60–70 mm), and a repent epicutis type of pileipellis
(Pegler 1986; Leelavathy et al. 2006). Hygrocybe diversicolor has a larger pileus (up to 50 mm) that is olivaceous
brown to blackish brown with appressed squamulose surface, longer stipe (30–35 mm), purplish grey to purplish
black, and epicutis type of pileipellis (Pegler 1986).
123
Fungal Diversity
Hygrocybe earlei, a species known only from Cuba and
Trinidad, has a pileus that is initially campanulate and
becomes expanded with a broad low umbo, and free to
adnexed lamellae. Hygrocybe cf. earlei, previously known
only from Panama, has an umbonate pileus, pale yellow
lamellae with white edge, a pure white, much longer stipe
(up to 40 mm), smaller macrobasidia (24–46 lm long) and
microbasidia (22–30 lm long), and presence of an intermittent thin gelatinous coating in the pileipellis (Lodge and
Ovrebo 2008).
Hygrocybe batistae Singer, described from Brazil and
later found in Colombia and Puerto Rico, has similarly
coloured basidiomes but differs by its rugulose to rugose
pileus, caespitose basidiomes, and presence of coralloid
hyphae in the pileipellis (Lodge and Pegler 1990; Cantrell
and Lodge 2001; Lodge et al. 2014). Hygrocybe neofirma
S.A. Cantrell & Lodge also has similarly coloured basidiomes but has a star-shaped perforation at the pileus centre,
light yellow to brilliant yellow lamellae, and larger macrospores (12.8–17.6 9 8–10.4 lm; Cantrell and Lodge
2001).
Hygrocybe hypohaemacta (Corner) Pegler, previously
placed within sect. Firmae and recently transferred to
sect. Velosae based on phylogenetic analysis by Lodge
et al. (2014), appears to be close to H. lucida in some
morphological characteristics. However, H. hypohaemacta
has a convex to plano-convex pileus that is covered by a
thick (up to 1.5 mm) greyish gluten, discolorous lamellae
with pale golden edge, longer stipe (up to 60 mm), somewhat differently sized macrospores (7–11 9 5–8 lm),
presence of polymorphic cheilocystidia, and an epicutistype pileipellis (Pegler and Fiard 1978).
Hygrocybe trinitensis (Dennis) Pegler and H. siparia
(Berk.) Singer are comparable in having similar smaller
pilei (Pegler and Fiard 1978). Hygrocybe trinitensis differs
by the presence of coral red lamellae that are broadly
adnate with a decurrent tooth, a longer stipe (up to 50 mm),
smaller macrospores (7–9 9 4.5–5.5 lm, Q = 1.56),
smaller macro- and micro-basidia, and an epicutis-type
pileipellis that contains red vacuolar pigments. Hygrocybe
siparia, described from Brazil, has an umbilicate pileus
with squamulose surface, lamellae with a decurrent tooth,
longer macrospores (10–13 lm long, Q = 1.53), and
smaller macro-basidia (40–48 lm long).
Among phylogenetically related taxa (Fig. 134), Hygrocybe firma (Berk. & Broome) Singer differs by its
yellow pileus, strongly decurrent lamellae, and longer
macrospores (11–18 lm long) (Berkeley and Broome
1871; Young and Mills 2002). Hygrocybe andersonii
Cibula & N.S. Weber, originally described from Horn
Island of Mississippi, has densely cespitose growth habit,
brownish orange to deep orange lamellae, longer stipe (up
to 42 mm long) reddish brown, and large bacilliform
Fig. 132 Hygrocybe lucida (CUH AM123, holotype). Basidiomes in
the field (lamellae facing side, in inset). Scale bars 10 mm. Photos by
A.K. Dutta
basidiospores (14–20.7 9 3.8–5.6 lm; Cibula and Weber
1996).
Marasmiaceae Roze ex Kühner
Marasmiaceae is a family of pale-spored agarics. It has
a worldwide distribution and comprises about 54 genera
and 1590 species (Kirk et al. 2008). Several of these taxa
were previously considered under Tricholomataceae sensu
lato (Moncalvo et al. 2002; Matheny et al. 2006). Most
species are saprobes that play a leading role in nutrientrecycling and form prominent components of forest
ecosystems (Cannon and Kirk 2007). A novel species,
Marasmius indojasminodorus is introduced within Marasmiaceae. An updated phylogenetic tree of ITS sequence
data based on maximum likelihood and Bayesian inference
analyses of the genus Marasmius is provided (Fig. 135).
Detailed literature and updated accounts of Marasmius
were provided by Wannathes et al. (2009) and Tibpromma
et al. (2017).
Marasmius Fr.
We follow the latest treatment and updated accounts of
Marasmius in Komura et al. (2016) and Tibpromma et al.
(2017).
Marasmius indojasminodorus A.K. Dutta, K. Acharya &
K. Das, sp. nov.
MycoBank number: MB820566; Facesoffungi number:
FoF03247, Figs. 136, 137
Etymology: Refers to an Indian look-a-like of the Thai
taxon Marasmius jasminodorus
Holotype: CAL 1514
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Pileus 12–21 mm diam., conical to convex when young,
becoming broadly convex at maturity, often with a small
123
Fungal Diversity
Fig. 133 Hygrocybe lucida
(CUH AM123, holotype).
a Macrospores. b Microspores.
c Macrobasidia. d Microbasidia.
e Basidioles. f Pileipellis
hyphae. Scale bars a–
f = 10 lm. Drawings by A.K.
Dutta
umbonate centre, smooth to minutely pruinose, moist to
semi moist, hygrophanous; disc rugulose, olive-brown
(4D5–8) or light brown (5D7) to yellowish brown (5D–E8,
5F7–8), turning translucent with KOH; margin smooth
when very young, rugulose-striate in age, slightly paler,
yellowish brown (5D4–6, 5E4–5), turning translucent with
KOH. Lamellae 2–2.5 mm broad, adnexed, subdistant
(L = 14–16, l = 2–3), white (1A1) to cream, non-marginate, concolorous, non-intervenose. Stipe 15–40 9 1–
1.5 mm, central, cylindrical, hollow, glabrous to slightly
velutinous, non-insititious, overall white when young, at
123
maturity apex becoming white to cream, base light brown
(6D6) to brown (6D8), strigose at base, yellowish white to
brownish white. Context 1 mm broad at centre, gradually
thinner towards margin, creamy white, turning translucent
with KOH. Odour strong, fragrant, sweet, like jasmine.
Taste slightly bitter. Lamellar trama composed of 6–7(–
7.5) lm broad, interwoven, cylindrical, hyaline, dextrinoid,
non-gelatinous, thin- to thick-walled hyphae. Basidia not
observed. Basidioles 21–22.5(–26) 9 (4–)6–6.5(–7.5) lm,
fusoid to clavate, hyaline, thin-walled. Basidiospores (6.5–
)8–8.7–10(–10.5) 9 3.5–3.7–4 lm,
Q = 1.8–2.4–2.8,
Fungal Diversity
ellipsoid, often curved in profile, hyaline, inamyloid, thinwalled. Pleurocystidia absent. Cheilocystidia composed of
Siccus-type broom cells; main body 10–12(–15) 9 (3.5–
)6–7(–7.5) lm, clavate to subclavate, hyaline, thin- to
moderately thick-walled; apical setulae (3–)6.5–7(–9.5) lm
long, cylindrical, acute to obtuse, yellowish in KOH, thickwalled. Pileipellis hymeniform, consisting of Siccus-type
broom cells; main body (10–)12.5–14(–16.5) 9 (5–)6.5–
7(–8) lm, cylindrical to clavate or broadly clavate, hyaline,
inamyloid, thin- to moderately thick-walled; apical setulae
(3.5–)5–7(–10.5) lm long, cylindrical, obtuse, yellowish
with KOH, thick-walled. Pileus trama hyphae 3.5–5.5 lm
broad, interwoven, cylindrical, often branched, hyaline,
dextrinoid, thin-walled. Stipitipellis hyphae 3.5–4.5 lm
broad, parallel, yellowish, cylindrical, dextrinoid, thinwalled. Stipe trama hyphae 4–7(–9) lm broad, parallel,
hyaline, dextrinoid, cylindrical, non-gelatinous, thin- to
moderately thick-walled. Caulocystidia of two types: a)
abundant non-setulose cells, (17–)28–32(–36) 9 6–7(–11)
lm, cylindrical to irregular in outline, often seldom branched, rarely bilobed at apex, thin- to moderately thickwalled; b) Siccus-type brooms cells with main body 20–
21.5(–25) 9 3.5–4.5(–5) lm, uncommon, scattered,
cylindrical to irregular in outline, hyaline, thin-walled;
apical setulae 10–12.5 lm, cylindrical to irregular in outline, thin- to moderately thick-walled. Clamp connections
present in all the tissues.
Material examined: INDIA, West Bengal, Howrah
District, Acharya Jagadish Chandra Bose Indian Botanic
Garden, 22°330 35.300 N, 88°170 21.500 E (alt. 12.2 m), 23
September 2015, A.K. Dutta & S. Paloi, AKD 135/2015
(CAL 1514, holotype); ibid., 22°330 38.400 N, 88°170 22.600 E
(alt. 15.1 m), 23 September 2015, A.K. Dutta & S. Paloi,
AKD 139/2015 (CAL 1515).
Fig. 134 Phylogram generated from maximum likelihood (RAxML)
analysis using a GTR ? I?G model of nucleotide evolution based on
ITS sequence dataset. Ampulloclitocybe clavipes and Cantharocybe
gruberi were used as the outgroup taxa following Lodge et al. (2014).
Maximum likelihood bootstrap support values greater than 50% are
indicated above or below the nodes. Sequences used in this study
mostly have been sampled from a previous study of Lodge et al.
(2014). The newly generated sequences are placed in blue font to
highlight its phylogenetic position in the tree. GenBank accession
numbers for all of the sequences are indicated in the tree
123
Fungal Diversity
Host and habitat: Uncommon, scattered to gregarious,
on dead and decayed leaves and wood of dicotyledonous
plants.
GenBank
numbers:
ITS = KY785172,
LSU =
KY785174 (CAL 1514); ITS = KY785171, LSU =
KY785173 (CAL 1515).
Notes: The most distinctive combination of features of
Marasmius indojasminodorus includes a small to medium
sized (12–21 mm diam.), convex to broadly convex,
rugulose-striate yellowish brown pileus with a rugulose,
olive brown to light brown or yellowish brown disc;
adnexed, subdistant (14–16), white to cream, lamellae with
lamellulae of 2–3 lengths; a non-insititious, glabrous to
slightly velutinous stipe coloured white to cream at apex
and light brown to brown at base, with yellowish white to
brownish white basal mycelium; strong, sweet, jasmine
like odour; ellipsoid basidiospores with a mean size of
8.7 9 3.7 lm (Q = 1.8–2.8); Siccus-type cheilocystidia
with main body 10–15 9 3.5–7.5 lm; absence of pleurocystidia; and two types of caulocystidia, Siccus-type in
combination with numerous non-setulose cells, cylindrical
to irregular in outline (17–36 9 6–11 lm). This combination of features undoubtedly place the present taxon
within Marasmius ser. Atrorubentes of sect. Sicci (Wannathes et al. 2009).
Among taxa that possess jasmine-like odour, Marasmius
jasminodorous Wannathes et al. (Wannathes et al. 2009),
described from Northern Thailand, differs by its colouration of pileus (dark reddish brown towards disc with light
brown to brownish orange margin), basal mycelium of the
stipe (brownish orange), and larger basidiospores with a
mean range of 10.1 9 3.6 lm (Qm = 2.8). Marasmius
odoratus V.A. Farook & Manim., recently described from
Kerala state of India, primarily differs by the presence of
‘‘Globulares-type’’ cells in the pileipellis and is categorized under sect. Globulares (Farook and Manimohan
2015).
Among other similar species belonging to sect. Sicci,
and phylogenetically close taxa (Fig. 135), M. midnapurensis A.K. Dutta et al., described from India, primarily
differs from M. indojasminodorus by the absence of jasmine-like odour, longer basidiospores (10.7–15 lm long,
Qm = 3), comparatively broader main-body of cheilocystidia (up to 10 lm broad), and longer (up to 50 lm), and
non-setulose caulocystidia (Dutta et al. 2014). The
Indonesian taxon, M. araucariae var. siccipes Desjardin
et al. differs by its non-rugulose pileus, orange brown all
over or with darker disc, discolorous lamellae with orangebrown edge, glabrous stipe, absence of any fragrant odour,
and distinctly longer (11–12 9 3–4 lm), subfusoid basidiospores (Desjardin et al. 2000). Marasmius ochroleucus
Desjardin & E. Horak, described from New Caledonia and
subsequently reported from Northern Thailand, primarily
123
differs by its light yellow to cream pileus, much crowded
(L = 20–24, l = 3–4), and intervenose lamellae, and presence of only one type of caulocystidia shaped cylindrical to
fusoid-ventricose (Wannathes et al. 2009). The Korean
taxon M. occultatiformis Antonı́n et al. has brownish
orange to brownish red pileus and slightly paler towards
margin, presence of more lamellae (L = 25) finely pubescent towards edge, whitish basal mycelium, somewhat
differently sized basidiospores (mean of 7.8 9 4 lm,
Q = 1.95), and absence of caulocystidia (Antonı́n et al.
2012). Marasmius araucariae Singer, described from
Argentina, has castaneous-ferruginous or ferruginous
pileus, more lamellae (16–24), comparatively longer stipe
(up to 70 mm long) with fulvous-white basal mycelium,
absence of any odour, larger basidiospores (9–12.5 9 2.7–
4.5 lm), and only one type of caulocystidia, subcylindric,
50 9 5.5 lm (Singer 1976). Marasmius napoensis Singer
differs by its larger pileus (up to 60 mm diam.) with deeply
sulcate margin, distant lamellae, longer stipe (60–77 mm
long), absence of any strong odour, and presence of only
Siccus-type caulocystidia (Singer 1976).
Marasmius pellucidus Berk. & Broome, although phylogenetically related as revealed in Fig. 135, possess
Globulares-type of cells in the pileipellis and is categorized
under a sect. Globulares (Wannathes et al. 2009). The other
distinguishing features of M. pellucidus from that of
M. indojasminodorus includes a plano-campanulate to
almost applanate pileus, ivory to pale orange towards disc
with white to cream or pale yellowish white margin, and
presence of comparatively more lamellae (up to 26;
Wannathes et al. 2009).
Omphalotaceae Bresinsky
The family Omphalotaceae was proposed by Bresinsky
in Kämmerer et al. (1985) to accommodate the genera
Omphalotus Fayod and Lampteromyces Singer. The
members of the family are characterized by fleshy, brightly
coloured and often luminescent basidiomes with a glabrous
or fibrillose pileus, decurrent lamellae, central or eccentric
stipe devoid of any veil, hyaline, smooth, globose or
ellipsoid and inamyloid basidiospores, clavate basidia often
intergrading with cystidioles, the absence of cystidia and
monomitic hyphal system with clamp connections (Cannon
and Kirk 2007). Species of Omphalotaceae are saprobic or
necrotrophic on wood or litter (Moncalvo et al. 2002).
Moncalvo et al. (2002), in their broad systematic treatment
of the euagarics using LSU sequence, observed that several
genera that were traditionally classified in various families
or tribes of Agaricales (Singer 1986) and Caripia montagnei, a reduced form that was generally placed in the
Stereales (Hawksworth et al. 1995) were clustered together
in Omphalotaceae clade together with lentinuloid and
omphalotoid species. In their overview of phylogeny of
Fungal Diversity
Fig. 135 Consensus phylogram (50% majority rule) obtained from
MCMC analysis of one million generations from a Bayesian inference
analysis based on ITS region for Marasmius taxa and two outgroup
sequences (Crinipellis brunneipurpurea and C. malesiana). Maximum likelihood (RAxML) bootstrap support (left) C 50% and
Bayesian posterior probabilities (right) C 0.50 are indicated above
the nodes. Sequences used in this study mostly have been sampled
Fig. 136 Marasmius indojasminodorus (CAL 1514, holotype).
Scale bar 5 mm. Photo by A.K. Dutta
major clades of Agaricales, Matheny et al. (2006) recovered Omphalotaceae as monophyletic family.
Marasmiellus Murrill
from a previous study (Tan et al. 2009; Wannathes et al. 2009). The
newly generated previously described type specimen sequence of
Marasmius midnapurensis and the sequences of the newly described
taxa for the present study are placed in blue font to highlight its
phylogenetic position in the tree. Ex-type strains are in bold.
GenBank accession numbers for all of the sequences are indicated
in the tree
Marasmiellus is a predominantly tropical and subtropical genus that encompasses more than 400 species known
worldwide (Wilson and Desjardin 2005; Blanco-Dios
2015). Traditionally, the genus had been placed in family
Tricholomataceae (Singer 1986). While Kirk et al. (2008)
treated it in the Marasmiaceae, DNA-based molecular
phylogenetic studies of Moncalvo et al. (2002) and Wilson
and Desjardin (2005) indicated that this genus belongs to
the Omphalotaceae. Marasmiellus is characterized by
collybioid, omphalioid or pleurotoid basidiomes, slightly
decurrent, intervenose lamellae, smooth, white or off-white
and inamyloid basidiospores, and pileipellis or stipipellis
showing a Rameales-structure (Singer 1973; Pérez-DeGregorio et al. 2011). Most of the species are saprobes
occurring on dead and rotting plant material and some are
host specific. A few species are found on living hosts.
According to Wilson and Desjardin (2005), the genus is
polyphyletic. In the course of our studies on the agarics of
Kerala State, India, we came across a remarkable species of
Marasmiellus. It is formally described here as a new
123
Fungal Diversity
Fig. 137 Marasmius
indojasminodorus (CAL 1514,
holotype). a Basidiomes.
b Basidiospores. c Basidioles.
d Siccus-type cheilocystidia.
e Siccus-type cells of the
pileipellis. f Caulocystidia
showing setulose and nonsetulose cells. Scale bars
a = 5 mm, b–f = 5 lm.
Drawings by A.K. Dutta
species based on both morphology and molecular phylogeny (Fig. 138).
Marasmiellus bicoloripes K.P.D. Latha, K.N.A Raj &
Manim., sp. nov.
MycoBank number: MB820689; Facesoffungi number:
FoF03251, Figs. 139, 140
Etymology: Referring to the bicoloured stipe of the
basidiomes.
Holotype: CAL 1524.
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978) and the Online Auction
Color Chart (Anonymous 2004).
Basidiomes small, somewhat omphalinoid. Pileus
5–20 mm diam., initially convex, becoming plano-convex
with a small umbo surrounded by a shallow depression at
123
maturity; surface brown (6F4/OAC733) all over when
young, becoming greyish brown (6D3/OAC730) at the centre
and brownish beige (6E3/OAC723) towards the margin, not
hygrophanous, faintly pellucid-striate, appressed-fibrillose
all over; margin decurved to somewhat straight, initially
entire, becoming somewhat wavy. Lamellae adnate or subdecurrent, somewhat thick and waxy, often furcate and
anastomosing, close, brownish orange (6E2/OAC724), up to
2 mm wide, with lamellulae of 2–3 tiers; edge finely torn,
concolorous with the sides. Stipe 12–14 9 1–2 mm, central,
terete, equal or slightly tapering towards the base, solid;
surface initially white all over, becoming dark grey (5E1,
5F1/OAC904) towards the base at maturity, appressed-fibrillose all over. Odour and taste not distinctive. Lamellar
trama subregular, hyphae 3–9 lm wide, hyaline, thin-walled, inamyloid. Subhymenium poorly developed
Fungal Diversity
pseudoparenchymatous. Basidia 20–31 9 4–6 lm, clavate
or occasionally pedicellate clavate, hyaline, thin-walled, 4or 2-spored; sterigmata up to 5 lm long. Basidiospores
4–8 9 3–4(–5) (6.5 ± 1.5 9 3.5 ± 0.6) lm, Q = 1.3–2.7,
Qm = 1.9, oblong-ellipsoid, pip-shaped or subfusiform, often
with a stretched-out apicular end, with prominent guttules,
hyaline, inamyloid, smooth, thin-walled. Lamellar edge
heterogeneous. Pleurocystidia absent. Cheilocystidia
20–32 9 4–8 lm, scarce, scattered, versiform: clavate with
short or long, apical diverticulate projections or with a
mucronate apex, cylindrical with a mucronate apex, utriform
with a rostrate apex or flexuous, thin-walled, hyaline. Pileus
trama interwoven; hyphae 3–10 lm wide, hyaline, thinwalled, inamyloid. Pileipellis a differentiated cutis with
occasional nodulose-diverticulate hyphae; hyphae 3–18 lm
wide, with a yellowish brown plasmatic pigment and strong,
yellowish brown spiral encrustions, thin- to slightly thickwalled. Stipitipellis an epicutis; hyphae 3–6 lm wide, hyaline or with a yellowish brown plasmatic pigment, thickwalled (up to 1 lm thick), often giving rise to short or long,
sometimes diverticulate side branches, 17–108 9 3–5 lm,
slightly thick-walled towards the base. Caulocystidia absent.
Clamp connections observed on all hyphae.
Material examined: INDIA, Kerala State, Ernakulam
District, Pooyamkutty, 22 November 2014, K.P.D. Latha,
DKP331 (CAL 1524, holotype).
Host and habitat: In small group among on a decaying
twig.
GenBank
numbers:
ITS = KY807129,
LSU = KY817233.
Notes: Marasmiellus bicoloripes is characterised by
omphalinoid basidiomes with a greyish brown, faintly
striate and fibrillose pileus, brownish orange, anastomosing
and somewhat thick and waxy lamellae, a white stipe
turning dark grey towards the base, oblong-ellipsoid to
subfusiform basidiospores, a hymenium devoid of pleurocystidia, a heterogeneous lamellar edge with scattered
cheilocystidia, a cutis-type pileipellis with occasional
nodulose-diverticulate hyphae, an epicutis-type stipitipellis
with frequent nodulose-diverticulate hyphae and presence
of clamp connections. Owing to the combination of characteristics such as the stipe tending to greyish at the base,
the cutis-type pileipellis with occasional diverticulate
hyphae and the short basidiospores (less than 10.4 lm),
this species can be placed either in sect. Candidi (Bat.)
Sing. or in sect. Dealbati Sing. (Singer 1986) of
Marasmiellus.
Marasmiellus cibodasensis Retnowati, an Indonesian
species described invalidly by Retnowati (2012), differs in
having a flattened pileus with a wrinkled surface, subdistant,
non-anastomosing lamellae, an apically tapered, granulose
and bulbous stipe which is not darkening towards the base,
ellipsoid basidiospores, a fertile lamellar edge devoid of
cheilocystidia and a gregarious habit on wood. Marasmiellus
rhizomorphigenus Antonı́n et al., known from Republic of
Korea (Antonı́n et al. 2010), differs in having a greyish to
whitish pileus with a pubescent to tomentose surface, whitish
to pale yellowish and distant lamellae, a stipe with a pubescent to furfuraceous surface, presence of rhizomorph, larger basidiospores (13.5–17 9 4.5–6.5 lm), well-developed,
larger hymenial cystidia (34–70 9 8–14 lm) and presence
of pileo- and caulocystidia. Marasmiellus koreanus Antonı́n
et al. also described from Republic of Korea (Antonı́n et al.
2010), has larger, robust basidiomes with a brownish orange,
rugulose pileus, light yellow lamellae, whitish to light yellow
stipe, versiform cheilocystidia and stipitipellis with copious
caulocystidia. Marasmiellus candidus (Fr.) Singer, a widespread species in Europe (Noordeloos 1995), differs in having a pure white, pruinose to tomentose pileus with a greybrown centre, a pruinose stipe with a bulbous base, oblong to
cylindrical and larger basidiospores [(10.5–)11.5–15(–
17.5) 9 (7.5–)8.0–12.5 lm], copious and larger cheilocystidia (45–90 9 6–12.5 lm), an irregular cutis-type
pileipellis with a transition to a trichoderm and the stipitipellis with caulocystidia.
The phylogenetic trees inferred from the ML and BI
analyses of the ITS sequence dataset show identical
topology and the ML phylogram is presented in Fig. 138.
The tree reveals two distinct clades, the Tetrapyrgos and
the Marasmiellus clades, as shown by Wilson and Desjardin (2005) and subsequently confirmed by Antonı́n et al.
(2010). Marasmiellus bicoloripes nests in the Tetrapyrgos
clade with 82% ML and 1.00 BYPP support. Within this
clade, M. bicoloripes together with collections of M. rhizomorphigenus, M. tenerrimus (Berk. & M.A. Curtis)
Singer and M. candidus formed a cluster where M. bicoloripes was found to be a lineage distinct from other species
with significant support (73% ML and 1.00 BYPP). It has
to be emphasized here that the clade Tetrapyrgos does not
exactly correspond to the genus Tetrapyrgos as the latter is
primarily characterised by tetrahedral basidiospores and a
stipe arising from a basal pad (Honan et al. 2015) while
some species including M. bicoloripes and M. candidus
that form part of the clade Tetrapyrgos do not show these
features.
Psathyrellaceae Vilgalys et al.
Based on molecular studies, Redhead et al. (2001)
introduced the family Psathyrellaceae to accommodate the
genera Psathyrella (Fr.) Quél. and Lacrymaria Pat., together with related species in the polyphyletic genus Coprinus sensu lato, which were transferred to the genera
Coprinellus P. Karst., Coprinopsis P. Karst. or Parasola
Redhead et al. A few species related to Agaricus remained
in Coprinus sensu stricto. Therefore, the concept of family
Coprinaceae Overeem & Weese was abandoned and
123
Fungal Diversity
Coprinus sensu stricto was transferred to Agaricaceae
Chevall. Afterwards, several smaller genera were included
in Psathyrellaceae. Kirk et al. (2008) consider the family to
contain 12 genera.
Coprinopsis P. Karst.
Coprinopsis is a large genus that contains around 200
species (Kirk et al. 2008). They are saprotrophic on soil,
wood, vegetable refuse, dung or burnt ground, and produce
agaricoid, rather fragile basidiomes which are often short
lived and/or deliquescent, with black to blackish spore
deposit (Vesterholt 2012). The phylogenetic tree is presented in Fig. 141.
Coprinopsis kubickae (Pilát & Svrček) Redhead, Vilgalys
& Moncalvo, in Redhead et al., Taxon 50(1): 229 (2001)
Facesoffungi number: FoF05721, Fig. 142
Basionym: Coprinus kubickae Pilát & Svrček, Česká
Mykol. 21: 142 (1967).
Holotype: Czechoslovakia, on culm of Juncus.
Pileus up to 10 mm wide when expanded, sphaerical at
first, then ellipsoid to ovoid, paraboloid, convex, finally
applanate or plano-concave with revolute margin, plicatesulcate except at the centre, whitish to pale brown when
young, then pinkish, finally brownish grey except brownish
central disc, non-deliquescent, covered with scarce, minute, brownish to brown flocculose veil. Lamellae free,
distant, L = 22–34, l = 1–3, white at first, becoming
pinkish, then reddish brown with whitish and flocculose
edge, finally brown-black and deliquescent. Stipe
16–28 9 0.8–1.2 mm, cylindrical or gradually thickened
towards the base, which is often (sub) bulbose or with basal
disc (up to 2 mm wide), central, hollow, dry, minutely
fibrillose-floccose, often tomentose at the base, white.
Odour and taste not observed. Spore print brown-black.
Basidia 14–31 9 9–12 lm, clavate, 4-spored, thin-walled,
hyaline, surrounded by 5–8 hymenophysalides (pseudoparaphyses). Basidiospores [100/2/1] 8–9.6–11.1 9 6.9–8.2–
9.3 lm, Q = (1–)1.05–1.17–1.30(–1.38), mostly subglobose, broadly ellipsoid or ovoid, but also globose and
ellipsoid, not flattened, with rounded to slightly conical
base and rounded apex, smooth, thin-walled to moderately
thick-walled (up to 0.7 lm), with 1.2–1.8 lm wide (inner
diameter), central to eccentric germ-pore, medium rusty
brown in H2O and NH4OH, medium brown in KOH, but
soon becoming medium brown grey, semi-transparent,
non-amyloid
and
non-dextrinoid.
Cheilocystidia
30–90 9 9–24 lm, narrowly utriform to utriform, oblong,
subcylindrical, conical or narrowly clavate, thin-walled to
moderately thick-walled (up to 0.7 lm), hyaline, scattered.
Pleurocystidia 50–110 9 13–24 lm, similar to cheilocystidia, but somewhat more elongated, mostly narrowly
utriform, narrowly conical or subcylindrical, scattered. Veil
composed of 2–6 lm broad, branched, thin-walled, rather
123
sparsely diverticulate, hyaline to light yellowish brown
hyphae, mostly with minutely to coarsely encrusted, light
to dark brown pigment. Pileipellis a cutis, composed of
repent, hyaline, thin-walled, 1.5–5 lm wide hyphae. Stipitipellis a cutis of parallel, repent, thin-walled, hyaline,
1–8 lm wide hyphae, sparsely diverticulate, with ascending hyphal tufts in places. Clamp connections present and
abundant in all tissues.
Material examined: CROATIA, Karlovac County,
Fishpond Draganići, 10 km NE from Karlovac,
45°330 4000 N, 15°380 2100 E, 107 m a.s.l., 7 September 2013,
leg. M. Čerkez, CNF 1/6614.
Host and habitat: Solitary or in small groups on dead
Phragmites, Typha, Juncus, Carex, Glyceria and Acorus
species in wet habitats, as well as on rich soil and rotten
straw in greenhouses (Uljé 2005; Nagy 2007; Gierczyk
et al. 2011). From Canada reported on wood submerged in
an alkali lake (Anastasiou 1967; as Coprinus amphibius
Anastasiou). Our collection was collected on wet remnants
of Phragmites australis, on the marshy edge of a fishpond.
Distribution: Known from about 15 European countries
and Canada, rare.
GenBank number: ITS = MH422562.
Notes: Coprinopsis kubickae is primarily characterized
by its branched, diverticulate and thin-walled veil elements, presence of clamp connections, smooth, globose to
broadly ellipsoid, not flattened spores, rather small basidiomes and by living on plant (almost always herbaceous)
remnants in wet habitats. Morphologically (according to
branched and diverticulate veil), it belongs to section
Alachuani (Singer) D.J. Schafer. However, the morphological concept of section Alachuani is not supported by
molecular data and several Alachuani species are mixed
with species of sections Atramentarii (Fr.) D.J. Schafer and
Lanatuli (Fr.) D.J. Schafer in our ITS based phylogram
(Fig. 141). Still, Coprinopsis kubickae nests in the clade
with the rest of Alachuani species. We report Coprinopsis
kubickae as new to Croatian mycobiota.
Boletales E.-J. Gilbert
Boletaceae Chevall.
Boletaceae is an important Basidiomycete family with a
global distribution, most of which form ectomycorrhizal
associations with trees. Most species have been described
from temperate regions, and little research had been carried
out on tropical boletes before 2010. In the past decade,
however, phylogenetic analyses based on multigene dataset
and including tropical taxa, have greatly improved our
understanding of the systematics of the group, with many
new genera and species being published (e.g., Halling et al.
2012; Hosen et al. 2013; Wu et al. 2015, 2016; Henkel
et al. 2016). This molecular revolution has left a number of
Boletaceae species (especially in the large genus Boletus)
Fungal Diversity
Fig. 138 Phylogram derived from maximum likelihood (RAxML)
analysis using ITS sequence dataset. Maximum likelihood (RAxML)
bootstrap support (left) C 50% and Bayesian posterior probabilities
(right) C 0.90 BYPP are indicated above the nodes. Marasmiellus
bicoloripes is highlighted in blue. Marasmius rotula and M. capillaris
are selected as the outgroup taxa
to be re-examined and recombined in the light of modern
classification of the family. A novel species, Baorangia
major is introduced based on distinct morphological characteristics coupled with phylogenetic analysis of a combined ATP6, RPB2, and TEF1-a dataset (Fig. 143). Two
new combinations, Baorangia rufomaculata and Lanmaoa
pallidorosea are also designated in this study.
Baorangia G. Wu & Zhu L. Yang
Baorangia was first recognized by Wu et al. (2014, as
‘clade 51’), and later formally described by Wu et al.
(2015). Typical characteristics are a thin hymenophore that
is 3–5 times thinner than the pileus context, yellow tubes
and pores that stain blue when bruised, a light yellow
context that slowly stains pale blue when cut, and a
123
Fungal Diversity
Fig. 139 Marasmiellus
bicoloripes (CAL 1524,
holotype). a, b Basidiomes in
the field. Scale bars a,
b = 10 mm. Photos by K.
P. Deepna Latha
trichodermium to interwoven trichodermium pileipellis
(Wu et al. 2015). Three species are recorded in the genus,
B. bicolor (Kuntze) G. Wu et al., B. emilei (Barbier) Vizzini et al. and B. pseudocalopus (Hongo) G. Wu & Zhu L.
Yang, which is the generic type (Kirk et al., continuously
updated).
Baorangia major Raspé & Vadthanarat, sp. nov.
MycoBank number: MB824250; Facesoffungi number:
FoF05722, Figs. 144, 145
Etymology: The specific epithet ‘‘major’’ from Latin,
refers to the size of mature basidiomes, the largest in the
genus.
Holotype: MFLU 12-0040
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes large when mature. Pileus (7–)16–22(–23) cm
diam., at first hemispherical to convex, becoming convex to
plano-convex and sometimes slightly depressed at the centre,
greyish red (11C–E5) to greyish ruby (12B–E5), more rarely
greyish pink (around 10A3), paler in spots and in age; margin
involute at first, later inflexed, slightly exceeding; concolorous or abruptly paler (yellowish white; 2A2); surface even,
dry, dull, subtomentose; context 1.7–3.8 cm thick half-way
to the margin, firm at first, soft in age, off-white to yellowish
white (1A2), slightly marmorated above the hymenophore,
blueing when bruised, sometimes only weakly, then slowly
getting yellow (3A5–6) with time, as in worm wounds. Stipe
central, terete, cylindrical to subclavate (4.8–)5.2–8.5(–
15) 9 1.5–3.3(–4.5) cm; surface finely and densely dotted,
subceraceous, dull, with off-white to pale yellow basal
tomentum; context solid, yellowish white (2A2) at the top,
more intensely yellow towards the base, marmorated, often
123
with dark red spots, especially in the lower half, quickly and
intensely blueing when cut. Hymenophore tubulate, decurrent; tubes 2–5 mm long, not separable, yellow (2–3A7),
sometimes stained red at places (old bruises) or when old,
quickly and intensely blueing when bruised; pores 2–5 mm
wide at mid-radius when mature, mostly angular, elongated
and radially arranged near the stipe, yellow (2A8), quickly
and intensely blueing when bruised, sometimes reddish when
very old. Odour fungoid to slightly fruity. Taste mild, fungoid. Spore print olive-brown. Macrochemical reactions:
KOH pale orange on pileus and stipe, paler on context, null or
merely slightly orangish on hymenium; NH4OH null. Basidia 4-spored, (36–)36–42.2–55(–55) 9 (8.5–)8.5–9.3–
11(–11) lm (n = 20), narrowly clavate, sometimes curved,
hyaline, with sterigmata up to 5 lm long, without basal
clamp connection. Basidiospores (6–)7.5–8.1–9(–10) 9 (4–
)4–4.6–5(–5.5) lm Q = (1.44–)1.5–1.8–2.1(–2.4) [N = 5/5/
265]. From the type (7.5–)8–8.6–9.5(–9.5) 9 (4–)4–4.5–5(–
5) lm, Q = (1.6–)1.69–1.9–2.13(–2.13) [n = 55], ellipsoid
to ovoid, thin-walled, smooth, yellowish to greenish hyaline
in water, KOH or NH4OH, yellowish in Melzer’s reagent.
Cheilocystidia thin-walled, hyaline, of two different shapes,
the first clavate to broadly clavate, (21–)21–29.3–37(–
37) 9 (11.5–)11.5–13.1–15(–15) lm, the second fusiform to
broadly fusiform, (26–)26–38.9–57(–57) 9 (10–)10–13.3–
15(–15) lm. Pleurocystidia infrequent, fusiform to narrowly
fusiform with obtuse apex, (44–)44–54.4–70(–70) 9 (8–)8–
9.5–12(–12) lm, thin-walled, hyaline. Hymenophoral trama
divergent, 90–112 lm wide, composed of 4–10 lm wide,
hyaline hyphae, with regular mediostratum 30–45 lm wide.
Pileipellis a trichoderm to tangled trichoderm, 170–190 lm
thick, made of moderately interwoven and somewhat anastomosing, thin-walled, hyaline hyphae; terminal cells
Fungal Diversity
Fig. 140 Marasmiellus bicoloripes (CAL 1524, holotype). a Basidiospores. b Cheilocystidia. c Basidium. d Clamped hypha of
pileipellis. e Pileipellis. f Stipitipellis. g Pileipellis hypha showing
spiral encrustations. Scale bars a–c = 10 lm, d–g = 20 lm. Photos
by K. P. Deepna Latha and K. N. Anil Raj
cylindrical with rounded apex, 20–58 9 4–9 lm, mostly
hyaline to slightly pale yellowish brown at places in KOH.
Pileus context made of moderately interwoven, hyaline, thinwalled hyphae, 4–13 lm wide. Stipitipellis a hymeniderm,
62–88 lm thick, mostly composed of basidiole-like cells,
12–43 9 4–10 lm. Caulocystidia infrequent, of three
shapes, the first clavate to broadly clavate, (13–)23.8(–
31) 9 (10–)12.5(–15) lm, thin-walled, hyaline, the second
fusiform to broadly fusiform, (21–)31.5(–45) 9 (9.5–
)11.6(–15), thin-walled, hyaline, and the third cylindrical,
slightly curved, mostly hyaline, at places slightly yellowish
brown in KOH. Stipe context composed of parallel, 4–12 lm
wide, hyaline and somewhat anastomosing hyphae. Clamp
connections not seen in any tissue.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng District, around Mushroom Research Center,
N19°07.10 –E98°44.00 , elev. 900 m, 2 July 2010, P.
Sysouphanthong, OR070 (MFLU!, BR!); ibid., N19°07.20 –
E98°43.90 , elev.915 m, 17 June 2011, O. Raspé & S.C.
Karunarathna, OR197 (MFLU!, BR!); N19°06.50 –
E98°44.50 , elev.1075 m, 18 June 2011, O. Raspé & S.C.
Karunarathna, OR209 (MFLU, holotype; BR, isotype);
N19°06.60 –E98°44.50 , elev.1055 m, 7 June 2012, O. Raspé
& K. Wisitrassameewong, OR404; N19°07.20 –E98°43.90 ,
elev. 910 m, 29 July 2013, O. Raspé & B. Thongbai,
123
Fungal Diversity
Fig. 141 Maximum likelihood
phylogenetic tree of
Coprinopsis kubickae and
related species based on ITS
sequence alignment, calculated
with MEGA6 software (Tamura
et al. 2013). The sequence
obtained in this study is shown
in blue. Ex-type strains are
indicated in bold. Maximum
likelihood bootstrap values
greater than 50% are indicated
at the nodes. The tree is rooted
with Coprinellus xanthothrix
and C. domesticus. The bar
indicates the number of
nucleotide substitutions per site
OR657 (MFLU!, BR!); CHINA, Yunnan Province, Cangyuan County, Banhongxiang, along the road, N23°18.60 –
E99°05.30 , elev. 1010 m, 10 July 2012, O. Raspé & R.L.
Zhao OR486 (HMAS).
Habit and habitat: Mostly gregarious, sometimes fasciculate or solitary, on soil in forests dominated by Lithocarpus spp., Castanopsis spp., sometimes mixed with
Dipterocarpus spp.
Distribution: Known from Chiang Mai Province, Thailand and Yunnan Province, China.
GenBank numbers: ATP6 = MG897421, RPB2 =
MG897441, TEF1-a = MG897431 (OR209, holotype);
ATP6 = MG897422,
RPB2 = MG897442,
TEF1a = MG897432 (OR404), ATP6 = MG897423, RPB2 =
MG897443, TEF1-a = MG897433 (OR486).
Notes: Baorangia major can easily be recognised in the
field by its large basidiomes when mature, decurrent
hymenophore with large, angular, radially arranged (at
least near the stipe), and 1–3 times compound pores, red
stipe that immediately and intensely turns blue when
injured or merely touched. In their diagnosis of Baorangia,
Wu et al. (2015) mentioned ‘‘context which stains pale blue
slowly when cut’’. While this fits B. major pileus context, it
does not fit its stipe context, which quickly and intensely
stains blue when cut. Baorangia bicolor (Kuntze) G. Wu
et al. and B. rufomaculata (see below) somewhat resembles
123
B. major, but the latter can easily be distinguished by the
large angular pores and intense blueing.
Baorangia rufomaculata (Both) Raspé & Vadthanarat,
comb. nov.
MycoBank number: MB824251; Facesoffungi number:
FoF05723, Fig. 146
Basionym: Boletus rufomaculatus Both, Bull. Buffalo
Soc. Nat. Sci. 36: 221 (1998)
Holotype: USA, Western New York, under Fagus
grandifolia Ehrh., Quercus rubra L., Tsuga canadensis (L.)
Carrière and Acer saccharum Marshall, Both 2831 (BUF).
Basidia 4-spored, (21–)21–24.6–31(–31) 9 (8–)8–8.9–
10(–10) lm, clavate, hyaline, sterigmata 3–5 lm long,
without basal clamp connection. Basidiopores (9–)9–10.6–
12(–12.5) 9 (3.5–)3.5–3.9–4.5(–4.5) lm Q = (2.1–)2.3–
2.7–3.2(–3.6) [N = 50], subcylindrical with slight
suprahilar depression, thin-walled, smooth, slightly greenish to yellowish hyaline in water, yellowish hyaline in 5%
KOH or NH4OH, brownish yellow in Melzer’s reagent.
Cheilocystidia (18–)18–31.4–45(–45) 9 (6–)6–7.9–10(–
10) lm, frequent, fusiform to broadly fusiform, thin-walled, hyaline. Pleurocystidia (42–)42–59.5–72(–72) 9 (7–
)7–8.2–9.5(–9.5) lm, infrequent, narrowly fusiform, thinwalled, hyaline. Hymenophoral trama regular to slightly
divergent, 90–190 lm wide, composed of (3–)5–7(–12)
lm wide hyaline hyphae. Pileipellis a subcutis to tangled
Fungal Diversity
Fig. 142 Coprinopsis kubickae (CNF 1/6614). a Basidiomes. b Veil on the pileus. c Basidiospores. d–f Cheilocystidia. Scale bars a = 5 mm,
b = 30 lm, c–f = 10 lm
trichoderm, 120–220 lm thick, made of loosely to moderately interwoven, thin-walled, hyaline hyphae; terminal
cells cylindrical with rounded apex, 22–45 9 5–8 lm,
mostly hyaline, at places yellowish hyaline in KOH. Pileus
context made of moderately interwoven hyaline, thin-walled hyphae, 3–12 lm wide. Stipitipellis a hymeniderm,
85–120 lm thick; terminal cells clavate to narrowly clavate with rounded apex, thin-walled, 14–47 9 3–12 lm;
below the hymeniderm parallel, hyaline hyphae anastomosing at places. Stipe context composed of parallel,
branched, hyaline, thin-walled hyphae, 7–14 lm wide.
Caulocystidia rare, 23–30 9 7–9 lm, fusiform or clavate,
thin-walled, hyaline. Clamp connections not seen in any
tissue.
Material examined: USA, New York, Erie Co., Town of
Orchard Park, Chestnut Ridge Park, N42°43.10 –W78°45.30 ,
elev. 330 m, 6 August 1997, E. Both 4144 (CFMR).
Distribution: North America.
GenBank numbers: ATP6 = MG897415, RPB2=
MG897435, TEF1-a = MG897425.
Notes: The specimen we studied (Both 4144) was
already sequenced by Nuhn et al. (2013) for LSU, RPB1,
and TEF1-a genes, but under the wrong number ‘‘4414’’.
Our TEF1-a sequence is 100% identical to the one published by Nuhn et al. (2013), except for the insertion of an
‘‘N’’ at position 215 in the latter, which is likely an artefact.
Our measurements of microscopic characteristics slightly
deviate from the original description of Both (1998).
However, the specimen we studied was collected and
identified by Both from the type locality. Therefore, we are
confident that it belongs to the same species.
Lanmaoa G. Wu & Zhu L. Yang
The genus Lanmaoa was recognized by Wu et al. (2014,
as ‘clade 49’), and later formally described by Wu et al.
(2015). The typical characteristics of the genus are a thin
hymenophore which is 3–5 times thinner than the pileus
context and stains blue when bruised, a light yellow context that slowly stains pale blue when cut, and an interwoven trichodermium to subcutis pileipellis (Wu et al.
2015). Seven species are recorded in the genus, L. angustispora G. Wu & Zhu L. Yang, L. asiatica G. Wu &
Zhu L. Yang (generic type), L. carminipes (A.H. Sm. &
Thiers) G. Wu et al., L. flavorubra (Halling & M. Mata) G.
Wu et al., L. fragrans (Vittad.) Vizzini et al., L. pseudosensibilis (A.H. Sm. & Thiers) G. Wu et al. and
123
Fungal Diversity
Fig. 143 Maximum likelihood tree inferred from three-gene dataset
(ATP6, RPB2 and TEF1-a). Retiboletus griseus, R. fuscus, and
Rhodactina rostratispora were used as the outgroup taxa. Sequences
retrieved from GenBank were originally published in Halling et al.
(2012), Wu et al. (2014, 2015, 2016), Zhao et al. (2014a, b), Raspé
123
et al. (2016), and Vadthanarat et al. (2018). Maximum likelihood
bootstrap values greater than 70% are indicated at the nodes. The bar
indicates the number of nucleotide substitutions per site. Type
specimens are indicated in bold. Newly generated sequences are in
blue
Fungal Diversity
Fig. 144 Baorangia major basidiomes in the field. a, b OR209 (holotype). c OR404. d OR486. Scale bars 3 cm
L. roseocrispans Bessette et al. (Kirk et al., continuously
updated).
Lanmaoa pallidorosea (Both) Raspé & Vadthanarat,
comb. nov.
MycoBank number: MB824252; Facesoffungi number:
FoF05724, Fig. 147
Basionym: Boletus pallidoroseus Both, Bull. Buffalo
Soc. nat. Sci. 36: 218 (1998).
Holotype: USA, New York, on ground under Quercus
rubra and Fagus grandifolia, Both 3026 (BUF).
Basidiospores and basidia not observed on the immature
specimen studied. Cheilocystidia (14–)14–19.4–25(–
25) 9 (5–)5–6.5–8(–8) lm, frequent, fusiform, thin-walled,
hyaline. Pleurocystidia (28–)28–33.9–39(–39) 9 (6.5–
)6.5–7.9–9(–9) lm, infrequent, narrowly fusiform, thinwalled, hyaline. Hymenophoral trama divergent,
110–175 lm wide, composed of (3.5–)4–7(–12) lm wide
hyaline hyphae. Pileipellis a trichoderm, 220–300 lm thick,
composed of thin-walled, hyaline hyphae; terminal cells
cylindrical with rounded apex, 28–70 9 4–8 lm, mostly
hyaline, at places brownish to yellowish hyaline in KOH.
Pileus context made of strongly interwoven, hyaline, thinwalled hyphae, 4–6 lm wide. Stipitipellis a hymeniderm,
55–80 lm thick, composed of pointing out, clavate, thinwalled terminal cells 13–28 9 4–9 lm, which are mixed
with caulocystidia. Stipe context composed of interwoven,
sub-irregular, thin-walled, hyaline hyphae, 4–9 lm wide.
Caulocystidia of two types, the first one very frequent, fusiform to broadly fusiform, (19–)19–28.4–39(–39) 9 (7–)7–
8.4–11(–11.5) lm, thin-walled, hyaline, the second one
infrequent, broadly clavate, (11–)11–20.5–27(–27) 9 (7–
)7–9.4–11(–11) lm, thin-walled, hyaline. Clamp connections not seen in any tissue.
Material examined: USA, New York, Erie Co., Orchard
Park, Chestnut Ridge Park, N42°43.10 –W78°45.30 , elev.
330 m, 19 August 2000, E. Both 4432 (CFMR).
Distribution: North America.
GenBank numbers: ATP6 = MG897417, RPB2 =
MG897437, TEF1-a = MG897427.
Notes: Although the specimen we examined is immature
and we could not check basidiospore and basidia dimensions, we are confident that it belongs to the species
described by Both (1998). Indeed, it was collected and
identified by E. Both himself from the type locality.
Cantharellales Gäum.
Clavulinaceae Donk
Clavulinaceae was established by Donk (1933) as a
tribe, but later Donk (1961) raised the tribe to the rank of
family. The family Clavulinaceae comprises four genera,
Burgella Diederich & Lawrey, Clavulina J. Schröt.,
Membranomyces Jülich and Multiclavula R.H. Petersen
(Kirk et al. 2008). We follow the treatment and updated
123
Fungal Diversity
Fig. 145 Baorangia major
microscopic features.
a Basidiospores. b Basidia.
c Two types of cheilocystidia.
d Pleurocystidia. e Two types of
caulocystidia. f Pileipellis.
g Stipitipellis. Scale bars
a = 10 lm, b–e = 20 lm, f,
g = 50 lm. All drawings were
made from the type OR209
accounts of Clavulinaceae in He et al. (2016) and Tibpromma et al. (2017). A novel species, Clavulina thindii is
introduced based on distinct morphology and maximum
likelihood phylogeny of ITS sequence data (Fig. 148).
Clavulina J. Schröt.
Clavulina is distributed worldwide and characterized by
clavarioid to coralloid, occasionally resupinate and effused,
123
simple or branched basidiomes, monomitic hyphal system,
with clamp connections in most species; smooth, hyaline,
subglobose to broadly ellipsoid basidiospores usually with
one large guttule; basidia with two or up to six cornuted
sterigmata and often develop a transversal septum (Corner
1950, 1970; Petersen 1988a; Henkel et al. 2005, 2011;
Uehling et al. 2012a; He et al. 2016). Species of Clavulina
Fungal Diversity
occur in forests dominated by ectomycorrhizal (ECM) trees
of the leguminous genera (Fabaceae) (Henkel et al. 2011).
Approximately 82 species have been described in the genus
Clavulina (Corner 1950, 1970; Thind 1961; Petersen
1967, 1983, 1985, 1988a, b; Thind and Sharda 1984;
Roberts 1999; Thacker and Henkel 2004; Henkel et al.
2005, 2011; Douanla-Meli 2007; Duhem and Buyck 2007;
Trappe and Castellano 2007; Olariaga and Salcedo 2012;
Uehling et al. 2012a, b; Wartchow 2012; He et al. 2016).
Clavulina thindii U. Singh, sp. nov.
MycoBank number: MB824527; Facesoffungi number:
FoF04878, Figs. 149, 150
Etymology: To commemorate Prof. K.S. Thind for his
contribution to Indian mycobiota.
Holotype: CAL 1661
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Basidiomes 41–66 9 30–49 mm, coralloid, terrestrial,
profusely branched, solitary to scattered; sterile stipe
12–28 9 7–12 mm, white to greyish white, glabrous; fertile
branches 30–38 9 2.5–4.5 mm, forming 4–7 ranks in multiple planes, branching pattern polychotomous to dichotomous-ascending, initially violet white to pale violet
(16A2–3), maturing to lilac grey to lilac (16B2–3), wax white
to greyish yellow (2B3–4) with drying, smooth; hymenium
ripening over distal two-thirds of basidiomes, amphigenous,
interface with stipe clearly demarcated; apices roundedacuminate to irregularly coronate or blunt, concolorous with
branches when young, becoming brownish orange (6C8) at
maturity; context subsolid to hollow, concolorous with
branches. Taste and Odour pleasant. Basidiospore deposit
not obtained. Basidia 22–48 9 6–10 lm, subcylindrical to
subclavate or clavate, with numerous granular contents, postpartial septa observed on some basidia, basally clamped,
2-spored; sterigmata up to 7 lm long, cornuted; basidioles
numerous, subcylindrical to clavate. Hymenophoral tramal
hyphae sub-regular to irregular; noninflated tramal hyphae
5–8 lm, smooth, thin-walled, hyaline, lacking internal contents; inflated tramal hyphae 11–15 lm, smooth, thin-walled,
hyaline, lacking internal contents. Cystidia absent. Basidiospores 6.5–7.5–8 9 6–6.4–7 lm (n = 30, Q = 1.07–1.15–
1.25), subglobose to broadly ellipsoid, smooth, pale yellow in
KOH, inamyloid, with one translucent guttule; apicules up to
1 lm long. Clamp connections present.
Material examined: INDIA, Uttarakhand, Pauri Garhwal
District, Gangoti-Adwani forest, alt. 1948 m, N 30°05.7990
E 078°43.9270 , 3 October 2016, U. Singh, US 1428 (CAL
1661, holotype); ibid., 31 August 2017, U. Singh, US
1585.
Host and habitat: Solitary to scattered, on ground, under
trees of Acacia dealbata A. Cunn. in mixed temperate
forest.
Distribution: Known only from India.
GenBank numbers: ITS = MG892054 (US 1428);
ITS = MG892055 (US 1585).
Notes: Clavulina thindii is distinct from all other species
of Clavulina by the ITS sequence data and combination of
morphological features: violet to lilac profusely branched
basidiomes (42–66 9 38–49 mm) with white to greyish
white stipe and rounded-acuminate to irregularly coronate
or blunt apices, subsolid to hollow context, subglobose to
broadly ellipsoid basidiospores, cylindrical to subclavate
basidia with two cornuted sterigmata, absence of cystidia,
noninflated to inflated tramal hyphae and presence of
clamp connections.
In field, Clavulina thindii may appear similar to C.
livida Shu Z. Yan et al. with both species possessing large
sized, branched basidiomes with rounded-acuminate apices
which turn brown at maturity, smooth basidiospores with
one large guttule, subcylindrical to clavate basidia with
sometimes post-partial septa, absence of cystidia, noninflated to inflated tramal hyphae and presence of clamp
connections. However, the latter has greyish basidiomes
with olive stipe, larger (11.6–12.9 9 10.7–12.5 lm), globose to subglobose basidiospores and larger basidia
(51.5–76.5 9 7–12.3 lm) (He et al. 2016).
Clavulina amethystina (Bull.) Donk shares similarities
with C. thindii in having lilac to violet, branched basidiomes with obtuse to blunt apices, but the former has
completely lilac to violet basidiomes (the stipe of Clavulina thindii is white to greyish white), becoming dark grey
with age and ellipsoid (Lm/Wm & 1.55–1.62) basidiospores (Corner 1950; Olariaga et al. 2009).
Clavulina cinerea (Bull.) J. Schröt, C. rugosa (Bull.) J.
Schröt, C. coralloides (L.) Schröt. (= C. cristata), C. subrugosa (Cleland) Corner and C. samuelsi R.H. Petersen are also
closely related to C. thindii. However, these five species show
morphological differences from C. thindii. Clavulina cinerea
differs from C. thindii by its white to dark grey basidiomes
with acute to blunt apices, indistinct or not well-delimited
stipe due to basitonic branching and ovoid (Lm/Wm =
1.1–1.3) basidiospores (Bulliard 1788; Olariaga et al. 2009).
Clavulina rugosa is distinguished by its simple to sparsely
branched, white basidiomes, noncristate and normally obtuse
apices and relatively larger basidiospores (9–14 9 8–12 lm,
Lm 9 Wm = 9.9–10.6 9 8.2–8.8) (Corner 1950; Olariaga
et al. 2009). Clavulina coralloides differs from C. thindii by
its white to grey basidiomes with not well-delimited stipe due
to basitonic branching, cristate apices, comparatively larger
and subglobose basidiospores (7–11 9 6.5–10 lm) (Holmskjold 1790; Corner 1950; Olariaga et al. 2009). Clavulina
subrugosa can be separated from C. thindii by its simple or
filiform to flattened branched, white to pale greyish brown
basidiomes
and
rather
large
basidiospores
(6.8–8.6 9 5.8–7.2 lm) (Corner 1950; Petersen 1988a).
123
Fungal Diversity
Fig. 146 Baorangia
rufomaculata microscopic
features. a Basidiospores.
b Basidia. c Cheilocystidia.
d Pleurocystidia.
e Caulocystidia. f Pileipellis.
g Stipitipellis. Scale bars
a = 10 lm, b–e = 20 lm, f,
g = 50 lm. All drawings were
made from Both 4144
Clavulina samuelsii differs from C. thindii in having tilleul
buff fruiting body, comparatively larger basidiospores
(7.2–9.4 9 6.1–7.6 lm) and presence of cystidia (Petersen
1988a).
123
Clavulina mussooriensis Corner et al. (an Indian species) shares similarities with C. thindii in possessing profusely branched basidiomes and sometimes subglobose
basidiospores but the former has cream-white to greyish
brown basidiomes with cristate to acute apices and larger
Fungal Diversity
Fig. 147 Lanmaoa pallidorosea microscopic features. a Cheilocystidia. b Pleurocystidia. c Two types of caulocystidia. d Pileipellis.
e Stipitipellis. Scale bars a–c = 20 lm, d, e = 50 lm. All drawings were made from Both 4432
basidiospores (7–9 9 5.6–8 lm) (Corner et al. 1958).
Clavulina limosa K.S. Thind & Sharda (also originally
reported from India) differs from C. thindii by simple
(unbranched), slimy, creamish white to pallid-cream
basidiomes and globose to subglobose basidiospores
(Thind and Sharda 1984).
Thind (1961) reported Clavulina amethystinoides (Peck)
Corner from India, but the conspecificity has not yet been
verified with help of sequence data from the Indian collections. Clavulina amethystinoides shares some similarities with C. thindii in possessing violet basidiomes and
2-spored basidia, but differs in having simple or sparsely
123
Fungal Diversity
branched basidiomes, comparatively larger and globose to
subglobose basidiospores (9–12.3 9 8.8–10.5 lm). Moreover, C. amethystinoides (originally reported from USA)
differs from C. thindii by having simple or irregular
branches, slightly palmate branching, pale drab, pinkish
tan, fawn or very pale lilac basidiomes, pale livid flesh,
eventually blackening tips with distinctly different colour
and larger basidiospores (7–10 9 6–8 lm) (Corner 1950).
Clavulina thindii is commonly collected and consumed
by local villagers. ‘Angalchyun’ (fingers like mushroom) is
the vernacular name used for this species in the local language (Garhwali). However, inhabitants refer this vernacular name (Angalchyun) for all clavarioid fungi. The ITS
dataset consisting of 49 sequences (including our isolates:
US 1428, represented by GenBank no. MG892054 and US
1585, represented by GenBank no. MG892055) of Clavulina and Hydnum L. was analyzed by maximum likelihood
phylogenetic inference (Fig. 148). Two sequences isolated
from Indian materials appear to be nested amongst other
sequences of Clavulina and form a distinct clade being
sister to Chinese collections of C. livida (represented by
KU219603, KU219604, KU219605 and KU219606). The
combination of morphological features and phylogenetic
analysis indicates that Clavulina thindii is a new species,
distinct from all the known taxa of Clavulina.
Polyporales Gäum.
Phanerochaetaceae Jülich
The family Phanerochaetaceae was described by Jülich
(1981) with Phanerochaete as generic type, and also
including the genera Meruliopsis Bondartsev, Phlebiopsis
Jülich and Scopuloides (Massee) Höhn. & Litsch. This
family was characterized by mostly resupinate basidiome,
thick-walled hyphae, often without clamps, basidia narrowly clavate and spores hyaline, smooth and thin-walled
(Larsson 2007; Ryvarden and Melo 2014; Yuan et al.
2017). Justo et al. (2017) revised the family on the basis of
molecular and morphological data and, in addition to
Phanerochaete, they include the following thirteen genera:
Bjerkandera P. Karst., Donkia Pilát, Hapalopilus P. Karst.,
Hyphodermella J. Erikss. & Ryvarden, Oxychaete Miettinen, Phaeophlebiopsis Floudas & Hibbett, Phanerina
Miettinen, Phlebiopsis Jülich, Pirex Hjortstam & Ryvarden, Porostereum Pilát, Rhizochaete Gresl. et al., Riopa
D.A. Reid and Terana Adans. Most of these genera are
corticioid and rarely polyporoid, with the hyphal system
monomitic or sometimes dimitic, and most have hyphae
without clamps; cystidia are often present and spores are
thin-walled, hyaline and smooth. All species of this family
produce a white-rot.
Phanerochaete P. Karst.
123
Phanerochaete is a genus of widespread corticioid fungi
that causes white rot on all kinds of decayed wood and
plays a key role in forest ecosystems. The genus contains
167 legitimate names (MycoBank 2019). Since 2008 eight
new species and one subspecies have been described
(Nakasone 2008; Ghobad-Nejhad et al. 2015; Floudas and
Hibbett 2015; Liu and He 2016; Sádlı́ková and Kout 2017;
Spirin et al. 2017), and six new combinations have been
proposed (Nakasone 2008; Hjortstam et al. 2009; Melo
et al. 2012; Floudas and Hibbett 2015; Volobuev et al.
2015; Miettinen et al. 2016).
Phanerochaete was described by Karsten (1889) to
accommodate Thelephora alnea Fr. and T. odora Fr., and
therefore based on two elements, without designation of a
type. Cooke (1953) considered T. alnea as generic type and
later Corticium decolorans (= Phanerochaete velutina)
was chosen as lectotype by Eriksson et al. (1978a).
Recently, Spirin et al. (2017) reinstated T. alnea as the
generic type. Long neglected, Phanerochaete was reintroduced by Donk (1957) with the two initial species. Several
years later, the taxonomy of the genus was discussed by
Donk (1962), Parmasto (1968), Eriksson et al. (1978b) and
Burdsall (1985). Donk (1962) amended Phanerochaete,
recognizing that the generic limits were uncertain and
suggesting that other taxa should be included. Parmasto
(1968) accepted twenty species separated in two subgenera,
Phanerochaete and Phanericium Parm.; cystidiate and
acystidiate species, respectively. Eriksson et al. (1978b)
recognized twelve species in Northern Europe divided into
three groups, Phanerochaete velutina group (= subgen.
Phanerochaete), P. tuberculata group (= subgen. Phanericium Parm.), and P. septocystidiata group, the last one
with only one species characterized by cystidia regularly
septate. Burdsall (1985) recognized 42 species in three
subgenera, Phanerochaete, Phanericium, and Scopuloides
(Massee) Burdsall, the latter with cystidia and subiculum
usually distinct.
Phylogenetic studies carried out in Phanerochaete sensu
Burdsall (1985) showed that the genus is polyphyletic (De
Koker et al. 2003; Wu et al. 2010a) and its species are
distributed in nine phylogenetic linages across the phlebioid clade within order Polyporales (Floudas and Hibbett
2015). According to Floudas and Hibbett (2015) one of
these linages, Phanerochaete sensu stricto, includes species mostly found in the subgenus Phanerochaete sensu
Burdsall (1985).
From a morphological point of view, the species included in the Phanerochaete sensu stricto clade in Floudas
and Hibbett (2015) and in our LSU analysis (Figs. 151,
152) are characterized by their basidiomes being resupinate, membraceous, white, yellowish orange, or red to
brown; the subiculum well-developed and the margin
mostly fibrillose, with or without cordons. The hyphal
Fungal Diversity
Fig. 148 Maximum likelihood phylogeny of Clavulina showing the
position of novel species Clavulina thindii (shown in blue and bold).
The ITS sequence dataset was aligned with online version of MAFFT
v. 7 (Katoh and Standley 2013) and no manual editing was done
within the alignment. One-thousand bootstrap replicates were
analyzed to obtain nodal support values. Analysis was conducted
with MEGA 6.0 (Tamura et al. 2013). Hydnum rufescens and H.
repandum were used as the outgroup taxa. Maximum likelihood
bootstrap values greater than 50% are indicated at the nodes. The bar
indicates the number of nucleotide substitutions per site. Ex-type
strains are indicated in bold
system is monomitic; the subicular hyphae thin to thickwalled, loosely interwoven, simple septate or with scattered single to multiple clamps; the subhymenial hyphae
thin-walled, without clamps. The cystidia are cylindrical to
subulate, thin-walled to almost metuloid, without a basal
123
Fungal Diversity
Fig. 149 Clavulina thindii (CAL 1661, holotype). a Fresh basidiomes. b Apices of basidiome. c Transverse section through
hymenium. d Basidium with post-partial septa and basal clamp
123
(indicated by white arrow). e Hymenophoral tramal hyphae showing
clamps (indicated by white arrows). f Basidiospores. Scale bars
a = 20 mm, c–f = 10 lm
Fungal Diversity
Fig. 150 Clavulina thindii (from holotype). a Basidia. b Basidiospores. c Hymenophoral tramal hyphae showing clamps. Scale bars
a–c = 10 lm
clamp. Spores are thin-walled, inamyloid and non
dextrinoid.
We follow the treatment and updated accounts of
Floudas and Hibbett (2015) and Miettinen et al. (2016); the
new species, Phanerochaete australosanguinea is
introduced.
Phanerochaete australosanguinea Telleria, M. Dueñas &
M.P. Martı́n, sp. nov.
MycoBank
number:
MB825276;
Facesoffungi:
FoF05725, Figs. 153, 154
Etymology: From Latin ‘‘australis’’ which means
southern and ‘‘sanguinea’’ refers to the morphological
similarity of this species with Phanerochaete sanguinea
(Fr.) Pouzar
Holotype: 20102Tell, MA-Fungi 91309
Colour codes follow: ISCC-NBS Centroid Colour Charts
(Kelly and Judd 1976)
Basidiomes resupinate, orbicular to confluent, membranaceous, closely adnate; hymenophore smooth to slightly
cracked in mature specimens; brownish orange (54. Br O–37.
m r O) when young, in mature specimens orange yellow (70.
l. OY–71. m. OY); the edge especially differentiated by dark
reddish orange (38. d. r O). Margin fibrillose and white; with
well-developed dark strands sometimes present. KOH 3%
reaction (?) turning green that quickly change to brown.
Subiculum byssoid, white to cream, lighter in colour than the
hymenophore. Hyphal system monomitic. Subicular hyphae
thick-walled, loosely interwoven, sparsely branched, growing parallel to the substrate, (5–)8–10 lm wide, sometimes
with orange brown content, with single clamps. Subhymenial
hyphae thin-walled and short-celled, branched, densely
interwoven, growing perpendicular to the substrate, 3–5 lm
wide. Cystidia thin-walled, cylindrical to tapering, sometimes with secondary septa, 44–66 9 4–5 lm. Basidia narrowly cylindrical to claviform, four sterigmata,
40–46 9 4–5 lm. Spores ellipsoid, 4.5–5.5 9 3–3.5 lm,
thin-walled, sometimes with oil drops in the protoplasm, Q
(L/W) = 1.29.
Material examined: CHILE, Los Lagos, Palena, Hornopirén, comuna Hualainhué, Huinay Biological Reserve,
path to Cerro del Tambor, 42°220 5400 S 72°240 5300 W, 202
msl, on unidentified wood, 8 May 2013, M. Dueñas, M.P.
Martı́n & M.T. Telleria, 20102Tell., (MA-Fungi 91309,
holotype); ibid., 20098Tell. (MA-Fungi 91308);
20114Tell. (MA-Fungi 91310).
Host and habitat: Known from the Valdivian temperate
rain forest, Chilean Patagonia, South America, growing on
decayed wood and associated with white rot.
GenBank numbers: ITS = MH233925, LSU = MH233928
(20098Tell., MA-Fungi 91308); ITS = MH233926, LSU =
MH233929, (20102Tell., MA-Fungi 91309, holotype);
ITS = MH233927 (20114Tell., MA-Fungi 91310).
Notes: Based on LSU analyses (Fig. 151), Phanerochaete australosanguinea clusters into the Phanerochaete sensu stricto clade in the P. sanguinea complex,
according to the maximum likelihood phylogeny of the
concatenated ITS, LSU, RPB1 and RPB2 obtained by
Floudas and Hibbett (2015). Most species of this complex
cause red–orange staining of the substrate, such as P.
australosanguinea (Fig. 153c). In the LSU analyses, P.
australosanguinea is a sister species of P. sanguineocarnosa Floudas & Hibbett. Morphologically, this is corroborated by both species sharing well-developed dark
strands, the KOH reaction turning quickly green changing
to brown on the basidiome; the subicular hyphae having
developed clamps, and cystidia in both species are cylindrical, tapering to the apex, with secondary septa and of
similar size. However, the spores are ellipsoid to broadly
ellipsoid in P. australosanguinea (Q = 1.29), while those
of P. sanguineocarnosa, described by Floudas and Hibbett
(2015), are sub-ellipsoid to ellipsoid (Q = 1.8).
In the ITS analyses (Fig. 152), Phanerochaete australosanguinea also clusters in the P. sanguinea complex.
However, the newly generated sequences are very distinct
123
Fungal Diversity
from those of the P. sanguinea complex of Floudas and
Hibbett (2015). These authors included mainly species and
specimens from the Northern hemisphere (except one
sequence from Tasmania). The three collections from Chile
form a basal and highly supported group (99% ML, 97%
MP and 1.00 PP) indicating that P. australosanguinea is
distinct from other species of the P. sanguinea complex by
its geographical distribution. So far, P. australosanguinea
has been reported from Valdivian temperate rain forest in
Chilean Patagonia, where it is probably frequent and could
have been confused with P. sanguinea (Fr.) Pouzar
reported from the Patagonian Andes forest of Southern
Argentina by Greslebin and Rajchenberg (2003).
Russulales Kreisel ex P.M. Kirk et al.
Russulaceae Lotsy
Russulaceae is one of the most speciose, and morphologically diverse ectomycorrhizal mushroom families
(Miller et al. 2006; Hyde et al. 2016). This family has a
worldwide distribution of more than 1200 species (Kirk
et al. 2008). Multigene molecular phylogenetic analyses
consistently recovered this family as a monophyletic clade
which is now comprised of four genera: Russula Pers.,
Lactarius Pers., Lactifluus (Pers.) Roussel and Multifurca
Buyck & Hofstetter (Buyck et al. 2008, 2010). We follow
the treatments and the updated accounts of taxa in Russulaceae in Li et al. (2016), Hyde et al. (2016, 2017), De
Crop et al. (2017) and Tibpromma et al. (2017). Four novel
species viz. Lactarius olivaceopallidus, Lactifluus midnapurensis, Russula choptae and R. uttarakhandia are introduced in Russulaceae based on morphological
distinctiveness and phylogenetic support. An updated
phylogenetic tree of Lactarius, Lactifluus, Russula subg.
Russula and subg. Incrustatula are provided. Detailed
discussion and reviewed literatures were provided in Li
et al. (2016), Hyde et al. (2016, 2017), De Crop et al.
(2017) and Tibpromma et al. (2017).
Lactarius Pers.
Lactarius is one of the most prevalent agarics which
form obligate symbiotic associations with many wide
spread genera of trees (Verbeken et al. 2001; Le et al. 2007;
Das and Verbeken 2011; Verbeken and Nuytinck 2013;
Das et al. 2015; Hyde et al. 2016; Uniyal et al. 2017).
Multigene based phylogenetic studies resulted segregation
of paraphyletic Lactarius sensu lato into Lactifluus Pers.
(Roussel) and Lactarius (including all angiocarpic species).
Lactarius sensu novo consists of three cosmopolitan subgenera: Lactarius subg. Lactarius, L. subg. Plinthogalus
(Burl.) Hesler & A. H. Sm. and L. subg. Russularia (Fr. ex
Burl.) Kauffman (Verbeken and Nuytinck 2013; Das et al.
2015). Recent studies indicate the high species richness
within L. subg. Lactarius in the Indian Himalayan region
123
(Sharma and Das 2003; Das and Sharma 2005; Das and
Verbeken 2011; Das 2013; Das et al. 2015; Uniyal et al.
2017). A novel species within L. subg. Lactarius
sect. Uvidi, collected from Himalayan temperate forest
(India) is presented here with morphological details and
ITS phylogeny (Fig. 155).
Lactarius olivaceopallidus Uniyal, sp. nov.
MycoBank number: MB820694; Facesoffungi number:
FoF04839, Figs. 156, 157
Etymology: Due to pale olive colour of basidiomes.
Holotype: CAL 1401
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Pileus 21–88 mm diam., convex, centrally depressed,
finally infundibuliform, olive yellow (3C6) at centre,
greyish yellow (4C4–7) toward margin, paler at margin,
surface slimy to gelatinous, viscid at centre, sticky when
dry, azonate to faintly zonate near margin, pale yellow
(3A3) then yellow (3A7) with 30% KOH; margin decurved
to inrolled. Lamellae yellowish white (1A2), pale yellow
(3A2) in age, often with greyish yellow (4B4) spots,
greyish violet (15D5) on bruising, adnate to broadly
adnate, close (10–17 L ? l/cm), including lamellulae),
some forked near stipe, lamellulae numerous. Stipe
21–45 9 10–23 mm, yellowish white (4A2) to pale yellow
(4A3), spotted with light yellow (4A5) scrobiculi, stuffed
to hollow. Context up to 14 mm thick in pileus, yellowish
white (1A2), pale yellow (3A3) with KOH, pale yellow
(3A2) in stipe, immediately turning violet (15C6) on
exposure. Latex yellowish white (3A2), rather abundant,
immediately turning dull lilac (15C3) on gills, context,
white paper and after 2–3 minutes on glass slide. Odour
pleasant. Taste acrid. Basidia 51–72 9 9.5–16 lm, subclavate, 2–4-spored, sterigmata 5–9 lm long. Basidiospores 7.5–8.7–9.5 9 7–7.7–8.5 lm, (n = 40, Q = 1.06–
1.12–1.21), subglobose to broadly ellipsoid; ornamentations amyloid, 1.7–2 lm high, mostly composed of broad
ridges with wavy to even margin, triangular narrow ridges
with round apices and small isolated warts, forming a dense
incomplete reticulum; plage distally amyloid. Pleuromacrocystidia 51–113 9 9–17.5 lm, abundant, subclavate
to fusiform, mostly round to blunt and acute apices,
sometimes mucronate, contents dense. Pleuropseudocystidia up to 9 lm thick, nonemergent, cylindrical with
rounded nonforked apex. Lamellae edge sterile composed
of cheilomacrocystidia and marginal cells. Cheilomacrocystidia 34–63 9 8.5–12 lm, broadly fusiform, round to
blunt apex. Marginal cells 13–23 9 4–6 lm, cylindric to
subfusiform with rounded apex. Hymenophoral trama
mainly composed of sphaerocytes and abundant lactifers,
lactifers up to 14 lm wide. Pileipellis an ixocutis,
300–630 lm thick, composed of densely packed septate
Fungal Diversity
Fig. 151 Phylogram generated from Bayesian inference analysis of
LSU sequence data of species of Phanerochaete sensu stricto from
Floudas and Hibbett (2015) and the new species is marked in grey
square. Maximum parsimony and maximum likelihood bootstrap
support C 50% (10.000 replicates both), and Bayesian posterior
probabilities C 0.50 BYPP are shown above the nodes. The tree is
rooted with Pirex concentricus and Hyphodermella rosae. Newly
generated sequences are in blue, types are in bold
hyphae, often shriveled, forming bundles in outer parts,
1–4 lm thick. Lactifers in pilear trama up to 11 lm wide.
Stipitipellis an ixocutis, 65–102 lm thick, hyphae 1–4 lm
wide, septate, not shriveled.
Material examined: INDIA, Uttarakhand, Pauri Garhwal, Chaubatta, alt 1904 m, N30° 09.6810 E78° 51.2220 , 19
July 2016, P. Uniyal, PU-1110 (CAL 1401, holotype);
ibid., Pauri Garhwal, Chaubatta, 16 July 2016, P. Uniyal,
PU 15-1090.
Host and habitat: Gregarious, under Quercus leucotrichophora A. Camus in temperate mixed forest.
GenBank numbers: ITS = KY440363 (CAL1401),
ITS = MF741705 (PU 15-1090).
Notes: Lactarius olivaceopallidus differs from other
members of Lactarius sect. Uvidi due to presence of pale
olive basidiomes, abundant yellowish white latex, very
high (B 2 lm) ornamentations of basidiospores and very
thick (B 630 lm) ixocutis type of pileipellis. The combination of characteristics like viscid to slimy, azonate to
slightly zonate pileus, ixocutis pattern of pileipellis and
presence of white latex turning dull lilac on exposure
undoubtedly place the present taxon in Lactarius
sect. Uvidi.
In the field, Lactarius olivaceopallidus can easily be
recognized by its pale olive basidiome, slimy to gelatinous
pilear surface, pale yellow lamellae turning lilac when
damaged, scrobiculate stipe surface and yellowish white
latex that immediately changes into lilac on gills. Microscopically, very thick pileipellis (B 630 lm) and basidiospores with very high prominences (up to 2 lm) in form
of a dense incomplete reticulum are also distinct.
Studies have suggested that poor molecular resolution in
Lactarius sect. Uvidi has been a problem, as species in this
group are dissimilar but genetically very close (Das et al.
2015; Barge et al. 2016). Species of L. sect. Uvidi in present study are well-separated (Fig. 155) from species
belonging to other sections of L. subg. Lactarius. Two
collections of the present species L. olivaceopallidus
(CAL1401 and PU 15-1090) are distinctly separated in a
relatively long branch from its closely related species.
Morphologically, L. olivaceopallidus can be confused with
L. uvidus (Fr.) Fr. (GenBank no. KJ742416, KJ742417;
Fig. 155), L. violascens (J. Otto) Fr. and L. aspideus (Fr.)
Fr. But, L. uvidus (originally described from Europe) differs in having greyish pink to light vinaceous grey pileus,
cream to pale pinkish buff lamellae, larger basidiospores
123
Fungal Diversity
Fig. 152 Phylogram generated from Bayesian inference analysis of
ITS sequence data of species of Phanerochaete sanguinea complex
from Floudas and Hibbett (2015) and the new species is marked in
grey square. Maximum parsimony and maximum likelihood bootstrap
support C 50% (10,000 replicates both), Bayesian posterior probabilities C 0.50 BYPP are shown above the nodes. The tree is rooted
with three Phanerochaete species
(8.4–11.3 9 6.6–8.5 lm), with shorter ornamentations (up
to 0.7 lm) in form of rounded warts and obtuse broad
ridges forming an incomplete reticulum. Lactarius violascens (originally reported from Europe) shares similarity in
occurrence under deciduous trees but quite different due to
greyish buff to brownish violet, azonate or zonate pileus,
extracellular pigmentation in pileipellis appearing as dark
brown granules and larger basidiospores (7.6 –
11.3 9 6.4–8.5 lm) (Heilmann-Clausen et al. 1998). Like
L. olivaceopallidus, a pale basidiome is also present in L.
aspideus (originally reported from Europe) which differs
from L. olivaceopallidus due to smaller (10–70 mm in
diam.) pileus, non-scrobiculate stipe, unchanging latex on
isolation from flesh, shorter ornamentations (up to 0.5 lm)
in basidiospores, zebroid pattern forming complete to
nearly complete reticulum and very thin (40–70 lm)
pileipellis (Heilmann-Clausen et al. 1998).
Lactarius salicis-reticulatae Kühner (GenBank no.
KR090958, KR090959; Fig. 155) is another lilac staining
species which is also phylogenetically close (94% similarity for 93–96% query coverage using BLAST) to the
present novel species, however, it has subdistant to distant
lamellae, mild tasting watery white latex, association with
Salix and larger basidiospores (8.5–11.5 9 8–10 lm)
(Barge and Cripps 2016).
Genetically, Lactarius repraesentaneus Britzelm.
(GenBank no. KR090950, KR090954, KR090948,
KR090953, JF908312 and KR090952; Fig. 155) is the
closest relative (only 95–96% similarity for 93–96% query
coverage using BLAST) to L. olivaceopallidus, but morphology of the two species is very different. Lactarius
repraesentaneus (originally reported from Europe) has
orange-yellow to honey pileus and conspicuously bearded
pileus margin (hairs up to 5 mm long). Moreover, L.
repraesentaneus is reported growing in ecological
123
Fungal Diversity
Fig. 153 Phanerochaete australosanguinea (20102Tell., MA-Fungi
91309, holotype). a Basidiome dry specimen. b Strands, dry
specimen. c Substrate stained red. d Basal hyphae thick-walled with
clamp. e Probasidia and cystidium. f Basidium stained in congo red.
g Cystidium with secondary septa stained in congo red. h Spores.
Scale bars a–c = 1 cm, d–h = 10 lm
123
Fungal Diversity
Fig. 154 Phanerochaete
australosanguinea (20102Tell.,
MA-Fungi 91309, holotype).
a Basal hyphae thin- to thickwalled with clamps.
b Subhymenial hyphae.
c Hymenium with probasidia,
basidia and cystidia. d Spores.
Scale bars a–d = 10 lm
association with Picea, Betula and Salix in temperate to
arctic-alpine areas of North America and Europe. Microscopically, it has longer basidiospores (8–10.5 lm) with
shorter (B 1 lm high) ridges and ixotrichoderm type of
pileipellis. Phylogenetic analyses have suggested 99.4%
similarity of L. repraesentaneus with L. dryadophilus
Kühner (GenBank no. KR090901), which is very similar to
L. repraesentaneus (Barge et al. 2016). Lactarius dryadophilus (originally reported from Europe) is also genetically
close to L. olivaceopallidus (95% similarity for 94% query
coverage using BLAST) but quite different due to white to
pale yellow pileus with bearded margin and larger basidiospores (9.3–11.8 9 7.2–9.2 lm) having very low
(B 0.3 lm high) ornamentations (Heilmann-Clausen et al.
1998; Barge et al. 2016).
123
In Asia, Lactarius sect. Uvidi is represented by L. formosus H.T. Le & A.Verbeken and L. pyriodorus K. Das &
Verbeken. Lactarius formosus (originally reported from
Thailand) is distinct from the present taxon by possessing
greyish yellow to yellowish buff, zonate, scaly pileus with
bundles of glutinous hairs and cutis to trichoderm type of
thick (120–140 lm) pileipellis (Le et al. 2007). Lactarius
pyriodorus K. Das & Verbeken (originally reported from
Indian Himalaya) cannot be mistaken in the field for L.
olivaceopallidus as the former displays brown vinaceous
pileus, watery white latex, distinctive pear-like fruity odour
and association with Abies. Microscopically, L. pyriodorus
has larger basidiospores (8–8.9–10.5 9 6.9–7.5–8.8 lm),
with lower (B 1.1 lm high) prominences and thinner
(B 320 lm) pileipellis (Das et al. 2015).
Fungal Diversity
Lactifluus (Pers.) Roussel
We follow the latest treatment and updated account of
Lactifluus in De Crop et al. (2017) and Hyde et al. (2017). The
new species Lactifluus midnapurensis is introduced in Lactifluus subgen. Lactifluus (Burl.) Hesler & A.H. Sm (Fig. 158).
Lactifluus midnapurensis S. Paloi & K. Acharya, sp. nov.
MycoBank number: MB820587; Facesoffungi number:
FoF03248, Figs. 159, 160
Etymology: The specific epithet refers to the type
locality, Purba Midnapur (India).
Holotype: CAL 1516
Colour codes follow: Methuen handbook of colour
(Kornerup and Wanscher 1978).
Pileus 11–48 mm diam., convex to broadly convex with
an umbo when young, becoming plain to infundibuliform
or applanate with a slight umbo toward centre at maturity,
semi-moist to dry when young, dry and shiny at maturity,
smooth, becoming slightly wavy and translucent with
maturity, margin incurved at young stage, brown (5E4;
6E4) to yellowish brown (5E5) toward centre, brownish
grey (5C2; 6D2) at early stage, becoming finally white
(1A1) toward margin at maturity, no colour change on
bruising, turns orange to reddish orange with guaiacol,
negative in phenol, SV and KOH; context 1 mm toward
centre, \ 1 mm toward margin, white (1A1), no colour
change on exposed in air. Partial veil present at the young
stage, white (1A1), but disappear when mature. Lamellae
ca. 3 mm broad, decurrent, distant, white (1A1), negative
in phenol and KOH; edge even, regular, concolorous;
lamellae two to three of length, forked toward margin.
Stipe 8–43 9 2–5 mm, central when young, becoming
central or excentric when mature, semi-moist at early stage,
becoming dry and shiny when mature, smooth, grey (4C1)
to golden grey (4C2) or brownish grey (4D2) when young,
becoming white (1A1) at maturity, turns light yellowish
brown on bruising at early stage but no change when
mature, orange to orange-red with guaiacol, negative with
KOH, phenol and SV; context hollow, white (1A1), no
colour change on exposed in air. Latex water-like (transparent), no colour change up to 5 minutes in exposed on
air. Odour unknown. Taste mild. Spore print white.
Lamellar trama mainly composed of irregularly arranged
sphaerocytes, hyphae and lactiflus hyphae. Basidia (43–
)46.5–51.5–53.7(–57.2) 9 (7.1–)8–8.9–10(–11) lm, clavate to subclavate, hyaline, thin-walled, oil droplets present
when
viewed
KOH,
4-spored;
sterigmata
3.6–7.2 9 1–2.7 lm.
Basidiospores
5.5–6.5–7.1(–
8) 9 (5–)6–6.6–6.9(–7.5) lm, (n = 30) Q = 1.02–1.07–
1.12, globose to subglobose, ornamentation amyloid,
composed of short (0.3–0.6 lm) and long warts
(1.5–2.0 lm), interconnected by a ridge line, forming a
complete reticulum, sometimes connected line are shown
wing-like pattern in side view, suprahilar spot amyloid or
not. Lamellae edge sterile mainly composed of numerous
cheilocystidia. Cheilocystidia 27–48.5 9 4.5–7.5 lm,
subclavate to subcylindrical with mostly capitate apex,
hyaline, thin-walled, oil droplet present when viewed with
KOH. Pleuromacrocystidia 43–61 9 7.5–11 lm, rare
toward cap margin but abundant toward cap centre, mostly
appendiculate or mucronate, hyaline, thin-walled. Pleuropseudocystidia 2–4.5 lm in diam., abundant, cylindrical
to subcylindrical with capitate or obtuse apex, dense with
cytoplasmic component. Pileipellis a palisade, context
made up with unequal, irregular shaped sphaerocytes and
loosely arranged hyphal cells, branched, septet; distantly
2-layers, 30.6–46.2 lm deep, subpellis non-gelatinous,
composed of tightly arranged globose to subglobose or
irregular shaped cell, 11.6–19.7 9 10.7–17.9 lm, thinwalled, 2–4-celled deep; suprapellis composed of erect to
sub-erect hyphal cell, 14.3–39.4 9 3.6–7.1 lm, hyaline,
thin-walled, 2–3-celled, underling attached with the subpellis region of globose cells, non-gelatinized. Stipitipellis
a palisade, 52.2–85.6 lm deep, upper layer composed of
thin-walled, septet (up to 4 times) hyphae, 4.5–7.5 lm in
diam., hyphal end subulate or obtuse, underling 2–3 globose like cells.
Material examined: INDIA, West Bengal, Purba Midnapur, Kasafaltalya, 21°430 0.5520 ’ N, 87°300 26.647200 E, S.
Paloi & K. Acharya, 23 August 2015, SOUMITRA-22 (CAL
1516, holotype); ibid., 21°430 6.2220 ’N, 87°300 7.95240 ’E, S.
Paloi & A.K. Dutta, 27 August 2015, SOUMITRA-40 (CAL
1523).
Habit and habitat: Common, solitary or gregarious,
growing under Shorea robusta C.F. Gaertn. tree
(Dipterocarpaceae).
GenBank numbers: ITS = KY785175, LSU = KY785177
(CAL 1516, holotype); ITS = KY785176, LSU =
KY785178 (CAL 1523).
Notes: The diagnostic features of the newly described
species includes small to medium sized (11–48 mm in
diam.), convex to infundibuliform or applanate pileus,
brownish grey to white that turns reddish orange with
guaiacol; distant lamellae, white; slightly excentric stipe;
watery latex (transparent); white spore print; mild taste;
globose to subglobose basidiospores with a mean size of
7.1 9 6.6 lm, Qm = 1.07, ornamentation composed of
warts (up to 2 lm), interconnected with each other forming
complete reticulate; pileipellis a palisade type. This combination of characteristics undoubtedly place Lactifluus
midnapurensis under subg. Lactifluus sect. Gerardii
(Stubbe et al. 2010; De Crop et al. 2017).
Lactifluus midnapurensis appears to be related to L.
coniculus (Stubbe & Verbeken) Verbeken, described from
Sri Lanka growing in association with Shorea sp. However,
L. coniculus differs by cigar brown to snuff brown pileus,
123
Fungal Diversity
123
Fungal Diversity
b Fig. 155 Phylogeny of Lactarius olivaceopallidus inferred from
maximum likelihood analysis of ITS sequence dataset using
RAxMLGUI 1.5 (Silvestro and Michalak 2012). Multiple sequence
alignment was performed using MAFFT v.7 (Katoh and Standley
2013) with default settings. Alignment was manually edited with
Bioedit v 7.2.5 (Hall 1999). To change the multiple alignment format,
Alignment Transformation Environment (ALTER) was used (GlezPeña et al. 2010). One-thousand bootstrap replicates were analyzed to
obtain nodal support values. TreeGraph2 was used for visualization of
the phylogram (Stöver and Müller 2010). The novel species is shown
in blue and type strains are in bold. Species of sister genus Lactifluus
were selected as the outgroup taxa
solid stipe context, comparatively larger basidiospores
(7.1–10.2 9 6.4–8.8 lm) with ornamentation like interconnected conical to triangular, round to acute warts and
absence of pleuromacrocystidia (Stubbe et al. 2012b).
Within the same section, among taxa with similar sized
basidiospores, colouration of the latex and growth in
association to Dipterocarpus plants, Lactifluus limbatus
(Stubbe & Verbeken) Stubbe (described from Malaysia) is
a phylogenetically close taxon (Fig. 158). Lactifluus limbatus has a larger pileus (30–60 mm), brownish grey
towards centre and grey towards margin, greyish brown
lamellar edge, spore ornamentation up to 0.8 lm high and
absence of pleuromacrocystidia (Stubbe et al. 2012a, b).
Lactarius atrovelutinus J.Z. Ying frequently encountered
from China and Malaysia, has a dark black-brown pileus,
cream (young) to yellow (mature) lamellae, latex that turns
pink or brownish pink on drying, presence of fruity or
sweet odour, absence of pleuromacrocystidia and very
thick pileipellis (80–100 lm; Ying 1991). Lactarius conchatulus Stubbe & Verbeken, described from northern
Thailand (alt. 1300 m), has smaller pileus (2–7 mm diam.)
that turns pale yellow on bruising, subdecurrent lamellae,
and broadly ellipsoid basidiospores (Q = 1.08–1.35) with
up to 0.9 lm long ornamentation (Stubbe et al. 2012a, b).
Being a well representative member of the sect. Gerardii; Lactifluus bicolor (Massee) Verbeken, distributed in
Malaysia and Singapore, differs from L. midnapurensis in
having a larger pileus (20–80 mm diam.), dark brown or
fuliginous, longer stipe (10–80 mm), and larger basidiospores (7.5–9.6 9 6.4–8.6 lm) with a ornamentation up
to 0.3–0.5 lm high (Massee 1914; Stubbe et al. 2012a, b).
A taxon with similar pileus colouration, L. leae Stubbe &
Verbeken differs by its very larger pileus (35–120 mm in
diam.), somewhat differently sized basidiospores
(6.6–12.1 9 5.9–9.1 lm), ornamentation up to 0.5–1 lm
high and absence of macrocystidia (Stubbe et al. 2012a, b).
Russula Pers.
Russula is species-rich and one of the most thoroughly
monographed genera of fungi that is mainly ectomycorrhizal with a diverse range of plants in deciduous or
evergreen, broadleaf or coniferous woods, scrubland and
even meadows from tropical to subalpine areas and has a
cosmopolitan distribution (Knudsen and Borgen 1982;
Singer 1986; Buyck et al. 1996; Rawla 2001; Das and
Sharma 2005; Das et al. 2006a, b, Das et al. 2010, Das et al.
2013, Das et al. 2014, 2017a, b; Miller et al. 2012; Li et al.
2013, 2015; Hyde et al. 2016; Ghosh et al. 2016, 2017;
Ghosh and Das 2017). Species within this genus are known
by the combination of their conspicuous and fleshy basidiomes, colourful fragile pileus, amyloid warty basidiospores, abundant sphaerocytes in a heteromerous trama
that can make these fungi brittle, absence of latex and the
hyphae that lack clamp connections (Romagnesi 1967;
Singer 1986; Sarnari 1998, 2005). Russula has 10 subgenera namely: R. subg. Compactae, R. subg. Heterophyllidia, R. subg. Ingratula, R. subg. Amoenula, R. subg.
Incrustatula and R. subg. Russula, R. subg. Archaea,
R. subg. Brevipes, R. subg. Malodora and R. subg. Crassotunicata Buyck & V. Hofstetter (Sarnari 1998; Hongsanan et al. 2015; Das et al. 2017b). Here, two novel species
(belonging to subg. Incrustatula and Russula) collected
from the northwestern part of Indian Himalaya are
described with morphological details and phylogenetic
placement (Figs. 161, 162).
Russula choptae A. Ghosh & K. Das, sp. nov.
MycoBank number: MB824343; Facesoffungi number:
FoF04953, Figs. 163, 164
Etymology: Referring to Chopta in Uttarakhand (India),
the type locality.
Holotype: CAL 1658
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Basidiomes 34–112 mm in height, small to medium sized.
Pileus 6–48 mm in diam., hemispheric when young, convex,
plano-convex, expanding to applanate when mature, centre
slightly depressed with age, finely wrinkled and areolate at
maturity; margin decurved to plane with maturity, entire,
slightly tuberculately striate, dry (viscid when moist), peeling 1/4 from the edge, brownish red (9C6–8) at young stage,
pale red (9–10A3), pastel red (9–10A4–5) to madder red
(9A6–7), intermixed with pink, orange to yellowish tinge of
coral red (9B7) at the centre, pinkish white (9A2), pale red to
pastel red (9A3–4) towards the margin; pale yellow to pastel
yellow (2A3–4) with KOH. Pileus context greyish white
(1B1), 2–4 mm thick, unchanging after bruising, turning
reddish brown (8D6–7) with guaiacol. Lamellae adnexed to
subdecurrent, crowded (10–16/cm), chalky white to yellowish white (1A1–2) with maturity, up to 3 mm thick,
narrowing towards the margin, forked near the stipe apex,
middle of lamellae and near cap margin, entire, edges concolorous, unchanging when bruised. Lamellulae present in
different lengths (up to 5 series). Stipe 37–88 9 6–13 mm,
123
Fungal Diversity
Fig. 156 Lactarius olivaceopallidus (CAL 1401, holotype). a, c
Fresh basidiomes in field and basecamp. b Latex on gills. d Transverse
section through thick pileipellis. e Hyphal arrangement in pileipellis.
f, g Pleurocystidia. h Cheilocystidia and marginal cells. i Pseudocystidia. j, k SEM images of basidiospores. Scale bars a = 20 mm,
d = 100 lm, e, g–i = 10 lm, f = 25 lm, j–k = 2 lm
cylindrical to subclavate, central, dry, smooth, brittle, pale
red, pastel pink to pink rose (11A3–5) with white (1A1) to
yellowish white (1A2) areas or orange white to pale orange
(5A2–3) tinge at the base; turning reddish brown (8D6–7)
with guaiacol. Stipe context stuffed to hollow, dirty white,
unchanging after bruising but reddish brown (8D6–7) and
greyish green (27C4–5) with guaiacol and KOH respectively.
Taste acrid. Spore print yellowish white. Subhymenium layer
up to 28 lm thick, pseudoparenchymatous. Basidia
30–40 9 9–14 lm, 4-spored, cylindrical, subclavate to clavate, sterigmata up to 6 lm long. Basidiospores (6.9–)7–7.4–
7.9(–8) 9 (6–)6.1–6.6–7 lm, [n = 30, Q = (1–)1.06–1.13–
1.2(–1.31)], subglobose to broadly ellipsoid, rarely globose
and ellipsoid, composed of amyloid warts becoming linked
as small crests and thick ridges, forming an incomplete network, intermixed with isolated warts (up to 1 lm) in height;
123
Fungal Diversity
Fig. 157 Lactarius olivaceopallidus (CAL 1401, holotype). a Fresh/dissected basidiomes. b Cheilocystidia. c Basidiospores. d Pseudocystidia.
e Basidia. f Marginal cells. g Pileipellis. h Pleurocystidia. Scale bars a = 20 mm, b–h = 10 lm
suprahilar plage inamyloid, apiculi up to 2 lm high. Pleurocystidia (35–)45–64–80(–96) 9 (7–)7.5–10–12(–13) lm,
abundant, cylindrical to subclavate with capitate, appendiculate, pointed, mucronate, moniliform or rounded apex,
emergent up to 47 lm, completely filled with dense, finely
crystalline content in congo red, turning grey-black with
sulphovanillin (SV). Lamellar edge fertile with basidia and
cystidia. Cheilocystidia (30–)35–45.5–56(–61) 9 (6–)7–9–
11(–12) lm, cylindrical to subclavate with mostly rounded
apex, completely filled with dense, finely crystalline content
in congo red, turning grey-black with sulphovanillin (SV).
Pileipellis orthochromatic in cresyl blue, a trichoderm, up to
260 lm thick, two-layered and sharply delimited from the
underlying sphaerocytes of the context; distinctly divided in a
120–160 lm deep, suprapellis of erect to suberect hyphal
ends and pileocystidia, containing branched, septate, hyphal
endings, and underneath this suprapellis a 140–100 lm deep,
subpellis of interwined, more or less horizontally oriented
hyphae. Hyphal extremities near the cap margin composed of
a single or a few cells only; the terminal cells subcylindrical
to cylindrical, with rounded or obtuse tips, measuring (15–
)18–24–30(–34) 9 (2.5–)3–3.5–4(–5) lm; subterminal cells
123
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123
Fungal Diversity
b Fig. 158 Phylogram generated from maximum likelihood (- InL =
10343.610778) (RAxML) analysis using a GTR ? I ? G model of
nucleotide evolution based on ITS sequence dataset for 47 Lactifluus
and six outgroup sequences (Lactarius scrobiculatus, L. olympianus,
L. hatsudak, L. miniatescens, L. fuliginosus, L. tenellus). Maximum
likelihood bootstrap support values (left) greater than 50% and
Bayesian posterior probabilities (right) greater than 0.50 BYPP are
indicated above the nodes. Sequences used in this study mostly have
been sampled from a previous study (Stubbe et al. 2010, 2012a, b).
The newly described taxon is indicated in blue and ex-type strains are
indicated in bold. Classification of the subgenus and sections follows
De Crop et al. (2017)
subcylindrical to cylindrical; in the cap centre, terminal cells
subcylindrical to cylindrical, with rounded or obtuse tips,
measuring (19–)20–25–29(–35) 9 3–4–4.5(–5) lm; subterminal cells cylindrical. Pileocystidia near the pileus margin aseptate, short to very long, measuring (2.5–)4–5–6(–7)
lm broad, subcylindrical to cylindrical, sometimes with
lateral projections, obtuse-rounded or sometimes swollen
apex, almost completely filled with dense granular content in
congo red, distinctly grey-black in sulfovanillin (SV), without incrustations. Pileocystidia near the pileus centre aseptate, short to very long, measuring (3)4–5–6(–8) lm broad,
slightly larger, subcylindrical to cylindrical, sometimes with
lateral projections, obtuse-rounded or swollen or sometimes
forked apex, almost completely filled with dense granular
content in congo red, distinctly grey-black in sulfovanillin
(SV), without incrustations. Clamp connections and laticiferous hyphae absent from all tissues.
Material examined: INDIA, Uttarakhand, Rudraprayag
District, Chopta, N30°28.9970 E79°10.2980 , alt. 2632 m,
under Quercus sp., 24 July 2016, A. Ghosh, AG 16-1186
(CAL, holotype; GUH, isotype); ibid., N30°28.6930
E79°11.6360 , alt. 2315 m, under Quercus sp., 25 August
2016, A. Ghosh, AG 16-1363; Pabdhar, N30°29.3760
E79°09.6730 , alt. 2356 m, under Quercus sp., 25 July 2016,
A. Ghosh, AG 16-1206; Bageshwar District, Dhakuri,
N30°04.9700 E79°55.1340 , alt. 2586 m, under Quercus sp.,
2 August 2016, A. Ghosh (AG 16-1269, paratype);
Baniyakund, N30°28.9980 E79°10.6570 , alt. 2614 m, under
Quercus sp., 7 August 2017, A. Ghosh, AG 17-1529;
Baniyakund, N30°29.0910 E79°10.4000 , alt. 2565 m, under
Quercus sp., 8 August 2017, A. Ghosh, AG 17-1549.
Host and habitat: Growing caespitose, under Quercus
sp. (Fagaceae) in mixed forests dominated by Quercus,
Rhododendron (Ericaceae) and Abies (Pinaceae).
GenBank numbers: ITS = MG897815 (AG 16–1186);
ITS = MG925207 (AG 16–1363).
Notes: The combination of macro- and micromorphological features such as, taste being usually acrid or very
acrid, epicutis containing well characterized dermatocystidia, yellowish white spore print, often pinkish or lilac
stipe, mostly adnate or subdecurrent lamellae and red pilei
growing in deciduous forests place Russula choptae under
subg. Russula sect. Russula subsect. Sardoninae (Sarnari
1998).
In the field Russula choptae is distinct from other species of this group by its brownish red, pale red, pastel red to
madder red finely wrinkled pileus with intermixed of pink,
orange to yellowish tinge of coral red at the centre, long
and slim stipe with yellowish white (1A2) areas or orange
white to pale orange (5A2–3) tinge at the base; lamellae
with bifurcate, different lengths at the apex of stipe,
lamellulae with different lengths (up to 5 series); stipe
cuticle and context turns reddish brown and greyish green
with guaiacol and KOH, respectively, and micromorphologically it is separated from allied species by different
types of cystidial apex (capitate, appendiculate, pointed,
mucronate, moniliform or rounded) and basidiospores with
warts that are linked as small crests and thick ridges,
forming an incomplete network and intermixed with isolated warts.
Based on the BLASTn search in NCBIs GenBank
nucleotide database, the closest hit using ITS sequences (AG
16-1186 and AG 16-1363) is Russula sp. (voucher HKAS
78400 GenBank no. KF002784) collected from China.
Unfortunately, there is no micro- and macromorphological
details available for this Chinese collection in published literature. Our phylogeny (Fig. 161) shows R. choptae as sister
to this unknown Chinese Russula sp.
In subsect. Sardoninae, three species Russula persicina
Krombh., R. exalbicans (Pers.) Melzer & Zvára and R.
gracillima Jul. Schäff. (also appeared to be close in ITS
phylogeny in Fig. 161) resemble R. choptae. However,
R. persicina, originally reported from Europe, has larger
pileus (40–100 mm), widely spaced rather sinuate lamellae,
basidiospores, mostly with isolated warts (0.8 lm) with few
crests (Galli 1996; Sarnari 1998; Kränzlin 2005; Knudsen
et al. 2012). Russula exalbicans (originally reported from
Europe), differs in having larger (50–100 mm diam.), rosered, pink, vinaceous, rapidly becoming paler to almost
entirely greenish white pileus usually with a trace of pink at
the extreme margin, firm white stipe which soon becomes
grey with age or in wet weather, frequent 0–1-septate dermatocystidia, larger basidiospores (7–10 9 5–7 lm) with
low (up to 0.75 lm high) ornamentations and ochre spore
print (Pearson 1950; Galli 1996; Sarnari 1998; Kränzlin
2005). Russula gracillima (also reported from Europe), differs from R. choptae in possessing moderately spaced
lamellae, presence of abundant, broad (5–10 lm) dermatocystidia and basidiospores with isolated blunt warts (Rayner
1985; Galli 1996; Sarnari 1998; Kränzlin 2005; Knudsen
et al. 2012; Kibby 2014). Russula renidens Ruots. et al.,
another taxon in subsect. Sardoninae and originally reported
from Europe, has bright crimson red, blood-red to copper red
123
Fungal Diversity
Fig. 159 Lactifluus midnapurensis (CAL 1516, holotype). a Fresh basidiomes in the field. b Basidiome showing lamellae side. c SEM
microphotograph of basidiospores. d Pleurocystidia. e Pileipellis. Scale bars a, b = 10 mm, c = 5 lm, d = 20 lm, e = 10 lm
123
Fungal Diversity
larger pileus (35–80(–100) mm), sulcate margin with age,
some intervening lamellae, cylindric to clavate stipe
(40–100 9 10–22 mm), basidiospores with low ornamentation (up to 0.5 lm) consisting of small warts with some
clusters or ridges, 0–2-septate and slender to clavate dermatocystidia (Galli 1996; Sarnari 1998; Knudsen et al. 2012).
Five more taxa of this subsection, Russula sanguinea
Fr., R. queletii Fr., R. sardonia Fr., R. torulosa Bres. and R.
thindii K. Das & S.L. Mill. also differ from R. choptae.
Russula sanguinea, originally reported from Europe, has a
coniferous habitat especially under pines and elliptical
larger basidiospores (7–9 9 6–7 lm), with ornamentation
consisting mostly of solitary warts which are somewhat
elongated (only a few with connections) (Rayner 1985;
Sarnari 1998; Kränzlin 2005; Knudsen et al. 2012). Russula
queletii (originally reported from Europe) can be distinguished by presence of a combination of characteristics i.e.
vinaceous red to violet pileus and stipe, strong fruity odour
and changing of pilear context to light vinaceous red with
FeSO4 (Rayner 1985; Saini and Atri 1989; Sarnari 1998;
Kränzlin 2005; Knudsen et al. 2012). Similarly, R. sardonia, originally reported from Europe, differs in having dark
violaceous purple, reddish purple or livid vinaceous pileus,
sometimes with greyish olivaceous or with yellowish patches; solid and firm stipe; slightly larger basidiospores
(7–9 9 6–7 lm) and is mycorrhizal with Pinus on acid soil
(Sarnari 1998; Knudsen et al. 2012). Russula torulosa
(originally reported from Europe) possess purple-violet,
purple-red, violet-red or dark red pileus sometimes with
lighter ochraceous or olivaceous spots, often blackish at
centre; solid and firm stipe; larger basidiospores
(7.2–9.5 9 5.6–7.3 lm)
with
low
ornamentations
(0.4–0.6 lm) and is found mostly under Pinus sp. (Sarnari
1998; Knudsen et al. 2012). Russula thindii (originally
reported from India) is distinct by its larger pileus
(38–77 mm diam.), pale yellow spore print and larger
basidiospores (7.5–9–10 9 6–7–8 lm), with ornamentation composed mostly of isolated conical to spinoid warts
being occasionally connected by fine connectors (Das et al.
2014).
In Eastern Asia, Russula choptae might be confused
with R. chiui G.J. Li & H.A.Wen, R. zhejiangensis G.J. Li
& H.A. Wen and R. minutula var. minor Bi (originally
described from China). However, R. chiui differs in having
brightly red tinged pileus, white stipe, septate pileocystidia,
densely reticulate larger basidiospores [(8–)8.5–10(–
10.5) 9 7–8(–8.5) lm] and sterile lamellar edge (Li et al.
2015). Russula zhejiangensis has a purely white stipe,
isolated conical to obtuse warts in basidiospores, shorter
pleurocystidia (35–74 9 6–11 lm) with mostly rounded or
mucronate apex, sterile lamellar edge and 0–3-septate
pileocystidia (Li et al. 2011). Russula minutula var. minor,
described in Guangdong Province of South China, differs
by its white to yellowish gills, white stipe, isolated spinoid
basidiospores and shorter pleurocystidia (Bi and Li 1986;
Li et al. 2011).
Russula uttarakhandia A. Ghosh & K. Das, sp. nov.
MycoBank number: MB820834; Facesoffungi number:
FoF04954, Figs. 165, 166
Etymology: Referring to the name of an Indian state,
Uttarakhand from where the species was collected.
Holotype: CAL 1537
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Basidiomes 70–90 mm high. Pileus 45–60 mm diam.,
planoconvex to applanate with slightly depressed centre,
margin decurved to plane with maturity, entire, tuberculately striate, surface dry, viscid when moist, gelatinous,
glossy, pinkish white or reddish white (10A2) to venetian
pink (10A3) and centrally olive yellow (2B6–7) to greyish
yellow (2A4–6), discoloring to colourless with KOH,
cuticle peeling 1/3 of the radius, chalky white (1–2A1)
beneath the cuticle; pileus context chalky white (1–2A1),
unchanging on bruising. Lamellae adnate to adnexed, close
to rather crowded (7–10 cm at pileus margin), chalky white
(1A1), entire, with concolorous edges, unchanging when
bruised. Lamellulae present, in 2 series. Stipe
55–70 9 10–12 mm, cylindric, central, dry, smooth, longitudinally striate, brittle, chalky white (1–2A1) turning
pink or red rose (11A3–5) with guaiacol; stipe context
hollow, chalky white (1–2A1), unchanging when bruised
but pink or red rose (11A3–5) sometimes after the applications of guaiacol. Odour indistinctive. Taste mild. Spore
print yellowish white (2A2). Hymenophoral trama mainly
composed of large nests of sphaerocytes and few hyphal
elements. Subhymenium layer up to 15 lm thick, pseudoparenchymatous. Basidia (26–)28.5–33–37(–45) 9 (9–
)12–14–15.6 lm, 4-spored, clavate, sterigmata up to 6 lm
long. Basidiospores (6–)6.5–7–8(–9) 9 (5–)5.5–6–6.8(–7)
lm, [n = 40, Q = (1–)1.1–1.2(–1.3)], broadly ellipsoid to
subglobose, rarely globose and ellipsoid; ornamentation
amyloid, composed of isolated spines (up to 1.1 lm high),
suprahilar plage amyloid; apiculus up to 2 lm high.
Pleurocystidia (48–)50–59–67(–71) 9 7–10–12(–14) lm,
cylindrical, subclavate to clavate with capitate, rounded,
moniliform, appendiculate (appendages up to 23 lm long)
or blunt apex; emergent up to 35 lm, partly filled with
heteromorphous, mostly thick coarsely crystalline content
in congo red, sulphovanillin (SV) negative. Lamellae edge
fertile with basidia. Cheilocystidia not found. Pileipellis up
to 65 lm thick, metachromatic in cresyl blue, not sharply
delimited from the underlying spherocytes of the context,
an ixotrichoderm type, distinctly divided in a 15–18 lm
deep and gelatinized suprapellis of erect or ascending
hyphal ends and primordial hyphae, gelatinous layer
123
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123
Fungal Diversity
b Fig. 160 Lactifluus midnapurensis (CAL 1516, holotype). a Basid-
iomes. b Basidia. c Pleuromacrocystidia. d Cheilocystidia. e Pleuropseudocystidia. f Pileipellis. g Basidiospores. Scale bars
a = 10 mm, b–f = 10 lm, g = 2 lm. Drawings by S. Paloi
containing branched, septate, hyphal endings, and underneath this suprapellis, 50–47 lm deep, subpellis of interwined, compact, more or less horizontally oriented hyphae.
Hyphal extremities near the cap margin composed of a
single or a few cells only; the terminal cells subcylindrical,
subclavate to clavate with tapered base, measuring (22–
)25–31–37(–41) 9 (3–)4–5–6(–7) lm, with rounded or
obtuse tips, subterminal cells cylindrical; in the cap centre,
terminal cells slightly shorter in length, subcylindrical,
subclavate to clavate with tapered base, measuring (20–
)21–26–30(–38) 9 (3–)4–5–6(–7) lm, with rounded or
obtuse tips; subterminal cells cylindrical. Primordial
hyphae comparatively longer than the other extremities and
often protruding or repent on the cap surface, occurring
mostly singly, composed of 5–7–9(–10) lm broad, cylindrical cells, obtuse apex, thick-walled (up to 1 lm thick);
incrustations present. Clamp connections and laticiferous
hyphae absent from all tissues.
Material examined: INDIA, Uttarakhand, Rudraprayag
District,
Baniyakund,
alt
2634 m,
N30°28.9140
0
E79°10.854 , 17 July 2015, A. Ghosh, AG 15-670 (CAL
1537, holotype); ibid., alt 2560 m, N30°292070
E79°10.2580 , 24 July 2016, A. Ghosh, (AG 16-1190
paratype).
Habitat and distribution: Under Quercus sp. in mixed
forests dominated by Quercus, Rhododendron and Abies.
GenBank numbers: ITS = KY873997 (CAL 1537);
ITS = MF684758 (AG 16-1190).
Notes: The combination of macro- and micromorphological features such as: epicutis with primordial hyphae,
absence of dermatocystidia, mild taste, yellow to ochre
spore print, nonpruinose cap and epicutis with clavate
hyphal ends undoubtedly place Russula uttarakhandia
under subg. Incrustatula, sect. Amethystinae subsect. Chamaeleontinae (Sarnari 1998). In the field R.
uttarakhandia is distinct from other known species of this
group by its pinkish white or reddish white to venetian pink
and centrally olive yellow to greyish yellow gelatinous,
glossy pileus; stipe that turns pink or red rose after sometime with application of guaiacol, pale yellowish cream
spore print. Micromorphologically, it is separated from
allied species by the combination of various shapes of
cystidial apex (distinctively appendiculate cystidia with
long appendages up to 23 lm long).
Based on the BLASTn search in NCBIs GenBank
nucleotide database, the closest hit using ITS sequences of
Russula uttarakhandia (collection numbers AG 15-670 and
AG 16-1190) is Russula sp. (voucher HMAS 277786 collected from China, GenBank no. LT602960), showing 96%
similarity with 98% query coverage. Unfortunately, there is
no morphological details available for this Chinese collection in any published literature. Our phylogeny
(Fig. 162) clearly shows that R. uttarakhandia is genetically closer and sister to unknown Chinese Russula sp.
(LT602960).
In subsection Chamaeleontinae, Russula postiana
Romell and R. risigallina (Batsch) Sacc. also appeared to
be close in ITS phylogeny with R. uttarakhandia
(Fig. 162). But, R. postiana (originally reported from
Europe), differs from R. uttarakhandia by its herbage green
to yellowish green, smooth, shining pileus; bright ochre
yellow lamellae, heavily encrusted primordial hyphae,
slightly larger basidiospores (8–10.2 9 7–8 lm) and its
occurrence under coniferous trees (Galli 1996; Kibby
2014). Russula risigallina (also originally reported from
Europe), differs from R. uttarakhandia by its red or orange
pileus often with paler yellow centre (sometimes entirely
yellowish opaque), strongly intervened egg yellow to
almost orange lamellae and deep yellow spore print (Galli
1996; Sarnari 2005; Kibby 2014). Another taxon of this
subsect., R. lutea (Huds.) Gray (also appeared in the
BLASTn search), originally reported from Europe, separated from R. uttarakhandia by its entirely yellow, apricot
or coral pileus, deep ochre gills and deep ochraceous spore
print (Rayner 1985; Kibby and Fatto 1990).
Considering the cap colouration, R. uttarakhandia may
also be confused in the field with R. gracillima Jul. Schäff.,
R. cremeirubra Murrill, R. robertii J. Blum, R. nitida
(Pers.) Fr. and R. decipiens (Singer) Svrcek. However,
R. gracillima (originally described from Europe) possess
moderately spaced lamellae, a little acrid taste, presence of
abundant broad dermatocystidia (5–10 lm) and basidiospores with isolated blunt warts (Rayner 1985; Galli
1996; Sarnari 1998; Chou and Wang 2005; Kibby 2014).
Russula cremeirubra (originally reported from Florida) has
rather distant straw coloured lamellae, basidiospores with
amyloid warts connected by fine lines or connectors which
combine to form few meshes and broken reticulum, pilear
hyphae (2–6 lm broad) with acute to rounded apex
(Murrill 1945; Hesler 1960; Kibby and Fatto 1990; Fatto
1998). Russula robertii (originally described from Europe)
has 1–2-septate, slender dermatacystidia (3–5 lm broad)
and spores with many crests and connectives forming
partial to complete reticulum (Sarnari 2005; Kibby 2014).
Russula nitida (originally reported from Europe) has pale
straw-yellow gills, sometimes with reddish edge near the
cap margin, larger basidiospores (8–10.5(–12) 9 6.5–8(–9)
lm) with rather sharp spines (up to 1.2 lm high), isolated
or variously connected, and abundant dermatocystidia with
1–3 septation (Rayner 1985; Galli 1996; Sarnari 2005).
123
Fungal Diversity
123
Fungal Diversity
b Fig. 161 The ITS sequence of the newly generated Russula species
(Russula choptae) plus those acquired from GenBank, UNITE
database and relevant literature, were aligned with the help of
AliView (Larsson 2014) using default settings. Sequences of ITS
were phylogenetically analyzed using Bayesian inference analysis
(BI). Bayesian inference was computed independently twice in
MrBayes v.3.2.2 (Ronquist et al. 2012). The best-fit substitution
model of nucleotide evolution (TIMef) was carried out in MrModeltest 3.7 (Posada and Crandall 1998). Posterior probabilities (PP) were
calculated in two simultaneous runs with Markov chain Monte Carlo
(MCMC) algorithm (Larget and Simon 1999). Markov chains were
run for 1000000 generations, saving a tree every 100th generation.
Default settings in MrBayes were used for the incremental heating
scheme for the chains (3 heated and 1 cold chain), unconstrained
branch length [unconstrained: exponential (10.0)] and uninformative
topology (uniform) priors. The analysis was terminated when the
average standard deviation of split frequencies fell below 0.01. The
first 25% of trees were discarded as burn-in (Hall 2004). The
convergence of runs was visually assessed using Trace function in
Tracer version 1.6 (Rambaut et al. 2013). The novel species having
GenBank no. MG897815 and MG925207 (ITS) are shown in blue and
bold on the tree and ex-type strains are in bold. Russula pseudoamoenicolor and R. violeipes are considered as the outgroup taxa
Russula decipiens (belonging to subg. Russula), originally
reported from Europe has slightly infundibuliform pileus,
yellowish white stipe that slightly becomes grey on bruising, acrid taste and presence of aseptate, subclavate to
clavate dermatocystidia (Rayner 1985; Sarnari 1998).
In field, Russula uttarakhandia may be confused with R.
rajendrae A. Ghosh & K. Das (originally described from
India and also collected from Uttarakhand, Rudraprayag
District, Baniyakund), but the latter possess an acrid taste,
numerous pileocystidia in cuticle and belongs to subg.
Russula (Ghosh and Das 2017).
Considering the acystidiate aspect of pileipellis and mild
taste (key characters of subg. Incrustatula), Russula
uttarakhandia is also similar to other taxa of subg. Incrustatula reported from India viz. R. sharmae K. Das et al., R.
dafianus K. Das & J.R. Sharma, R. dhakuriana K. Das et al.,
R. hookeri S. Paloi et al. and R. kewzingensis K. Das et al.
However, R. sharmae (reported for the first time from West
District of Sikkim) possess red to scarlet with yellow blotched larger pileus (70–140 mm), absence of lamellulae,
smaller basidiospores (6.8–7.3–7.9 9 6–6.6–7 lm) and
palisade to trichopalisade type of pileipellis (Das et al. 2013).
Russula dafianus (reported from Dafia Dhura, Uttarakhand)
has smaller basidiomes (35–45 mm), yellowish white to pale
yellow lamellae and encrusted pilear hyphae (Das and
Sharma 2005). Russula dhakuriana (reported from Dhakuri,
Uttarakhand) has more robust basidiomes (pileus
80–120 mm diam., stipe 50–125 9 20–28 mm), longer
basidiospores (6.5–10.2 9 6–7.7 lm) and pleurocystidia
(60–115 9 7–12 lm) and cellular nature to the subpellis
(Das et al. 2006a). Russula hookeri (reported from Darjeeling
Hill, Eastern Himalaya) has small to medium basidiomes
(20–35 mm), greyish red to greyish rose pileus with brownish red centre, concolorous stipe; smaller basidiospores
(5.4–7.2 9 4.5–6.4 lm) and white spore print (Paloi et al.
2015). Russula kewzingensis (reported from Kewzing, Sikkim Himalaya) has deep and intense red pileus, red-flushed
stipe and pseudoparenchymatous nature of subpellis (Das
et al. 2017b).
Stereaceae Pilát
The family Stereaceae was proposed by Pilát (1930)
with Stereum Hill ex Pers. as generic type. This family is
one of the most widespread and diverse in the Russulales,
together with Russulaceae. It includes species with basidiome appressed, effuse-reflexed or discoid, rarely stalked;
pileus often zoned; monomitic or dimitic hyphal system;
hymenophore smooth to tuberculate; basidia and spores
hyaline, smooth, amyloid or non-amyloid. The family
comprises species that grow in exposed positions, such as
dead branches still attached to hardwood or conifer trees,
causing white rot (Miller et al. 2006; Larsson 2007).
Aleurodiscus Rabenh. ex J. Schröt.
Aleurodiscus is one of the largest genera in Stereaceae.
It was proposed by Rabenhorst (1874) without diagnosis,
and validated by Schröter in (1888), with Peziza amorpha
Pers. (: Aleurodiscus amorphus (Pers.) J. Schröt.) as type
species. Pilát (1926) included Aleurodiscus corticioid fungi
with large basidia and large spores, as well as variable
sterile elements and this concept was largely accepted
(Bourdot and Galzin 1912, 1928; Burt 1918). Lemke
(1964a, b) considered the genus to be artificial and proposed to exclude species with inamyloid spores, and
described the following new genera: Aleurocystidiellum
Lemke, Aleurocorticium Lemke and Licrostroma Lemke.
On the basis of morphological and molecular data, Wu
et al. (2000) described Acanthofungus Sheng H. Wu et al.
with Acanthofungus rimosus Sheng H. Wu et al. as type
species, and Neoaleurodiscus Sheng H. Wu with
Neoaleurodiscus fujii Sheng H. Wu as type species. From a
phylogenetic perspective, Wu et al. (2001) analyzed the
limits of Aleurodiscus sensu lato and the monophyly of
previously segregated genera, concluding that Aleurocystidiellum and Acanthobasidium are monophyletic. However, subsequent studies (Larsson and Larsson 2003; Miller
et al. 2006; Larsson 2007) have shown that Aleurocystidiellum does not belong to Stereaceae. Wu et al. (2001) set
the boundaries of Aleurodiscus sensu stricto and confirmed
that Aleurodiscus sensu lato is paraphyletic.
According to Wu et al. (2001), Aleurodiscus sensu lato
includes species with highly variable characteristics; these
are not congruent and usually overlap, so the species in
Aleurodiscus sensu lato show all possible combinations of
spore surface from smooth or ornamented, presence or
absence of acanthophyses, and hyphae with clamps or
123
Fungal Diversity
Fig. 162 The ITS sequence of the newly generated Russula
uttarakhandia plus those acquired from GenBank, UNITE database
and relevant literature, were aligned with the help of AliView
(Larsson 2014) using default settings. Sequences of ITS were
phylogenetically analyzed using Bayesian inference analysis (BI).
Bayesian inference was computed independently twice in MrBayes
v.3.2.2 (Ronquist et al. 2012). The best-fit substitution model of
nucleotide evolution (TIMef) was carried out in MrModeltest 3.7
(Posada and Crandall 1998). Posterior probabilities (PP) were
calculated in two simultaneous runs with Markov chain Monte Carlo
(MCMC) algorithm (Larget and Simon 1999). Markov chains were
run for 1000000 generations, saving a tree every 100th generation.
Default settings in MrBayes were used for the incremental heating
scheme for the chains (3 heated and 1 cold chain), unconstrained
branch length [unconstrained: exponential (10.0)] and uninformative
topology (uniform) priors. The analysis was terminated when the
average standard deviation of split frequencies fell below 0.01. The
first 25% of trees were discarded as burn-in (Hall 2004). The
convergence of runs was visually assessed using Trace function in
Tracer version 1.6 (Rambaut et al. 2013). The novel species having
GenBank no. KY873997 and MF684758 (ITS-rDNA) are shown in
blue and ex-type strains are in bold. Russula fellea, R. sanguinea and
R. persicina are considered as the outgroup taxa
simple-septate. The novel species, Aleurodiscus patagonicus is introduced following the treatment in Wu et al.
(2001), Dai and He (2016), and Dai et al. (2017b, c).
MycoBank number: MB823981; Facesoffungi number:
FoF05726, Figs. 167, 168
Etymology: Named after the Chilean Northern Patagonian region where the holotype and paratypes were
collected.
Aleurodiscus patagonicus Nogal, Telleria, M. Dueñas &
M.P. Martı́n, sp. nov.
123
Fungal Diversity
Fig. 163 Russula choptae (AG 16–1186, holotype). a–e Fresh and
dissected basidiomes in the field. f, g Radial section through
pileipellis showing elements. h Image of basidiospores under SEM.
i–k Transverse section through lamellae showing pleurocystidia.
l Transverse section through lamellae showing cheilocystidia. Scale
bars b = 55 mm, f = 25 lm, g = 10 lm, h = 2 lm, i = 25 lm, j–
l = 10 lm
Holotype: 19609Tell., MA-Fungi 90714
Colour codes follow: ISCC-NBS Centroid Colour Charts
(Kelly and Judd 1976).
Basidiomes first discoid then confluent; margin determinate, involute; hymenophore smooth to slightly reticulate, pale orange yellow to medium orange yellow (73. p.
OY–71. m. OY). Hyphal system monomitic; hyphae thick-
123
Fungal Diversity
Fig. 164 Russula choptae (AG
16-1186, holotype). a Dissected
basidiome. b Radial section
through pileipellis.
c Basidiospores. d Basidia.
e Pleurocystidia.
f Cheilocystidia. Scale bars
a = 10 mm, b–f = 10 lm
walled, 3–6 lm wide; paraphysoid hyphae cylindrical,
occasionally branched, with scattered clamps, 3–5 lm
wide. Basidia clavate, thin-walled, stalked, with basal
clamp, 150–190 9 (22 -)24–27 lm, with four sterigmata,
5–7 lm wide. Basidiospores citriform, smooth, thin-walled, hyaline, strongly amyloid, 19–22(-24) 9 14–16 lm,
Q = 1.38.
Material examined: CHILE, Palena, Comuna Hualaihué,
Huinay Reserve, path to Cerro del Tambor, 42°220 4400 S,
72°240 2500 W, 100 m asl., on unidentified wood, 7 May
2013, M. Dueñas, M.P. Martı́n & M.T. Telleria, 19609Tell.
(MA-Fungi 90714, holotype); ibid., 14537MD (MA-Fungi
90713); Comuna Hualaihué, Base Paula, path of Geyser, on
unidentified wood, 42°240 16.100 S, 72°440 0.5900 W, 52 msl.,
28 May 2012, M. Dueñas, M.P. Martı́n & M.T. Telleria,
14080MD (MA-Fungi 90711).
Habitat and distribution: On unidentified wood in Valdivian temperate rainforest in Chilean Northern Patagonian
region.
123
Additional material examined: Aleurodiscus limonisporus—AUSTRALIA, Victoria, Cumberland falls, on
Nothofagus cunninghamii (Nothofagaceae), 6 June 1954,
A. Miller (PDD 16691, isotype); NEW ZEALAND, Bay of
Plenty, Te Urewera, Tarapounamu, west of road, on
decaying branch, 19 May 2005, B. Paulus & M. Fletcher
(PDD 83502); Te Urewera, Tarapounamu, east of road, on
decaying wood, 17 May 2005, B. Paulus (PDD 83552); Te
Urewera, Tarapounamu, west of road, on fallen branch, 19
May 2005, B. Paulus & M. Fletcher (PDD 83553); Buller,
North of Reefton, Perseverance Bridge, 5 May 2001, E.
Johannesen (PDD 72991); Paparoa National Park, Bullock
Creek Farm, on N. fusca, 26 April 1987, P.K. Buchanan
(PDD 55241); Paparoa Ranges, Tiropahi Walk, on N.
menziesii, 27 April 1987, P.K. Buchanan (PDD 55264);
Victoria Forest Park, east of Maruia, on N. menziesii, 22
April 1986, R.E. Beever (PDD 53413); Canterbury,
Arthur’s Pass National Park, Dobson Nature Walk, on
Olearia capillaris (Asteraceae), 20 November 1988, P.K.
Fungal Diversity
Fig. 165 Russula uttarakhandia (CAL 1537, holotype). a–d Fresh
basidiomes in the field. e, f Radial section through pileipellis showing
elements. g Basidia. h Basidiospores. i SEM image of basidiospores.
j–m Transverse section through lamellae showing pleurocystidia.
Scale bars a = 80 mm, e–h = 10 lm, i = 2 lm, j–m = 10 lm
Buchanan (PDD 55021); Jollie’s Bush Reserve, Christchurch, on standing dead tree, 4 July 2010, J.A. Cooper (PDD
95980);
Gisborne,
Moanui
Conservation
Area,
38°240 24.8400 S, 177°230 58.9200 E, on Nothofagus, 13 May
2013, S.R. Pennycook (PDD 97004); Hawke’s Bay, Upper
Mohaka River, Kaimanawa Range, on Nothofagus fusca,
May 1953, J.M. Dingley (PDD 12600); Nelson, Abel
Tasman National Park, Hardwoods Hole, on decaying
branch of N. solandri, 15 April 2008, A.J. O’Donnell &
B.C. Paulus (PDD 94131); Kahurangi National Park, Flora
Saddle, on decaying bark, 14 April 2008, A.J. O’Donnell &
B.C. Paulus (PDD 94144); Murchison, on N. fusca, April
1956, S.D. & P.J. Brook (PDD 17122); Southland, Porakino Valley, on bark of N. solandri, 7 May 2012, Lloyd
(PDD 96617); Wairarapa, Tararua Forest Park, Mt Holdsworth, Gentle Annie Track, on N. fusca, 10 May 2007, A.J.
O’Donnell (PDD 92582); Tararua Forest Park, Mt Holdsworth, Donnelly Flat track, on decaying branch, 10 May
2007, B.C. Paulus (PDD 92616); Tararua Forest Park, Mt
Holdsworth track, on decaying wood, 7 May 2007, G.
123
Fungal Diversity
Fig. 166 Russula
uttarakhandia (CAL 1537,
holotype). a Dissected
basidiome. b Basidia.
c Basidiospores.
d Pleurocystidia. e Radial
section through pileipellis.
Scale bars a = 10 mm, b–
e = 10 lm
Gates & D. Ratkowsky (PDD 92829); Wellington, Kaimanawa Range, on N. menziesii, April 1955, J.M. Dingley
(PDD 15227); Mangatorutoru Stream, on N. solandri var.
cliffortioides, March 1948, J.M. Dingley (PDD 7452);
Tongariro National Park, on N. solandri var. cliffortioides,
February 1951, G.H. Cunningham (PDD 15230); Ohakune,
Lake Surprise Track, on Nothofagus sp., April 1935, E.E.
Chamberlain (PDD 15229); Whakapapa-iti Stream, Ruapehu, on N. solandri var. cliffortioides, Sep 1955, J.M.
Dingley (PDD 15355): Whakapapa-iti Stream, Tongariro
National Park, on N. solandri var. cliffortioides, January
1951, J.M. Dingley (PDD 11188); Turangi, near Beggs
Pool, Kaimanawa Range, on Weinmannia racemosa
(Cunoniaceae), 25 May 1970, J.M. Dingley (PDD 28631);
Westland, Granville Forest, Orwell Creek, Ahaura, on N.
fusca, 2 April 1963, J.M. Dingley (PDD 20997). Aleurodiscus sp.–Otago Lakes, Beyond Paradise, on Nothofagus small branch, 7 May 2016, P. Catcheside (PDD
123
110288) [identified as A. limonisporus]. Unidentified–
Glacier Burn Track, on Nothofagus, 8 May 2016, A. Chinn
(PDD 109766) [identified as A. limonisporus].
GenBank numbers: ITS = MF631175, LSU = MF631191
(MA-Fungi 90711); ITS = MF631176, LSU = MF631192
(MA-Fungi 90713); ITS = MF631177, LSU = MF631193
(MA-Fungi 90714); ITS = MF631152, LSU = MF631178
(PDD 7452); ITS = MF631153 (PDD 12600); ITS =
MF631154 (PDD 15229); ITS = MF631155 (PDD 15230);
ITS = MF631156, LSU = MF631179 (PDD 16691); ITS =
MF631157 (PDD 17122); ITS = MF631158 (PDD 28631);
ITS = MF631159 (PDD 53413); ITS = MF631160 (PDD
55021); ITS = MF631161 (PDD 55241); ITS = MF631162
(PDD 72991); ITS = MF631163 (PDD 83502); ITS =
MF631164, LSU = MF631180 (PDD 83552); ITS =
MF631165, LSU = MF631181 (PDD 92582); ITS =
MF631166, LSU = MF631182 (PDD 92616); ITS =
MF631167, LSU = MF631183 (PDD 92829); ITS =
Fungal Diversity
MF631168, LSU = MF631184 (PDD 94131); ITS =
MF631169, LSU = MF631185 (PDD 94144); ITS =
MF631170, LSU = MF631186 (PDD 95980); ITS =
MF631171, LSU = MF631187 (PDD 96617); ITS =
MF631172, LSU = MF631188 (PDD 97004); ITS =
MF631173, LSU = MF631189 (PDD 109766); ITS =
MF631174, LSU = MF631190 (PDD 110288).
Notes: Phylogenetic analysis based on a combined ITS
and LSU sequence dataset (Fig. 169) shows that Aleurodiscus patagonicus clusters within Stereaceae, far away
from the Aleurodiscus sensu stricto clade, which includes
only A. amorphous (Pers.) J. Schröt. (type species) and A.
grantii Lloyd (Wu et al. 2001). Aleurodiscus sensu lato is
paraphyletic; sequencing shows species of apparently
unrelated genera intermingled with the species of Aleurodiscus sensu lato (Wu et al. 2001, 2010b; Larsson and
Larsson 2003; Binder et al. 2005; Miller et al. 2006). In the
phylogeny (Fig. 169), A. patagonicus groups in a highly
supported clade (99% ML, 99% MP and 1.00 PP), which is
sister to the A. limonisporus D.A. Reid clade and this
relationship is also highly support (92% ML, 90% MP and
1.00 BYPP).
Aleurodiscus patagonicus is similar to A. limonisporus.
Both species have a monomitic hyphal system, cylindrical
paraphysoid hyphae, and characteristic citriform smooth
spores. However, A. patagonicus differs in the following
characteristics: clamps present in paraphysoid hyphae,
longer and wider basidia, with basal clamp, and, spores
with higher length/width ratio, as well as, in the geographical distribution (Table 3). Aleurodiscus patagonicus
is reported from Chilean Northern Patagonia, while
A. limonisporus occurs in Australia (Reid 1955) and New
Zealand (McKenzie et al. 2000).
Trechisporales K.H. Larsson
Hydnodontaceae Jülich
Hydnodontaceae was described by Jülich (1981) with
Hydnodon Banker as generic type, and includes Brevicellicium K.H. Larss. & Hjortstam, Cristelloporia I. Johans. &
Ryvarden and Trechispora P. Karst. Hydnodon is now
considered as a synonym of Trechispora (Ryvarden 2002).
This family was characterized by the resupinate, effusedreflexed or pileate basidiome, hymenophore smooth to
hydnoid or poroid, as well as by the ampulate septa, short
cylindrical basidia and ornamented spores. The family
concept was revised by Larsson (2007), on the basis of
morphological and molecular data, and Brevicellicium,
Fibriciellum J. Erikss. & Ryvarden, Fibrodontia Parmasto,
Luellia K.H. Larss. & Hjortstam, Porpomyces Jülich,
Subulicystidium Parmasto, Trechispora and Tubulicium
Oberw. were included.
Trechispora P. Karst.
Trechispora was erected by Karsten (1890) to include only
one species, Trechispora onusta P. Karst. (= Trechispora
hymenocystis). This genus was characterized by its adnate
basidiomes, poroid hymenophore, with hyaline and ornamented spores. Trechispora was treated by Bondartsev and
Singer (1941), and Rogers (1944, 1951), and the initial
description was amended by Liberta (1966) to include species with clamped hyphae, ampulate septa and smooth
spores. Liberta (1973), in his monograph, introduced new
morphological criteria for species delimitation, such as shape
and size of spores, as well as the spore surface (smooth or
ornamented), hyphal system (monomitic or dimitic), and
hymenophore configuration (smooth, poroid or hydnoid),
and this species concept was followed in all subsequent
treatments. Larsson (1994, 1995a, b, 1996) gave detailed
micromorphological studies, described new taxa, proposed
new combinations and recognized the morphology of crystal
deposits on subicular hyphae as a reliable character for species delimitation.
Most species of Trechispora have been described from
boreal and temperate zones, growing on deeply decayed
wood and other debris on the ground, although Matssura
and Yashiro (2010) reported an insect-fungus association
between a termite species (Reticulotermes termites) and an
undescribed trechisporoid fungus (Trechispora sp.) from
Japan. Trechispora contains 82 legitimate names (Index
Fungorum 2019). According to Hjortstam and Ryvarden
(2007), 27 species have been reported from tropical and
subtropical areas. Trechispora echinospora sp. nov. is
introduced from Equatorial Guinea.
Trechispora echinospora Telleria, M. Dueñas, I. Melo &
M.P. Martı́n, sp. nov.
MycoBank number: MB825275; Facesoffungi number:
FoF05727, Fig. 171
Etymology: From Latin ‘‘echinus’’ which means
hedgehog and spore; referring to the characteristic ornamented spores with straight prickles
Holotype: 13858Tell., MA-Fungi 82485
Colour codes follow: ISCC-NBS Centroid Colour Charts
(Kelly and Judd 1976).
Basidiomes resupinate, effused, confluent, adnate;
hymenophore farinaceous to grandinioid, yellowish grey
(93. y Grey) to cream (92. y White–89. p. Y). Margin not
differentiated and strands not seen. Hyphal system monomitic. Subicular hyphae thin-walled, loosely interwoven,
moderately branched, straight, 1–2 lm wide, long-celled,
with clamps and ampullate septate, up to 4 lm wide.
Sphaerocysts thin-walled, with basal clamp, (11–
)15–18 lm diam., sometimes present. Subhymenial hyphae
thin-walled, branched, short-celled, isodiametric, up to
6 lm wide, with clamps in all septae. Crystals not seen.
Basidia short-cylindrical to urniform, four sterigmata,
123
Fungal Diversity
Fig. 167 Aleurodiscus
patagonicus (19609Tell., MAFungi 90714, holotype).
a Basidiome. b Section of
basidiome under SEM;
hymenial layer (1), basal layer
(2). c Probasidia with basal
clamp. d Spores under SEM. e,
f Basidiospores. Notes scanning
electron microscope (SEM) was
used after coating basidiome
samples in gold with Balzers
SCD 004 sputter coater with a
Hitachi S-3000 N SEM. Scale
bars a = 5 mm, b = 50 lm,
c = 10 lm, d–f = 5 lm
13–16(–17) 9 5–6 lm, with basal clamp. Spores globose,
4–5 lm (excluding the ornamentation), thin-walled, echinulate with straight prickles up to 1 lm long, sometimes
with one oil drop in the protoplasm, Q (L/W) = 1.02.
Material examined: EQUATORIAL GUINEA, Centro
Sur Province, road Niefang to Evinayong, next to Nkumekie, on Oxythenantera abyssinica (Poaceae), 7
December 1990, M.T. Telleria, 13858Tell. (MA-Fungi
82485, holotype); ibid., 13861Tell. (MA-Fungi 82486);
13870Tell. (MA-Fungi 91312).
123
Habitat and distribution: Only known from the type
locality in the Continental Region (Mbini) of Equatorial
Guinea, West Tropical Africa, on dead stem of bamboo
(Oxythenantera, Poaceae)
GenBank numbers: ITS = JX392845, JX392847,
LSU = JX392846, JX392848, JX392849 (13858Tell., MAFungi 82485, holotype); ITS = JX392853, JX392850,
JX392852, LSU = JX392851, JX392854 (13861Tell., MAFungi 82486).
Notes: In the framework of our study Telleria et al.
(2013), a number of ITS and LSU sequences of
Fungal Diversity
Fig. 168 Aleurodiscus
patagonicus (19609Tell., MAFungi 90714, holotype). a Basal
layer. b Hymenial layer with
basidia and paraphysoid hyphae.
c Basidiospores. Line drawings
were made with a Leyca
DM2500 microscope with aid of
a drawing tube by M. Dueñas.
Scale bars = 10 lm
Trechispora obtained by our team were deposited in
GenBank under Trechispora sp. because many specimens
were collected in tropical areas, and require a thorough
study of their morphology to clearly identify them to the
species level.
In this paper, ITS sequences from 13858Tell. and
13861Tell. collected during our study of the corticoid fungi
from Equatorial Guinea (Centro Sur Province), growing on
Oxythenantera abyssinica (A.Rich.) Munro, were compared with sequences available in the EMBL/GenBank/
DDBJ (Cochrane et al. 2011) and UNITE (Abarenkov et al.
2011; Kõljalg et al. 2013) databases, mainly from Larsson
et al. (2004), Lutzoni et al. (2004), Krause et al. (2011),
Brazee et al. (2012, 2014), Sjökvist et al. (2012), Telleria
123
Fungal Diversity
et al. (2013), and Ordynets et al. (2015). The sequences
were analyzed under maximum parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI). The BI
tree is shown in Fig. 170, including the MP and ML
bootstrap values, as well as the posterior probabilities. The
five sequences from these collections, group together in a
highly supported clade (85% MP, 99% ML and 1.00
BYPP), sister group of the clade formed by Trechispora
araneosa (Höhn. & Litsch.) K.H. Larss., T. farinacea
(Pers.) Liberta, T. hymenocystis (Berk. & Broome) K.H.
Larss. and T. mollusca (Pers.) Liberta, although this relationship is not well-supported. These collections from
Equatorial Guinea show unique morphological characteristics, such as the spores being globose and echinulate with
straight prickles, striking under SEM (Fig. 171). The
morphological characteristics and molecular phylogenetic
results, its habitat on dead stem of bamboo, and its geographical distribution led us to propose the new species, T.
echinospora.
Subclass Auriculariomycetidae Jülich
Auriculariales J. Schröt.
Auriculariaceae Fr. Ex Lindau
Auriculariaceae, typified by Auricularia mesenterica
(Dicks.) Pers. (= Helvella mesenterica Dicks.), belongs to
Auriculariales and comprises nine genera and 155 species
(Kirk 2017). The family includes species with resupinate,
effused-reflexed, hydnoid, cerebriform and pileate basidiomes; thin or thick-walled probasidia and metabasidia,
both varying from globose to cylindrical and thin-walled
basidiospores that germinate by tubes or producing conidia
(Lowy 1971; Martin 1952). The most common genera of
Auriculariaceae are Auricularia Bull., Heterochaete Pat.,
Exidia Fr., Eichleriella Bres., Exidiopsis (Bref.) A. Møller
and Hirneolina (Pat.) Bres.
Tremellochaete Raitv.
Tremellochaete was described by Raitviı́r (1964) with T.
japonica (Lloyd) Raitv. (= Exidia japonica Lloyd) as the
type species and comprises two legitimate species (Kirk
2017). The genus is characterized by tough-gelatinous
basidiomes, often becoming softer with age, varying from
tuberculate-erumpent and effused-tuberculate; papillate
hymenium, papillae heavily encrusted with irregular, hyaline crystals; metabasidia with 2- or 4-celled; and allantoidsuballantoid, hyaline, aseptate basidiospores. Tremellochaete is similar to Exidia, which has fewer papillae
(Raitviı́r 1964), and has been considered a synonym of the
latter. However, Tremellochaete was reinstated after morphological and phylogenetic analyses (Malysheva and
Spirin 2017). We follow the treatments and updated
accounts in Malysheva and Spirin (2017). The new species,
T. atlantica is introduced based on phylogenetic analysis of
123
a combined ITS and LSU sequence dataset coupled with
morphological characteristics (Fig. 172).
Tremellochaete atlantica Alvarenga, sp. nov.
MycoBank number: MB823716; Facesoffungi number:
FoF05728, Fig. 173
Etymology: The name refers to the phytophysiognomy
where it was collected.
Holotype: URM 90199
Colour codes follow: Methuen Handbook of Colour
(Kornerup and Wanscher 1978).
Basidiomes yellowish to greyish or brown (1B, 2A, 5E,
4D) when fresh, greyish brown to yellowish brown (3F,
8E) when dry, foliose, 2.5–3 9 3.5 cm, gelatinous, densely
papillate, papillae 50–107.5 9 40 lm, encrusted with
irregular, hyaline crystals not dissolving in KOH, margin
detaching from the substrate, marginal hairs present in
tufts, 50–212 lm or absent, subhymenial hyphae with
ochraceus, granular content. Basidia with complete septum, 2- or 4-celled, hyaline, ovoid, 9–12(–13) 9 6–9(–10)
lm, clamped at the base. Basidiospores allantoid to reniform, 7.8–10(–12) 9 2–5 lm, thin-walled, IKI-. Hyphal
system monomitic, hyphae clamped, thin-walled, hyaline,
1–2 lm in diam. Cystidia absent.
Material examined: BRAZIL, Pernambuco: Igarassu,
Refúgio Ecológico Charles Darwin, May 2017, R.L.M.
Alvarenga, RLMA 491 (URM 90198); ibid., Pernambuco:
Recife, Centro de Biociências, near the Departamento de
Zoologia, Universidade Federal de Pernambuco, June
2017, R.L.M. Alvarenga, RLMA 477 (URM 90199 holotype; isotype in O).
GenBank
numbers:
ITS = MG594382,
LSU =
MG594384 (URM 90198); ITS = MG594381, LSU =
MG594383 (URM 90199).
Notes: Tremellochaete atlantica is recognized by the
marginal hairs ranging from 50–212 lm and the heavily
encrusted papillae with hyaline, irregular crystals. It is
similar to T. japonica and T. nigerima (Viégas) Spirin & V.
Malysheva. Tremellochaete japonica (type locality Japan)
differs by the darker basidiomes, fewer dark hyphae,
smaller papillae (60–75 lm), absence of marginal hairs and
slightly smaller basidiospores (9.5–12 9 4–4.5 lm)
(Roberts 2006), while T. nigerima (type locality Brazil) has
larger metabasidia and basidiospores (20–25 9 10–15 lm
and 17–20 9 7–8 lm, respectively) (Viégas 1945).
Tremellochaete hispidula (Lowy) Raitv. (= Exidia hispidula Lowy, from USA) also has marginal hairs (up to
125 lm), but lacks clamps, has larger metabasidia [(7–)8–
10 9 (10–)12–16] and ovoid to allantoid basidiospores
[(9–13(–14.5) 9 5–7)] (Lowy 1957). This species, however, is illegitimate, because Exidia hispidula Berk. has
priority over E. hispidula Lowy.
Fungal Diversity
Fig. 169 Bayesian majority rule consensus topology based on a
combined ITS and LSU DNA sequence dataset. Two Gloeodontia
species were included as the outgroup taxa. Maximum parsimony and
maximum likelihood bootstrap support (C 50%), and posterior
probability (C 0.95 BYPP) are indicated above branches. Newly
Table 3 Comparison of
morphological characteristics
and distribution between
Aleurodiscus patagonicus and
A. limonisporus
generated sequences are marked in blue and ex-type strains are in
bold. Scale bar indicates substitution per site. Sequences were
retrieved from EMBL Nucleotide Sequence Database (Cochrane et al.
2015), and they were published in Wu et al. (2001), Larsson and
Larsson (2003), Wu et al. (b), Dai and He (2016), Dai et al. (2017a, b)
Morphology and distribution
A. patagonicus
A. limonisporus
ANOVA
Basidia length (lm)
150–190
130–160(-190)
F = 31.26, p < 0.05
F = 21.45, p < 0.05
Basidia width (lm)
(22 -)24–27
20–25(-26)
Basal clamp in basidia
Present
Absent
–
Spore length, L (lm)
Spore width, W (lm)
19–22(-24)
14–16
20–24(-26)
12–16
F = 0.808, p = 0.379
F = 4.195 p = 0.0532
Q Index, L/W
1.36–1.40
1.41–1.60
F = 10.02, p < 0.05
Distribution
Chile: Huinay Reserve
Australia, New Zealand
–
ANOVA analyses were performed to assess the significance of basidia and spore morphology using the
function ‘‘aov’’ of the stats R package v 3.2.1 (R Core Team 2015)
Phylogenetic analysis of a combined ITS and LSU
sequence dataset shows that Tremellochaete atlantica
forms a well-resolved clade in Tremellochaete clade as a
sister group to T. japonica, collected in Russia (Fig. 172).
123
Fungal Diversity
Class Dacrymycetes Doweld
The classification of the families in Dacrymycetes follows Oberwinkler (2014), Zhao et al. (2016) and Shirouzu
et al. (2017).
Dacrymycetales Henn.
Dacrymycetaceae J. Schröt.
Dacrymycetaceae was introduced by Schröter (1889),
with the genera Calocera (Fr.) Fr., Dacrymyces Nees,
Dacryomitra Tul. & C. Tul. and Guepinia Fr. Nowadays,
according to Oberwinkler (2014) this family contains ten
genera: Arrhytidia Berk. & M.A. Curtis, Calocera,
Dacrymyces, Dacryomitra, Dacryonaema Nannf., Dacryopinax G.W. Martin, Ditiola Fr. (= Guepinia), Femsjonia
Fr., Guepiniopsis Pat. and Heterotextus Lloyd.
The species included in this family are characterized by
the form of basidiome: pustulate, cupulate, clavate, or
sometimes branched; gelatinous when fresh and usually
yellow or orange. The hyphal system is monomitic, the
basidia bifurcate, with two well-developed sterigmata and
the spores are usually septate germinating by conidia or
germ tubes. All species are wood-decaying and cause
brown rot (Oberwinkler 1993). The genera in this family
have been delimited according to morphological criteria
such as the shape of basidiome, the position of the hymenium, and presence or absence of a specialized cortex
(Kobayasi 1939; McNabb 1973; Reid 1974). Phylogenetic
studies (Weiß and Oberwinkler 2001; Shirouzu et al.
2007, 2009, 2013, 2017) showed that Calocera, Cerinomyces, Dacrymyces, and Dacryopinax are polyphyletic,
suggesting that the traditional taxonomy should be revised.
We follow the treatment and updated accounts in Shirouzu et al. (2017) as well as following Reid (1974) for
taking into account Dacrymyces sensu lato. The species of
this genus are characterized by the basidiomes being
gelatinous when fresh, at first pustular, becoming pulvinate, discoid or turbinate, sometimes stipitate; hymenium
amphigenous, or restricted to the upper surface; hyphal
system monomitic; hyphidia simple, or somewhat branched, with or without clamp connections; basidia clavate
becoming bifurcate; spores cylindrical, ellipsoid, allantoid,
ovate or subglobose, becoming variously septate at maturity and germinating by conidia or germ-tubes. The novel
species, Dacrymyces invisibilis is introduced based on
morphological characteristics coupled with ITS phylogenetic analyses (Fig. 174).
Dacrymyces invisibilis M. Dueñas, Telleria & M.P. Martı́n,
sp. nov.
MycoBank number: MB825361; Facesoffungi number:
FoF05729, Figs. 175, 176
Etymology: From Latin ‘‘invisibilis’’ not visible, refers
to dry basidiome difficult to see.
123
Holotype: 14597MD, MA-Fungi 91306.
Basidiomes pustulate at first, pulvinate to discoid, confluent, 0.5–3 mm diam., gelatinous, white, becoming varnish-layered when dry; hymenophore amphigenous,
smooth; margin determinate. Hyphal system monomitic.
Hyphae thin- to thick-walled, smooth, or incrusted, loosely
interwoven, sparsely ramified, 2–4 lm wide, without
clamps. Basidia 37–48 9 5–6 lm, hyaline, guttulate,
subcylindrical to subclavate, without basal clamp, two
sterigmata, 21–30 9 3 lm. Spores (14–)15–18 9 5–6(–7)
lm (
x = 15.8 9 5.95 lm), hyaline, ellipsoid, 3-septate at
maturity, thin-walled, smooth, with oil drops, Q (L/
W) = 2.65.
Material examined: CHILE, Los Lagos, Palena, comuna
Hualaihué, Hornopirén, Huinay Biological Reserve, behind
the hydroelectric power station, next to the beach,
42°220 53.400 S 72°240 55.600 W, on unidentified wood, 9 May
2013, M. Dueñas, M.P. Martı́n & M.T. Telleria, 14597MD,
(MA-Fungi 91306, holotype); ibid., CHILE, Los Lagos,
Palena, comuna Hualaihué, Hornopirén, Huinay Biological
Reserve, ‘‘Cementerio de los Alerces’’ experimental plot,
42°220 01.500 S 72°240 57.800 W, 50 msl, on unidentified wood,
10 May 2013, M. Dueñas, M.P. Martı́n & M.T. Telleria,
14617MD (MA-Fungi 91307).
Habitat and distribution: Known from the Valdivian
temperate rainforest in the Chilean Northern Patagonian
region, growing on unidentified wood.
GenBank numbers: ITS = MH230100 (14597MD, MAFungi 91306); ITS = MH230101 (14617MD, MA-Fungi
91307).
Notes: Previous ITS/LSU analyses (data not shown) of
Dacrymyces invisibilis, and 119 homologous sequences
from Shirouzu et al. (2017), included this taxon with D.
subantarcticensis Burds. & Laursen described from New
Zealand (Burdsall and Laursen 2004). However, morphological and molecular features clearly separate the two
taxa. In this paper, a shorter and more accurate alignment is
reported, including the species closest to D. subantarcticensis in Shirouzu et al. (2017): D. aureosporus Shirouzu
& Tokum., D. chrysospermus Berk. & M.A. Curtis, D.
dictyosporus G.W. Martin, D. minor Peck, D. subalpinus
Kobayasi, D. stillatus Nees and Guepiniopsis buccina
(Pers.) L.L. Kenn. In the UPGMA tree (Fig. 174) based on
Kimura-2-parameters, the genetic distances proposed in
Schoch et al. (2012) as barcode analyses, are shown,
including the parsimony bootstrap and maximum likelihood support values, as well as the Bayesian posterior
probabilities. The two collections from Chile form a wellsupported group (81% MP, 78% ML and 1.00 BYPP)
indicating that D. invisibilis is distinct from D. subantarcticensis. In addition, in this figure a small part of the
alignment (around positions 189 and 246) is included to
show some of the variable positions between these two
Fungal Diversity
Fig. 170 Phylogram generated
from Bayesian inference
analysis of ITS sequence dataset
of Trechispora sequences.
Maximum parsimony and
maximum likelihood bootstrap
support values C 50% (10,000
replicates both), and Bayesian
posterior probabilities C 80%
are shown above the nodes. The
tree is rooted with
Brevicellicium exile and B.
olivascens. Newly generated
sequences are marked in blue
and ex-type strains are in bold
species. Dacrymyces invisibilis and D. subantarcticensis
have basidiomes nearly invisible when dry, and thin-walled
spores, with three septa, but they differ in the size of
basidia and spores, smaller in D. subantarcticensis
(20–30 9 5 lm and 10–13 9 4.5–6 lm, respectively).
Dacrymyces subalpinus, D. aureosporus, D. chrysospermus and D. dictyosporus form a subclade close to D.
subantarcticensis and D. invisibilis; all of them have spores
with more than three septa at maturity. Dacrymyces subalpinus and D. aureosporus are known only from Japan
(Kobayasi 1939; Shirouzu and Hosoya 2017). Dacrymyces
chrysospermus has been widely reported and D. dityosporus is known from Honduras, and Mexico (Lowy 1971).
Mucoromycota Doweld
We follow the latest treatment and updated accounts of
Mucoromycota in Tedersoo et al. (2018) and Wijayawardene et al. (2018b).
Mucoromycetes Doweld
The classification of the families in Mucoromycetes
follows Tedersoo et al. (2018) and Wijayawardene et al.
(2018b).
Mucorales Fr.
123
Fungal Diversity
Fig. 171 Trechispora echinospora (13858Tell., MA-Fungi 82485,
holotype). a, b Basidiome dry specimen. c Apical hypha of aculeus.
d Basidia. e Spores. f, h Sphaerocysts (SEM). g Spores (SEM) of T.
123
echinospora (13861Tell., MA-Fungi 82486). i Spores (SEM). Scale
bars a = 1 cm, b = 5 mm, c–f = 5 lm, g, i = 2.5 lm, h = 10 lm
Fungal Diversity
Fig. 172 Phylogenetic tree of the Tremellochaete obtained by analyses of DNA sequences. The phylogenetic tree was generated by
partial concatenated analyses of the ITS and LSU sequence dataset.
Only species with at least the LSU sequences were considered for the
phylogeny. The sequences obtained in this study were aligned with
other from GenBank in MEGA6 (Tamura et al. 2013) and edited
using the Staden Package 2.0 software (Staden et al. 1998).
Sistotrema brinkmannii (Bres.) J. Erikss. was used as outgroup taxa.
Prior to phylogenetic analyses, the model of nucleotide substitution
was estimated using Topali 2.5 (Milne et al. 2004). Bayesian
inference analyses [two runs over 2 9 106 generations with a burn in
value of 25% and maximum likelihood (1000 bootstrap)] were
performed, respectively, in MrBayes 3.1.2 (Ronquist et al. 2012) and
PhyML (Guindon and Gascuel 2003) launched from Topali 2.5, using
the GTR ? G model. Sequences obtained in this study are in blue.
Support values are maximum likelihood (BSML) and Bayesian
posterior probability (BYPP). Only support values of at least 50% are
shown
Mucoraceae Dumort.
We follow the latest treatment and updated accounts of
Mucoraceae in Spatafora et al. (2016), Tibpromma et al.
(2017) and Wanasinghe et al. (2018). A high-level classification of Mucoraceae was updated by Tedersoo et al.
(2018) based on an evolutionary ecological analysis.
collected from the gardens of Chonnam National University, Gwangju, Korea, a new species of Mucor was isolated
and is described here. Recently, only three new Mucor
species have been reported in Korea: M. koreanus Hyang
B. Lee et al. from tangerine fruit (Li et al. 2016); M.
stercorarius Hyang B. Lee et al. from rat faeces (Tibpromma et al. 2017); and M. fluvii Hyang B. Lee et al. from
freshwater (Wanasinghe et al. 2018) (Fig. 177).
Mucor Fresen.
The genus Mucor was described by Fresenius (1850),
and comprises the largest number of species within the
Mucorales (Benny et al. 2014). They are readily isolated
from soil, fruits, vegetables, stored grains, insects or dung
(Benny 2008). Several species are of great interest to the
biotechnology industry because of their ability to produce
proteolytic enzymes (de Souza et al. 2015), while some
species are considered as the causal agents of cutaneous
zygomycosis in humans (Ribes et al. 2000). During a study
of Mucorales from fecal samples of praying mantis
Mucor orantomantidis Hyang B. Lee, P.M. Kirk & T.T.T.
Nguyen, sp. nov.
Index Fungorum number: IF555163; Facesoffungi
number: FoF05733, Fig. 178
Etymology: The specific epithet ‘‘orantomantidis’’ refers
to the origin of the fecal pellet of a praying mantis from
which the species was first isolated.
Holotype: CNUFC-MID1-1
123
Fungal Diversity
Fig. 173 Tremellochaete atlantica (URM 90199, holotype). a Basidiome when fresh. b Papillae encrusted with crystals. c Clamped
hyphae of the context. d, e 2-celled, metabasidia, hyphae with
ochraceus, granular content. f, g Basidiospores. Scale bars a = 1 cm,
b = 1 mm, c–g = 10 lm. Photos by R.L. Alvarenga
Fig. 174 UPGMA tree based on
Kimura-2-parameters of ITS
sequence dataset of seleceted
Dacrymycetales species from
Shirouzu et al. (2017).
Maximum parsimony and
maximum likelihood bootstrap
support values C 50% (10,000
replicates) and Bayesian
posterior probabilities C 0.95
BYPP are shown above the
nodes. The analyses were done
without rooting the tree. The
189-246 positions in the
alignment to D. invisibilis and
D. subantarcticensis are shown.
New species is indicated in blue
bold
Colonies on SMA reaching 38–41 mm diam. at 25 °C
after 3 days of incubation, yellowish; reverse yellow
brownish. Sporangiophores 6.5–16.5 lm wide, arising
123
from aerial hyphae, simple or sympodially branched, with
short or long branches. Sporangia globose to subglobose,
multispored, initially light yellow then turning brownish
Fungal Diversity
Fig. 175 Dacrymyces invisibilis (MA-Fungi 91306, holotype). a, b Appearance of basidiome on host surface. c Hyphae stained in congo red.
d Basidia stained in congo red. e Basidia. f–h Spores. Scale bars a, b = 2 mm, c = 5 lm, d, e = 10 lm, f–h = 5 lm
123
Fungal Diversity
Fig. 176 Dacrymyces invisibilis
(MA-Fungi 91306, holotype).
a Hyphae. b Basidia. c Spores.
Scale bars = 10 lm
yellow, 29.5–60.5 9 28.5–58.5 lm. Columellae globose to
subglobose, ellipsoidal, 25–58.5 9 22.5–45.5 lm. Sporangiospores ellipsoidal, sometimes one side slightly flattened, 8–11.5 9 4.5–6 lm. Chlamydospores abundantly
produced in aerial hyphae, in chains or irregular, globose,
fusiform, barrel-shaped, or irregular in shape. Zygospores
borne near the agar surface, reddish brown when young,
later blackish brown, 38.5–60.5 9 31.5–49 lm.
123
Culture characteristics: The isolate grew over a wide range
of temperatures with varying growth rates on SMA, PDA, and
MEA of 12.5, 14.5, and 12 mm per 24 h, respectively. Optimal growth was observed around 25–30 °C, slow growth was
observed below 10 °C, and no growth at 40 °C.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, garden of Chonnam National University located
in Gwangju (35° 090 16.9900 N, 126° 540 56.0200 E), from
fecal sample of praying mantis, October 2017 (CNUFC-
Fungal Diversity
Fig. 177 Phylogenetic tree of
Mucor orantomantidis
(CNUFC-MID1-1 and CNUFCMID1-2), and related species
based on a maximum likelihood
analysis of a combined ITS and
LSU sequence dataset. The
sequence of Backusella
oblongispora was used as
outgroup taxon. Numbers at the
nodes indicate the bootstrap
values ([ 50%) from 1000
replications. The bar indicates
the number of substitutions per
position. New taxa are shown in
blue and ex-type strains in bold
MID1-1, holotype); isotype in Korean Agricultural Culture
Collection (KACC, Wanju, Korea); ex-type living culture,
JMRC:SF:013605 (deposited at Jena Microbial Resource
Collection, University of Jena and Leibniz Institute for
Natural Product Research and Infection Biology, Jena,
Germany and preserved as glycerol stock at -80 °C in the
CNUFC).
GenBank number: ITS = MH594737, LSU = MH591457
(CNUFC-MID1-1); ITS = MH594738, LSU = MH591458
(CNUFC-MID1-2).
Notes: The phylogenetic analyses of a combined ITS
and LSU sequence dataset (Fig. 177) show that Mucor
orantomantidis belongs to M. amphibiorum group as
defined by Walther et al. (2013). Mucor orantomantidis is
closely related to Mucor sp. ‘hiemalis’. However, M.
orantomantidis differs from Mucor sp. ‘hiemalis’ by producing zygospores with equal or unequal suspensors, and
by having larger sporangiospores. M. orantomantidis
grows and sporulates at 35 °C, whereas, the maximum
temperature for Mucor sp. ‘hiemalis’ is around 30 °C. In
addition, the strain forms a separate branch from other
species of the genus, showing it represents a new species in
the phylogenetic tree (Fig. 177).
123
Fungal Diversity
Fig. 178 Mucor orantomantidis (CNUFC-MID1-1, holotype). a A
praying mantis. b Colony in potato dextrose agar. c Colony in
synthetic mucor agar. d Colony in malt extract agar. e, f
Sporangiophores and sporangia. g, h Columellae with collarette.
i Sporangiospores. j–m Zygospores. (e, g–k: LM; f, l, m: SEM).
Scale bars e–k = 20 lm, l, m = 50 lm
Acknowledgements We express sincere appreciations to the Key
Laboratory for Plant Diversity and Biogeography of East Asia,
Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan, China and the Centre of Excellence in Fungal
Research, Mae Fah Luang University, Chiang Rai, Thailand for
providing the facilities of taxonomic laboratory. 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. Prof. Dr. Shaun Pennycook is
thanked for his essential nomenclatural review. We gratefully thank
Prof. Dr. Jian-Kui Liu, Drs. Dimuthu S. Manamgoda, Dhanushka
Udayanga, Komsit Wisitrassameewong, Nalin N. Wijayawardene,
Philippe Callac and Miss Na Zhou for valuable suggestions and
general assistance. Rungtiwa Phookamsak thanks CAS President’s
International Fellowship Initiative (PIFI) for Young Staff 2019–2021
(grant number 2019FY0003), 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 research support.
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). Jianchu Xu thanks the Key Research Program of
Frontier Sciences of the Chinese Academy of Sciences (Grant No.
QYZDY-SSW-SMC014). Peter E Mortimer would like to thank the
National Science Foundation of China and the Chinese Academy of
Sciences for financial support under the following grants:
41761144055, 41771063 and Y4ZK111B01. Olivier Raspé is grateful
to the Fonds de la Recherche Scientifique-FNRS (Belgium) for travel
grants. 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). Dhanushaka N.
Wanasinghe would like to thank CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (Grant
No. 2019PC0008). E.B. Gareth Jones is supported under the Distinguished Scientist Fellowship Program (DSFP), King Saud University,
Kingdom of Saudi Arabia. Aniket Ghosh, Kanad Das, Priyanka
Uniyal, Rajendra P. Bhatt, Tahir Mehmood, and Upendra Singh are
grateful to the Head, Department of Botany & Microbiology & USIC
Dept. HNB Garhwal University, Srinagar Garhwal and thank to the
Director, Botanical Survey of India, Kolkata for providing all kinds of
facilities during the present study and UGC for providing fellowship
to Aniket Ghosh, Priyanka Uniyal and Tahir Mehmood. K. P. Deepna
Latha acknowledge the Kerala State Council for Science, Technology
123
Fungal Diversity
and Environment (KSCSTE) in the form of a PhD fellowship (Grant
No. 001/FSHP/2011/CSTE) and the Principal Chief Conservator of
forests, Kerala State, for granting permission (No. WL10- 4937/2012,
dated 03-10-2013) to collect agarics from the forests of Kerala. K. N.
Anil Raj thanks the Council of Scientific & Industrial Research
(CSIR), New Delhi, India, in the form of an award of CSIR Research
Associateship (09/043(0178) 2K17 dated: 31/03/2017). Mao-Qiang
He and Rui-Lin Zhao thank the National Natural Science Foundation
of China (Project ID: 31470152 and 31360014) and the Foundation of
Innovative Group of Edible Mushrooms Industry of Beijing (Project
ID:BAIC05-2017). Mingkwan Doilom would like to thank the 5th
batch of Postdoctoral Orientation Training Personnel in Yunnan
Province and the 64th batch of China Postdoctoral Science Foundation. Tatiana Baptista Gibertoni and Renato Lúcio Mendes Alvarenga
acknowledge Dr. Viacheslav Spirin for the taxonomic and morphological advises, CNPq for the Ph.D scholarship of RLMA (140283/
2016-1), Pós-Graduação em Biologia de Fungos (UFPE, Brazil),
Capes (Capes-SIU 008/13), CNPq (PQ 307601/2015-3) and FACEPE
(APQ 0375-2.03/15) for funding the research. Margarita Dueñas, M.
Teresa Telleria and Marı́a P. Martı́n acknowledge financial support
from the Agreement ENDESA and San Ignacio de Huinay Foundations and Consejo Superior de Investigaciones Cientı́ficas, CSIC
(Projects No. 2011HUIN10, 2013CL0012, 2014CL0011), the AECID
(Agencia Española de Cooperación Internacional para el Desarrollo)
and Plan Nacional I ? D?i project no. CGL2015–67459–P; to
Reinhard Fitzek (San Ignacio del Huinay Foundation, Chile) for his
invaluable help during fieldwork, and M. Glenn (Seton Hall University, US) for revising the text and Miguel Jerez and Yolanda Ruiz
León for SEM technical assistance. Sandra Nogal-Prata was supported by a Predoctoral Grant from the Ministerio de Economı́a y
Competitividad (Spain) (BES-2016-077793). Armin Mešić and
Zdenko Tkalčec thank Croatian Science Foundation for their partial
support under the project HRZZ-IP-2018-01-1736 (ForFungiDNA)
and are grateful to Milan Čerkez for collecting specimens of coprinoid taxa for this study. Hyang Burm Lee was supported by the
Graduate Program for the Undiscovered Taxa of Korea, and 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, and BK21 PLUS program funded by Ministry of Education,
Republic of Korea. Mubashar Raza would like to thank the CASTWAS for the PhD Fellowship. Sanjay K. Singh, Paras Nath Singh,
Shiwali Rana and Frank Kwekucher Ackah thank Director, MACS,
Agharkar Research Institute, Pune, India for providing facilities.
Shiwali Rana and Frank Kwekucher Ackah thank UGC (Junior
Research Fellowship) and DST, Govt. of India (CV Raman Fellowship for African Researchers), respectively. Gen-Nuo Wang, Huang
Zhang, Wei Dong and Xian-Dong Yu thank the National Natural
Science Foundation of China (Project ID: NSF 31500017). Bandarupalli Devadatha and V. Venkateswara Sarma thank The Ministry
of Earth sciences, Govt. of India (Sanction order: MOES/36/OO1S/
Extra/40/2014/PC-IV dt.14.1.2015) for a funding of the project, T,
District Forest Office, Tiruvarur, Tamil Nadu and PCCF (Head of
Forest Force), Chennai, Tamil Nadu Forest Department for providing
permission to collect samples from Muthupet mangroves, and
Department of Biotechnology, Pondicherry University is thanked for
providing the facilities. Myung Soo Park, Seung-Yoon Oh and Young
Woon Lim thank the Marine Bio Resource Bank Program of the
Ministry of Ocean & Fisheries, Korea. Olinto Pereira thanks the
CAPES, CNPq and FAPEMIG for financial support. Neven Matočec,
Ivana Kušan and Margita Jadan express their gratitude to Livio
Lorenzon, Enrico Bizio and Raffaella Trabucco (MCVE) for their
kind help with loan of Sarcopeziza sicula type material; parts of their
research were financed by Public Institutions Sjeverni Velebit
National Park and Paklenica National Park. Dong-Qin Dai and LiZhou Tang thank the National Natural Science Foundation of China
(No. NSFC 31760013, NSFC 31260087, NSFC 31460561), the Scientific Research Foundation of Yunnan Provincial Department of
Education (2017ZZX186) and utilization of endophytes and the
Thousand Talents Plan, Youth Project of Yunnan Provinces. Ting-Chi
Wen, Chada Norphanphoun and Shi-Ke Huang are grateful to the
National Natural Science Foundation of China (No. 31760014) and
the Science and Technology Foundation of Guizhou Province (No.
[2017]5788). Monika C. Dayarathne thanks to Thailand Research
Fund (TRF) Grant No MRG6080089 for financial research support.
Napalai Chaiwan thanks The Royal Golden Jubilee Ph. D. Program
(PHD60K0147) under Thailand Research Fund, for financial research
supports on project entitle ‘‘Fungi on limestone outcrops from
southern Thailand to lower himalyas’’. Saranyaphat Boonmee and
Sirinapa Konta would like to thank the National Research Council of
Thailand (Grant No. 61215320023, 61215320013) and the Thailand
Research Fund (Grant No. TRG6180001) for research financial support. Saisamorn Lumyong would like to thank the Thailand Research
Fund (RTA 5880006) and Chiang Mai University for partially support
this research work. Itthyakorn Promputtha would like to thank Chiang
Mai University for partially supported this research work. Lei Cai
acknowledges China-Thailand Joint Lab on Microbial Biotechnology
(Most KY201701011) for financial support. PhD students from Mae
Fah Laung and Chiang Mai Universities thank the Mushroom
Research Foundation for research financial support and PhD
Fellowships.
Open Access This article is distributed under the terms of the Creative
Commons Attribution 4.0 International License (http://creative
commons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give
appropriate credit to the original author(s) and the source, provide a
link to the Creative Commons license, and indicate if changes were
made.
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Dictyosporium, new combinations in Dictyocheirospora and an
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Affiliations
Rungtiwa Phookamsak1,2,3,4,6 • Kevin D. Hyde1,3,4,5 • Rajesh Jeewon7 • D. Jayarama Bhat8 • E. B. Gareth Jones17,48 •
Sajeewa S. N. Maharachchikumbura9 • Olivier Raspé10,11 •
Samantha C. Karunarathna1,2,4 • Dhanushka N. Wanasinghe1,2,3,4 • Sinang Hongsanan3,12 • Mingkwan Doilom1,2,3,4,6 •
Danushka S. Tennakoon1,3,5,13 • Alexandre R. Machado14 • André L. Firmino15 • Aniket Ghosh16 •
Anuruddha Karunarathna1,3,13,17 • Armin Mešić18 • Arun Kumar Dutta19 • Benjarong Thongbai20 •
Bandarupalli Devadatha21 • Chada Norphanphoun3,5,13,22 • Chanokned Senwanna3,5,17 • Deping Wei1,3,5,17 •
Dhandevi Pem3,5,12 • Frank Kwekucher Ackah23 • Gen-Nuo Wang24 • Hong-Bo Jiang1,3,5 • Hugo Madrid25 •
Hyang Burm Lee26 • Ishani D. Goonasekara1,3,5 • Ishara S. Manawasinghe3,27 • Ivana Kušan18 •
Josep Cano28 • Josepa Gené28 • Junfu Li1,3 • Kanad Das29 • Krishnendu Acharya19 • K. N. Anil Raj30 •
K. P. Deepna Latha30 • K. W. Thilini Chethana3,27 • Mao-Qiang He31 • Margarita Dueñas32 • Margita Jadan18 •
Marı́a P. Martı́n32 • Milan C. Samarakoon3,6,33 • Monika C. Dayarathne1,3,5 • Mubashar Raza31,34 •
Myung Soo Park35 • M. Teresa Telleria32 • Napalai Chaiwan1,3,5 • Neven Matočec18 • Nimali I. de Silva1,3,5,6 •
Olinto L. Pereira36 • Paras Nath Singh37 • Patinjareveettil Manimohan30 • Priyanka Uniyal16 • Qiu-Ju Shang3,33 •
Rajendra P. Bhatt16 • Rekhani H. Perera3,5,33 • Renato Lúcio Mendes Alvarenga38 • Sandra Nogal-Prata32 •
Sanjay K. Singh37 • Santhiti Vadthanarat6 • Seung-Yoon Oh35 • Shi-Ke Huang1,3,5,22 • Shiwali Rana37 •
Sirinapa Konta1,3,5 • Soumitra Paloi19 • Subashini C. Jayasiri1,3,5 • Sun Jeong Jeon26 • Tahir Mehmood16 •
Tatiana Baptista Gibertoni38 • Thuong T. T. Nguyen26 • Upendra Singh16 • Vinodhini Thiyagaraja1,3,5,17 •
V. Venkateswara Sarma21 • Wei Dong3,5,17,39 • Xian-Dong Yu39 • Yong-Zhong Lu3,5,22 • Young Woon Lim35 •
Yun Chen40 • Zdenko Tkalčec18 • Zhi-Feng Zhang31,34 • Zong-Long Luo3,5,41 • Dinushani A. Daranagama42 •
Kasun M. Thambugala52 • Saowaluck Tibpromma1,2,3,4 • Erio Camporesi43,44,45 • Timur S. Bulgakov46 •
Asha J. Dissanayake3 • Indunil C. Senanayake3,12 • Dong Qin Dai47 • Li-Zhou Tang47 • Sehroon Khan1,4 •
Huang Zhang39 • Itthayakorn Promputtha6,50 • Lei Cai31 • Putarak Chomnunti3,49 • Rui-Lin Zhao31 •
Saisamorn Lumyong6,51 • Saranyaphat Boonmee3 • Ting-Chi Wen22 • Peter E. Mortimer1 • Jianchu Xu2,4
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
12
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Science and Oceanography, Shenzhen
University, 3688, Nanhai Avenue, Nanshan,
Shenzhen 518060, People’s Republic of China
2
Institute of Animal Science, State Key Laboratory of Animal
Nutrition, Chinese Academy of Agricultural Sciences,
Beijing 100193, People’s Republic of China
13
Department of Plant Medicine, National Chiayi University,
300 Syuefu Road, Chiayi City 60004, Taiwan
14
3
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
Departamento de Micologia, Universidade Federal de
Pernambuco, Recife, Pernambuco, Brazil
15
4
World Agroforestry Centre, East and Central Asia,
Kunming 650201, Yunnan, People’s Republic of China
Instituto de Ciências Agrárias, Universidade Federal de
Uberlândia, Monte Carmelo, Minas Gerais, Brazil
16
5
Mushroom Research Foundation, 128 M.3 Ban Pa Deng T. Pa
Pae, A. Mae Taeng, Chiang Rai 50150, Thailand
Department of Botany & Microbiology, H.N.B. Garhwal
University, Srinagar, Garhwal 246174, Uttarakhand, India
17
Department of Entomology and Plant Pathology, Faculty of
Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
6
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
7
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit, Mauritius
18
Rud̄er Bošković Institute, Bijenička cesta 54, 10000 Zagreb,
Croatia
8
No. 128/1-J, Co-Op Azad Housing Society, Curca,
Goa Velha, India
19
9
Department of Crop Sciences, College of Agricultural and
Marine Sciences, Sultan Qaboos University, PO Box 8,
123 Al Khoud, Oman
Molecular and Applied Mycology and Plant Pathology
Laboratory, Department of Botany, University of Calcutta,
Kolkata, West Bengal 700019, India
20
Department Microbial Drugs, Helmholtz Center for Infection
Research, Inhoffenstrasse 7, 38124 Brunswick, Germany
21
Fungal Biotechnology Laboratory, Department of
Biotechnology, Pondicherry University,
Kalapet 605014, Pondicherr, India
22
Engineering Research Center of Southwest BioPharmaceutical Resources, Ministry of Education, Guizhou
University, Guiyang 550025, People’s Republic of China
10
Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium
11
Fédération Wallonie-Bruxelles, Service général de
l’Enseignement universitaire et de la Recherche scientifique,
Rue A. Lavallée 1, 1080 Brussels, Belgium
123
Fungal Diversity
23
Department of Crop Science, School of Agriculture, College
of Agriculture and Natural Sciences, University of Cape
Coast, Cape Coast, Ghana
24
Faculty of Environmental Science & Engineering, Kunming
University of Science & Technology, Kunming 650500,
People’s Republic of China
25
Centro de Genómica y Bioinformática, Facultad de Ciencias,
Universidad Mayor, Camino La Pirámide 5750, Huechuraba,
Santiago, Chile
26
Environmental Microbiology Lab, Division of Food
Technology, Biotechnology & Agrochemistry, College of
Agriculture and Life Sciences, Chonnam National University,
Gwangju 61186, Republic of Korea
27
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences,
Beijing 100097, People’s Republic of China
28
29
30
31
Mycology Unit, Medical School and IISPV, Universitat
Rovira i Virgili, 43201 Reus, Spain
Cryptogamic Unit, Botanical Survey of India, P. O. Botanic
Garden, Howrah 711103, India
Department of Botany, University of Calicut, Kerala 673 635,
India
State Key Laboratory of Mycology, Institute of
Microbiology, Chinese Academy of Sciences,
Beijing 100101, People’s Republic of China
38
Departamento de Micologia, Centro de Biociências (CB),
Departamento de Micologia, Universidade Federal de
Pernambuco, Avenida da Engenharia, S/N 50740-600 –
Cidade Universitária, Recife, Pernambuco, Brazil
39
Yunnan Institute of Food Safety, Kunming University of
Science and Technology, Kunming 650500, People’s
Republic of China
40
Law Enforcement of Agricultural Bureau, Xiuwen district,
Guiyang City 550200, Guizhou, People’s Republic of China
41
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, Yunnan, People’s Republic of
China
42
Department of Botany, University of Kelaniya, Kelaniya, Sri
Lanka
43
A.M.B, Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314,
Brescia, Italy
44
A.M.B. Gruppo, Micologico Forlivese ‘‘Antonio
Cicognani’’, Via Roma 18, Forlı́, Italy
45
Società per gli Studi Naturalistici della Romagna,
C.P. 143, Bagnacavallo, RA, Italy
46
Russian Research Institute of Floriculture and Subtropical
Crops, Yana Fabritsiusa Street 2/28, Sochi, Krasnodar region,
Russia 354002
47
Center for Yunnan Plateau Biological Resources Protection
and Utilization, College of Biological Resource and Food
Engineering, Qujing Normal University,
Qujing 655011, Yunnan, People’s Republic of China
32
Department of Mycology, Real Jardı́n Botánico-CSIC, Plaza
de Murillo 2, 28014 Madrid, Spain
33
Guizhou Key Laboratory of Agricultural Biotechnology,
Guizhou Academy of Agricultural Sciences,
Guiyang 550006, Guizhou, People’s Republic of China
48
Department of Botany and Microbiology, College of Science,
King Saud University, P.O Box 2455, Riyadh 11451,
Kingdom of Saudi Arabia
34
University of Chinese Academy of Sciences, Beijing 100049,
People’s Republic of China
49
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
35
School of Biological Sciences and Institute of Microbiology,
Seoul National University, Seoul 08826, Republic of Korea
50
36
Departamento Fitopatologia, Universidade Federal de
Viçosa, Viçosa, Minas Gerais, Brazil
Center of Excellence in Bioresources for Agriculture,
Industry and Medicine, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
51
Center of Excellence in Microbial Diversity and Sustainable
Utilization, Faculty of Science, Chiang Mai University,
Chiang Mai 50200, Thailand
52
Faculty of Science and Technology, Thammasat University,
Rangsit Campus, Klong Luang, Pathumthani 12121, Thailand
37
Biodiversity and Palaeobiology Group, National Fungal
Culture Collection of India, MACS Agharkar Research
Institute, G.G. Agarkar Road, Pune 411 004, India
123