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 sexual-asexual 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 camelliae-yunnanensis, 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, 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.
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.
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.
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
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.
Diaporthales Nannf.
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.
Pseudoplagiostomataceae Cheew. et al.
987.Pseudoplagiostoma mangiferae Dayarathne, Phookamsak & K.D. Hyde, sp. nov.
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.
991.Proliferophorum thailandicum G.N. Wang, H. Zhang & Senan., sp. nov.
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 leptobactrumvar.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.
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.
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 Gӓum.
Phanerochaetaceae Jülich
1026.Phanerochaete australosanguinea Telleria, M. Dueñas & M.P. Martín, 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. Martín, sp. nov.
Trechisporales K.H. Larss.
Hydnodontaceae Jülich
1032.Trechispora echinospora Telleria, M. Dueñas, I. Melo & M.P. Martín, 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. Martín, sp. nov.
Mucoromycota Doweld
Mucoromycetes Doweld
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, 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 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.
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)
Hyweljonesia indica P.N. Singh & S.K. Singh, sp. nov.
MycoBank number: MB821804; Facesoffungi number: FoF03526, 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 barsd, f–j = 10 µm
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 µm wide. Conidiophores arising from loose to compact hyphal tufts, macronematous, lateral, unbranched to rarely branched at base, 0–1-septate, straight, smooth-walled, light olivaceous, 4.5–24.5 µm long (\( \bar{x} \) = 9.6 µm, n = 30); base flared, 3.5–8.5 µm wide (\( \bar{x} \) = 5.87 µm, n = 30); apex narrow, cylindrical, 1.5–4.5 µm wide (\( \bar{x} \) = 2.74 µm, n = 30). Conidiogenous cells terminal, 1(–2), straight, smooth-walled, subhyaline to olivaceous, cylindrical to clavate, scars inconspicuous, 5.5–12.8 × 2–5 μm (\( \bar{x} \) = 10.3 × 3.28 µm, n = 30). Conidia acrogenous to rarely acropleurogenous, produced in apical whorl of 1–12 conidia, simple, aseptate, obovoid to pyriform, smooth-walled, hyaline, apex rounded, base truncate, hilum refractive, 2.4–6.8 × 1.5–2.6 µm (\( \bar{x} \) = 4.6 × 2 µm, 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 × 132–220 µm. 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 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
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.
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 μm diam. (\( \bar{x} \) = 130 μm, n = 10), velvety, greyish 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 × 3–5 μm (\( \bar{x} \) = 5.2 × 4.6 μm, n = 10), holoblastic, integrated, terminal, determinate, pale brown, smooth-walled. Conidia 45–69 × 15–29 μm (\( \bar{x} \) = 61 × 20 μm, 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 × 5–7 μm (\( \bar{x} \) = 47 × 6 μm, n = 10) (Fig. 3).
Maximum likelihood majority rule consensus tree for the analysed Dictyosporiaceae isolates based on a dataset of combined ITS, LSU and TEF1-α 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
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), ex-type living culture, MFLUCC 15-0282.
GenBank numbers: ITS = MH742321, LSU = MH742313, SSU = MH742317, (MFLUCC 15-0282a); ITS = MH742322, LSU = MH742314, SSU = MH742318, TEF1-α = MH764301 (MFLUCC 15-0282b); ITS = MH742323, LSU = MH742315, SSU = MH742319, TEF1-α = MH764302 (MFLUCC 15-0282c); ITS = MH742324, LSU = MH742316, SSU = MH742320, TEF1-α = 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-α 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).
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 barsa = 500 μm, b, c = 100 μm, g–p = 20 μm, d–f = 10 μm
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.
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 morphAscomata 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.
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
Vicosamyces venturisporus Firmino, A.R. Machado & O.L. Pereira, sp. nov.
MycoBank number: MB822578; Facesoffungi number: FoF03787, 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. e Mature ascus. f Immature ascospore. g Brown and smooth-walled ascospores. Scale barsc = 50 μm, d–g = 10 μm
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 morphAscomata 240–340 × 250–310 μm, 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 μm wide, numerous, cylindrical, filiform, septate, unbranched, hyaline pseudoparaphyses. Asci 125–152.5 × 14–15 µm, 8-spored, bitunicate, fissitunicate, cylindrical, subsessile to short pedicellate, with furcate pedicel, apically rounded with well-developed ocular chamber. Ascospores 22.5–30 × 6–8 µm, 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: ITS = MF802825, LSU = MF802828 (CDA1494); ITS = MF802826, LSU = MF802829 (CDA1495); ITS = MF802827, LSU = MF802830 (CDA495).
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 TEF1-α 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).
Phylogram generated from maximum likelihood analysis based on the combined LSU, SSU, ITS and TEF1-α 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
Keissleriella caraganae Chaiwan, Phookamsak, Wanas. & K.D. Hyde, sp. nov.
Index Fungorum number: IF555523; Facesoffungi number: FoF04965, 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 Ascospores. m Ascospores stained in Indian ink. n, o Culture on PDA after one week (n = from above, o = from below). Scale barsa = 200 µm, b, c = 50 μm, d–i = 20 μm, j–m = 5 μm
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 morphAscomata 140–175 μm high, 170–235 μm diam., scattered, solitary or in groups, semi-immersed, visible as raised, black dots on host surface, globose to subglobose, glabrous, ostiolate at centre, with minute papilla, filled with short, brown, aseptate periphyses. Peridium 15–25 μm 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 μm wide, broad filamentous, distinctly septate, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci 39–75 × 10–12 μm (\( \bar{x} \) = 60.1 × 11.1 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores 14–20 × 3–7 μm (\( \bar{x} \) = 16.9 × 5.1 μm, 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, 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-α = MK214377 (KUMCC 18-0163); ITS = MK359434, LSU = MK359439, SSU = MK359444 TEF1-α = 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 well-resolved 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
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 barsa = 200 µm, b = 50 μm, c, l = 20 μm, d–k = 10 μm
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 morphConidiomata 95–150 μm high, 110–170 μm 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 μm 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 × (3–)4–8 µm (\( \bar{x} \) = 11 × 5.5 μm, 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) × 7–10(–12) µm (\( \bar{x} \) = 15.6 × 9.4 μm, 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, smooth-walled 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-α = MK214379 (KUMCC 18-0167); ITS = MK359436, LSU = MK359441, SSU = MK359446, TEF1-α = MK359075 (KUMCC 18-0168); ITS = MK359437, LSU = MK359442, SSU = MK359447, TEF1-α = MK359076 (KUMCC 18-0169); ITS = MK359438, LSU = MK359443, SSU = MK359448, TEF1-α = 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, 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. 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
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 barsc = 50 µm, d, h = 10 μm, e–g, i–l = 5 μm
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.
Associated with leaf spots on Cycas (Cycadaceae). Sexual morph Undetermined. Asexual morphConidiomata 130–170 μm high, 180–270 μm 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 μm, 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) × (1.5–)2–5(–7) µm (\( \bar{x} \) = 4.3 × 3.7 μm, n = 30), holoblastic, phialidic, hyaline, smooth, aseptate, ampulliform, arising from the inner cavity of the conidioma wall. Conidia (8–)10–13(–14) × 3–4 µm (\( \bar{x} \) = 12 × 4.2 μm, 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.
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-α = MK214378 (KUMCC 18-0165); ITS = MK359434, LSU = MK359439, SSU = MK359444, TEF1-α = 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 × 1.5–2.5 μm in the type collection) and longer conidia (9–11 × 3–4.5 µm in the type collection). Phragmocamarosporium platani has smaller conidiogenous cells (1.5–3 × 1.5–2.5 μm) and narrower conidia (12–13 × 5–7.5 μm) (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) re-circumscribed 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
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 barsa, b = 500 µm, c = 200 μm, d = 100 μm, e, g–i = 50 μm, j–m = 10 μm, f = 5 μm
Etymology: Name reflects the host from which the fungus was isolated.
Holotype: MFLU 18-0591
Saprobic on dead branches of Urtica dioica. Sexual morphAscomata 100–130 high, 70–110 μm 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 μm diam., papillate, black, shiny, smooth. Peridium 25–50 μm 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 relatively thin-walled cells of textura angularis, cells near the base comparatively larger. Hamathecium comprising numerous, dense, 1.5–2 μm wide, filamentous, septate, cellular pseudoparaphyses, branched and anastomosing, embedded in gelatinous matrix. Asci 60–140 × 9–11 μm (\( \bar{x} \) = 104.5 × 10 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, rounded at the apex, pedicellate, numerous, with ocular chamber. Ascospores 35–40 × 4–6 μm (\( \bar{x} \) = 38.3 × 5.2 μm, 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-α = 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 × 9–11 μm versus 60–112 × 7–12 μm, L. italica), longer ascospores (L. urticae, 35–40 × 4–6 μm versus 12–18 × 4–6 μm, 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.
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 glauco-punctata (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
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
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 barsa = 500 μm, b, c = 100 µm, d, e = 50 µm, f–i, n = 20 µm, j–m = 10 µm
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 morphAscomata 140–280 μm high, 300–450 μm diam., black, shiny on the host surface, solitary to aggregated, immersed at the base, becoming superficial, uni-loculate, 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 μm 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 μm wide, distinctly septate pseudoparaphyses, anastomosing, embedded in a hyaline gelatinous matrix. Asci (64–)70–90(–100) × (9.5–)10–13 μm (\( \bar{x} \) = 82.7 × 11.2 μm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical, short, pedicellate, apically rounded with a distinct ocular chamber. Ascospores (28–)30–40 × (4.5–)5–6 μm (\( \bar{x} \) = 34.4 × 5.5 μm, 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). ex-type living culture, KUMCC 18-0151.
GenBank numbers: ITS = MK387920, LSU = MK387958, SSU = MK387928, TEF1-α = 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 × 3.1–4.3 μm), 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 × 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.
Plenodomus lijiangensis Phookamsak, A. Karunarathna & K.D. Hyde, sp. nov.
Index Fungorum number: IF556137; Facesoffungi number: FoF05697, 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 barsa = 200 μm, b, n, o = 100 µm, c, p = 50 µm, u = 10 µm, g, q–t = 5 µm, h, i = 2 µm
Etymology: The specific epithet “lijiangensis” refers to Lijiang prefecture-level city, of Yunnan Province, China where the holotype was collected.
Holotype: KUN-HKAS 102249
Saprobic on dead fronds of fern. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata 140–290 µm high, 135–240 µm 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 µm 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 × 5–8 µm (\( \bar{x} \) = 6.4 × 6.5 μm, 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 × 1.7–2.3 µm (\( \bar{x} \) = 4.3 × 2 µm, 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 µm high, 130–230 µm diam., scattered, solitary to gregarious, semi-immersed in culture colony, or embedded in agar medium, perithecial, pycnidial, with short stipe (19–49 µm long), black, glabrous, globose to subglobose, lacking ostioles. Conidiomata walls 3–8 µm 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 × 1.5–3 µm (\( \bar{x} \) = 2.5 × 2.2 μm, 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) × 1.5–2.5 µm (\( \bar{x} \) = 3.9 × 2, n = 50), hyaline, oblong to ellipsoidal, or obovoid, aseptate, smooth-walled, 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), ex-type living culture, KUMCC 18-0186.
GenBank numbers: ITS = MK387921, LSU = MK387959, SSU = MK387929, TEF1-α = 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
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 barsa = 200 µm, b = 100 µm, c = 50 µm, d, i–k = 20 µm, e–h = 10 µm
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 barsa = 500 µm, c = 50 µm, b, d = 20 µm, e, f, j = 10 µm, g–i = 5 µm, k–n = 2 µm
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 morphAscomata 250–290 µm high, 300–360 µm 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 µm wide, comprising 2–3 strata of flattened, pale brown, thin-walled, pseudoparenchymatous cells, arranged in textura angularis to textura prismatica, middle layer 25–100 µm 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 µm wide, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (75–)80–95(–107) × (8.5–)9–11(–12) µm (\( \bar{x} \) = 88.1 × 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 30–39 × 4–6(–6.5) µm (\( \bar{x} \) = 34.1 × 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 µm high, 50–110 µm diam., pycnidial, black, shiny, scattered, gregarious, superficial, uni-loculate, globose to subglobose, with short stipe (5–10 × 7–11 µm), glabrous, ostiole central, with pore-like opening, apapillate. Conidiomata walls 5–10 µm 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) × 4–6(–8) µm (\( \bar{x} \) = 5.6 × 5.1 μm, 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 × 1–2 µm (\( \bar{x} \) = 3.8 × 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-α = MK435602, RPB2 = MK435608 (KUMCC 18-0153); ITS = MK387923, LSU = MK387961, SSU = MK387931, TEF1-α = 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 (MFLU17-0757) 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, 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 thick-walled 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
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 barsa = 500 µm, b = 100 µm, c = 50 µm, d–n = 20 µm
Etymology: The specific epithet “isthmospora” refers to the fungus having isthmospores.
Holotype: KUN-HKAS 102225
Saprobic on dead branches of herbaceous plant. Sexual morphAscomata 230–330 μm high, 260–510 μm diam., 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 μm 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 µm wide, hyaline, filamentous distinctly septate pseudoparaphyses, laterally becoming, 6–18 µm wide, broadly cellular, hyaline, septate catenophyses, deeply constricted at the septa. Asci (120–)130–150(–165) × (15–)19–23(–28) µm (\( \bar{x} \) = 141.4 × 21.6 µm, 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) × 4–7 µm (\( \bar{x} \) = 87.1 × 5.9 µm, 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) µm long, 5–7-septate, cylindrical, with rounded end; lower part 28–35(–50) µm 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-α = MK435604 (HKAS 102225A); ITS = MK387926, LSU = MK387964, SSU = MK387934, TEF1-α = 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.
Sphaerellopsis paraphysata Crous & Alfenas, in Trakunyingcharoen et al., IMA Fungus 5(2): 411 (2014)
Facesoffungi number: FoF04968, Fig. 18
Sphaerellopsis paraphysata (KUN-HKAS 101483). aLiriope 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 barsj = 100 µm, c, d, f = 50 µm, k = 20 µm, e, l, m = 10 µm, n = 5 µm, o–r = 2 µm
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 µm high, 90–150 µm 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 µm wide, composed of 2–5 layers, of dark brown pseudoparenchymatous cells of textura angularis. Conidiophores 15–21 × 2.4–3 µm (\( \bar{x} \) = 18 × 2.7 µm, n = 20), arising from the basal cavity, 1–2-celled, hyaline, curved, cylindrical, or reduced to conidiogenous cells. Conidiogenous cells 3.5–6 × 2.5–4.5 µm (\( \bar{x} \) = 4.7 × 3.5 µm, 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 × 3–5 µm (\( \bar{x} \) = 15.5 × 4.5 µm, 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-α = MK435606.
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 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.
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 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 mucilaginosis M. Raza & L. Cai, sp. nov.
Index Fungorum number: IF555333; Facesoffungi number: FoF04941, 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 barsb = 100 μm, c = 50 μm, d, p, q = 20 μm, e–i = 10 μm, o = 5, j–n = 2 μm
Etymology: In reference to the mucilaginous sheath around spores.
Holotype: HMAS 255437
Saprobic on dead wood. Sexual morphAscomata 220–340 μm high, 240–400 μm 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 × 25–60 μm, apically with crest-like papilla, filled with periphyses, carbonaceous, with beak-like opening. Peridium 55–80 μm 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 μm wide, filamentous, indistinct septate, anastomosed pseudoparaphyses, embedded in a hyaline, gelatinous matrix. Asci (83.5–)102–144(–210) × (10.5–)13–15(–21.5) μm (\( \bar{x} \) = 127.5 × 14.5 μm, 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) × (5.5–)6–11(–12.5) μm (\( \bar{x} \) = 39.1 × 8.6 μm, n = 40), overlapping 2-seriate, hyaline, subfusoid to fusiform, with rounded or obtuse ends, 1(–3)-septate, smooth-walled, guttulate when young, with an entire mucilaginous sheath (9–20.5 μm 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-α = MH822893.
Notes: Multi-loci phylogenetic analyses based on a concatenated ITS, LSU, SSU, TEF1-α and RPB2 sequence dataset show that Lophiotrema mucilaginosis forms a well-supported 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 from Yunnan and Guizhou Provinces in China respectively. Lophiotrema mucilaginosis can be distinguished from L. hydei in having larger asci (78–89(–99) × 6.9–8.8 μm, L. hydei) and larger ascospores (23–28 × 3–4 μm, L. hydei) (Zhang et al. 2018). In addition, L. mucilaginosis has conical to mammiform ascomata, 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 × 7–10 μm, L. neohysterioides) and larger ascospores (17–25 × 3–5 μm, L. neohysterioides) and presence of mucilaginous sheath surrounding the ascospores, a feature not observed in L. neohysterioides (Tanaka and Harada 2003a).
Phylogenetic tree generated from maximum likelihood analysis (RAxML) based on a combined ITS, LSU, SSU, TEF1-α 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)
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).
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 morphAscomata 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, 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 slit-like openings (Zhang et al. 2009b; Hashimoto et al. 2017).
Maximum likelihood phylogenetic tree generated from analysis of a combined LSU, SSU, ITS, RPB2 and TEF1-α 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
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 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.
Brunneofusispora sinensis S.K. Huang & K.D. Hyde, sp. nov.
Index Fungorum number: IF555600; Facesoffungi number: FoF04863, 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 barsc = 500 µm, d = 200 µm, f = 50 µm, g–i = 20 µm, e, j–m = 10 µm
Etymology: The specific epithet “sinensis” refers to the country, China, where the taxon was collected.
Holotype: KUN-HKAS 97451.
Saprobic on dead wood. Sexual morphAscomata 325–370 μm 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 μm wide, equally thick-walled, composed of 5–8 strata, of blackened pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, 1.5–3.5 μm wide, filamentous, septate pseudoparaphyses, embedded in a gelatinous matrix. Asci 53–110 × 9–18 μm (\( \bar{x} \) = 75 × 14 μm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate to clavate, short pedicellate, rounded at the apex, with an ocular chamber. Ascospores 18–22 × 5–8.5 μm (\( \bar{x} \) = 20 × 7 μm, 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-α = 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.
Simplified phylogram showing the best RAxML tree obtained from a combined multigene (SSU, ITS, LSU and TEF1-α) 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 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 morphAscomata 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 cylindric-clavate, 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-α 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-α). However, we could not compare the morphological characteristics of Lonicericola with Pseudomonodictys and Paratrimmatostroma as they are represented by different morphs.
Lonicericola hyaloseptispora Phookamsak, Jayasiri & K.D. Hyde, sp. nov.
Index Fungorum number: IF556140; Facesoffungi number: FoF04963, 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 barsa, b = 500 µm, c = 100 µm, d, e = 50 µm, f–m = 20 µm
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 morphAscomata 170–240 µm high, 165–250 µm 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 pore-like opening, filled with hyaline periphyses. Peridium 8–25 µm 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 µm wide, filamentous, septate, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (90–)100–120(–145) × (24–)25–30(–33) μm (\( \bar{x} \) = 115.1 × 27 μm, 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 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 × 9–12 μm, l/w), (8–)9-septate, constricted at the septa, smooth-walled, with small to large guttules, surrounded by entire mucilaginous sheath (3.5–13 μm 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-α = 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 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
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 immersed in hyaline, cellular pseudoparaphyses, stained in Indian ink. f Ascus. g–i Ascospores. j, k Ascospores stained in Indian ink. Scale barsa = 500 µm, b = 100 µm, c, d = 50 µm, e–k = 20 µm
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 morphAscomata 130–170 µm high, 430–600 µm 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 µm 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 µm wide, filamentous, septate pseudoparaphyses, anastomosing above the asci, embedded in a hyaline gelatinous matrix. Asci (50–)80–120 × (10–)25–33 µm (\( \bar{x} \) = 107 × 28 µm, 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 × 7.5–11 µm (\( \bar{x} \) = 46.5 × 9 µm, 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), ex-type living culture, KUMCC 18-0147 (IS003A), KUMCC 18-0148 (IS003B).
GenBank numbers: ITS = MK098190, LSU = MK098199, SSU = MK098205, TEF1-α = MK098209 (KUMCC 18-0147); ITS = MK098193, LSU = MK098198, SSU = MK098202, TEF1-α = 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 × 150–300 µm in P. bambusina), absence of a beak-like structure and wider peridium (10–20 µm in P. bambusina). Ascospores of P. thysanolaenae are 5-septate and shorter than the 3–5-septate ascospores of P. bambusina (54.7 × 8.4 μm, 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 thick-walled.
Type species: Paratrimmatostroma kunmingensis Jayasiri, Phookamsak, D.J. Bhat & K.D. Hyde
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 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-α 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-α). 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
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 barsa = 200 µm, b = 50 µm, d–q = 10 µm
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) × 2–4 µm (\( \bar{x} \) = 19.2 × 3.2 µm, 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) × 2–4.5 µm (\( \bar{x} \) = 7.1 × 3.1 µm, 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) × (6–)10–20 µm (\( \bar{x} \) = 16.8 × 13.7 µm, n = 40)], cylindrical to sigmoid, or reniform [(8.5–)15–50 × 6–8(–10) µm (\( \bar{x} \) = 29.7 × 7.7 µm, n = 50)], solitary, tapering near apex and base, rounded at tip, multi-septate, 6–13-septate 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 (KUN-HKAS 102224, holotype).
GenBank numbers: ITS = MK098192, LSU = MK098196, SSU = MK098204, TEF1-α = 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).
Phylogram generated from the best scoring of the RAxML tree based on combined ITS, LSU and TEF1-α 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
Periconia cortaderiae Thambug. & K.D. Hyde, in Thambugala et al., Mycosphere 8(4): 734 (2017), 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 barsb, c = 50 µm, d–f = 20 µm, g, j = 10 µm, h, i = 5 µm
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 morphColonies on the substratum superficial, numerous, effuse, black, floccose. Mycelium immersed on the substrate, composed of septate, branched, smooth, dark hyphae. Conidiophores 100–225 × 8.5–11 μm, 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 × 4.1–5.6 μm (\( \bar{x} \) = 4.9 × 5.1 μm, n = 20), mono- to polyblastic, discrete, terminal, globose, light brown. Conidia (5.5–)6–9(–12) × (4.5–)5–8 μm (\( \bar{x} \) = 7.8 × 6.4 μm, 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).
GenBank numbers: ITS = MH892348, LSU = MH892401 (MFLUCC 18-0667); ITS = MH892349, LSU = MH892402, TEF1-α = MH908964 (MFLUCC18-0668).
Notes: Phylogenetic analyses of a concatenated LSU, ITS and TEF1-α sequence datset show that our strains (MFLUCC 18-0667 and MFLUCC 18-0668) form a well-resolved clade, clustering with Periconia cortaderiae (100% ML and 1.00 BYPP; Fig. 27). A comparison of ITS and TEF1-α 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-α). 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 μm high, holotype) and slightly larger conidia (4–6.6 × 4.1–7.1 μm, holotype) (Thambugala et al. 2017). Periconia species are morphologically variable in different hosts and geographic distributions, but the molecular data of informative genes can 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. (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 previous phylogenies derived from Hyde et al. (2018b) and Wanasinghe et al. (2018).
Phylogram generated from maximum likelihood analysis based on a concatenated LSU, SSU, TEF1-α 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
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. 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
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 barsa = 200 μm, b = 50 μm, c–h = 20 μm, i–p = 10 μm
Holotype: ITALY, Forlì-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 morphAscomata 170–230 μm high, 160–260 µm diam., scattered, solitary, semi-immersed to superficial, visible as raised, black dots on the host surface, globose to subglobose, uni-loculate, glabrous, ostiole central, with minute, papilla (20–45 × 25–55 μm, l/w), lacking periphyses. Peridium 13–23 μm 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 μm wide, filamentous, septate, pseudoparaphyses, anastomosing above the asci, embedded in a hyaline gelatinous matrix. Asci (70–)75–95(–113) × (9.5–)10–13(–14) μm (\( \bar{x} \) = 88.9 × 12 μm, 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) × 4.5–6 μm (\( \bar{x} \) = 26.1 × 5.5 μm, 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 (KUN-HKAS 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 morphAscomata scattered, solitary or in groups, semi-immersed to erumpent, globose to subglobose, or irregular in shape, uni-loculate, 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, smooth-walled, 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-α and ITS sequence dataset (Fig. 29) show that our new strains (KUMCC 18-0157 and KUMCC 18-0158) 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 μm thick), smaller, cylindrical asci (70–85 × 9–17 μm) and smaller, 1-seriate, muriform ascospores, with 2–4 transverse septa and 1 longitudinal septum (10–15 × 5–6.5 μm) (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 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
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 barsa = 500 μm, b = 200 μm, c = 100 μm, d = 50 μm, e–i = 20 μm, j–o = 10 μm
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 morphAscomata 170–280 μm high, 230–330 µm 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 μm 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 μm wide, filamentous, septate, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (67–)80–110(–132) × (13–)16–20(–24) μm (\( \bar{x} \) = 98 × 18.8 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindrical to cylindric-clavate, or clavate, short pedicellate, with truncate pedicel, apically rounded, with inconspicuous ocular chamber, clearly seen when immature. Ascospores (12–)14–18(–20)(–23) × (4–)5–8(–13) μm (\( \bar{x} \) = 17 × 7.7 μm, 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), ex-type living culture KUMCC 18-0157 (KIB030A), KUMCC 18-0158 (KIB030B).
GenBank numbers: ITS = MK356372, LSU = MK356346, SSU = MK356360, TEF1-α = MK359064, RPB2 = MK359079 (KUMCC 18-0157); ITS = MK356373, LSU = MK356347, SSU = MK356361, TEF1-α = MK359065 (KUMCC 18-0157).
Galiicola Tibpromma et al.
Galiicola was introduced by Ariyawansa et al. (2015a) to accommodate a single species G. pseudophaeosphaeria 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 baoshanensis Phookamsak, Wanas. & K.D. Hyde, sp. nov.
Index Fungorum number: IF556167; Facesoffungi number: FoF05701, 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 barsa = 200 μm, b = 50 μm, d = 20 μm, c, g–m = 10 μm, e, f = 5 μm
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 μm high, 90–120 μm 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 μm 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) × (2–)3–6 μm (\( \bar{x} \) = 5.7 × 4.7 µm, 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 × (10–)11–14(–15.5) μm (\( \bar{x} \) = 39.6 × 12.9 µm, 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-α = MK359066.
Notes: Multigene phylogenetic analyses of a combined LSU, SSU, TEF1-α 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-α 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. 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-α 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
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 barsa = 200 μm, b = 50 μm, c–h, o = 10 μm, i–n = 5 μm
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 μm high, 80–160 μm diam., pycnidial, scattered, solitary to gregarious, immersed to semi-immersed, uni-loculate, 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 μm 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 × (3–)5–10 μm (\( \bar{x} \) = 6.2 × 6.7 µm, 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) × 4–5 μm (\( \bar{x} \) = 11 × 4.8 µm, 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 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-α = MK359067 (KUMCC 18-0155); ITS = MK356376, LSU = MK356350, SSU = MK356364, TEF1-α = 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-α 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 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).
Ophiobolus malleolus S.K. Huang, Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF554782; Facesoffungi number: FoF04686, Fig. 34
Ophiobolus malleolus (MFLU 15-2230, holotype) a Herbarium 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 barsc = 500 µm, d = 100 µm, e = 50 µm, f–m = 20 µm
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 morphAscomata 230–270 μm high, 265–310 μm diam., scattered, solitary to gregarious, immersed, eventually erumpent, uni-loculate, globose to subglobose, dark brown to black, papillate. Ostiole central, short papilla, filled with periphyses. Peridium 25–60 μm 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 μm wide, filamentous, septate pseudoparaphyses, anastomosing at the apex, embedded in a gelatinous matrix. Asci 125–155 × 12–15 μm (\( \bar{x} \) = 140 × 12.5 μm, 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 × 2.5–4.5 μm (\( \bar{x} \) = 107.5 × 3.5 μm, 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 (47.8547249˚N, 40.2318907˚E), 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, 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 μm diam., and 120–130 × 2.5–3 μm; 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
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 barsa = 100 μm, b = 50 μm, c, d = 20 μm, e, g = 10 μm, f, h–k = 5 μm
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 μm high, (60–)135–200 μm diam., pycnidial, visible as black dot on host surface, scattered to clustered, solitary to gregarious, semi-immersed to erumpent, uni-loculate, subglobose to ampulliform, or irregular in shape, ostiolate, with minute papilla. Conidiomata walls 10–20(–30) μm 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 × 4–8(–10) μm (\( \bar{x} \) = 6.4 × 5.7 μm, 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) × 3–5 μm (\( \bar{x} \) = 12.9 × 3.9 μm, n = 50), pale brown to light yellowish, oblong to ellipsoidal, or subcylindrical, (0–)1–2-septate, 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-α = MK359069 (KUMCC 18-0161); ITS = MK356379, LSU = MK356353, SSU = MK356367, TEF1-α = 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-α 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-α 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) × 3–5 μm versus 8–12 × 2.4–3.5 μm, 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 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
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 barsa = 200 μm, b = 50 μm, c, i = 20 μm, h, j–l = 10 μm, d–g = 5 μm
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 μm high, 110–190 μm 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 μm 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) × (2.5–)3.5–6(–8) μm l/w (\( \bar{x} \) = 5 × 5.3 µm, 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) × 4–6(–7) μm l/w (\( \bar{x} \) = 22 × 5.6 µm, n = 50), dark brown, subcylindrical to fusiform, or reniform, slightly curved, with acute to rounded apex, and rounded to truncate base, 3–7-septate, not constricted at the septa, thick-walled, smooth-walled, 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 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-α = MK359071 (KUMCC 18-0159); ITS = MK356381, LSU = MK356355, SSU = MK356369, TEF1-α = 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-α genes could not resolve the novelty of the Wojnowiciella species in this study. A comparison of ITS and TEF1-α 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-α 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) × 4–6(–7) μm versus 26–36 × 4–6 µm; Hernandez-Restrepo 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).
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 Hernandez-Restrepo 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 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 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 morphAscomata 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-α 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.
Phylogram generated from maximum likelihood analysis based on LSU, SSU, ITS, RPB2 and TEF1-α 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
Pseudoastrosphaeriellopsis shares morphological similarity with Pseudoastrosphaeriella in having immersed ascomata underneath the host epidermis, erumpent, uni-loculate, 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.
Index Fungorum number: IF555791; Facesoffungi number: FoF05707, 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 barsb = 100 μm, c, d = 50 μm, e–p = 10 μm
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 morphAscomata 225–335 µm high, 220–345 µm diam., immersed to erumpent, globose to subglobose, solitary to gregarious, coriaceous, brown, short papillate, ostiolate. Ostioles 110–165 µm long, 50–90 µm diam., central, cylindrical, straight to oblique, with ostiolar canal lined by hyaline periphyses. Peridium 15–25 µm 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 µm wide, filamentous, septate, branched, hyaline pseudoparaphyses, embedded in a gelatinous matrix, anastomosing between and above the asci. Asci 85–145 × 15–18 µm (\( \bar{x} \) = 108 × 17 µm, 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 × 5–10 µm (\( \bar{x} \) = 34 × 7 µm, 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-α = 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.
Maximum likelihood phylogenetic tree based on a combined LSU, SSU, ITS, TEF1-α 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 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
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 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
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 barsa = 200 µm, b = 100 µm, c = 20 µm, d–j = 5 µm
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 μm high, 115–180 μm 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 µm 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 × 2–5 μm (\( \bar{x} \) = 5.2 × 3.4 μm, 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 × 2–4 µm (\( \bar{x} \) = 4 × 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.
Neoroussoella leucaenae Jayasiri, E.B.G. Jones & K.D. Hyde, Mycosphere 10(1): 1–186 (2019), 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 barsb–d = 200 µm, g = 30 µm, e = 20 µm, f, h, i = 10 µm
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 morphAscomata 130–195 µm high, 150–170 µm 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 µm, 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 µm wide, hyaline, septate, filamentous, anastomosed pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (35–)42–68(–75) × 4–5(–6) µm (\( \bar{x} \) = 53 × 5.5, n = 15), 8-spored, bitunicate, cylindrical, with a short pedicel, apically rounded, with an indistinct ocular chamber. Ascospores 7–10(– 15) × 3–4.5(– 6.5) µm (\( \bar{x} \) = 9.4 × 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-α = 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 and TEF1-α among N. leucaenae strains MFLUCC 18-1544, MFLUCC 17-0927 and MFLUCC 17-0346, which confirms that they are conspecific.
Roussoella Sacc.
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 elaeicola Konta & K.D. Hyde, sp. nov.
Index Fungorum number: IF555291; Facesoffungi number: FoF04834, Fig. 42
Roussoella elaeicola (MFLU 15-0022, holotype). a, b Ascomata 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 barsa, b = 1000 μm, c = 100 μm, d = 50 μm, e–h, q = 20 μm, i–p = 5 μm
Etymology: Name reflects the host genus Elaeis.
Holotype: MFLU 15-0022
Saprobic on dead petiole of Elaeis guineensis. Sexual morphAscomata 315–410 µm high, 325–350 μm 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 µm 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 μm broad, filamentous, septate, trabeculate pseudoparaphyses, embedded in mucilage. Asci 70–140 × 6–9 μm (\( \bar{x} \) = 100 × 8 µm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, pedicellate, rounded apex with an indistinct ocular chamber. Ascospores 10–15 × 3–6 μm (\( \bar{x} \) = 12 × 4 µm, 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 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).
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 ≥ 50% are indicated at the nodes. The new species is indicated in blue. Ex-type strains are indicated in bold
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.
Index Fungorum number: IF555293; Facesoffungi number: FoF05708, 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. l Apex of the ascus. m–q Ascospores. Scale barsd = 50 μm, f = 20 μm, e, g–k = 10 μm, l–q = 5 μm
Etymology: The specific epithet “brunnea” refers to the brownish ascospores.
Holotype: MFLU 18-1393
Saprobic on dead stem of herbage. Sexual morphAscomata 80–205 μm high, 90–260 μm diam., semi-immersed, blackish, irregular in shape, scattered on surface of stem, uni-loculate, glabrous, ostiolate, apapillate. Ostioles 35.5–68 μm diam., dark, circular and sunken at the apex of ascoma. Peridium 30–40 μm thick, two layered, thick-walled, outer layer irregular, comprising dark brown cells of textura angularis and inner layer slightly, irregular of light brown cells. Hamathecium comprising 1–2 μm wide, filamentous, branched or simple, septate, anastomosed, cellular pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci 70–90 × 8–10 μm (\( \bar{x} \) = 83.6 × 9 μm, n = 10), 4- or 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with an ocular chamber. Ascospores 11–18 × 5–6 μm (\( \bar{x} \) = 12.9 × 5.8 μm, 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
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-α 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 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). Therefore, we introduce Liua as a new genus to accommodate L. muriformis based on its morphological distinct with Cycasicola.
Liua muriformis Phookamsak, H.B. Jiang & K.D. Hyde, sp. nov.
Index Fungorum number: IF556176; Facesoffungi number: FoF05710, 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 barsa = 200 μm, b = 50 μm, c = 20 μm, l, m = 10 μm, d–k = 5 μm
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 μm high, 110–190 μm 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, with pore-like opening. Conidiomata walls 7–17 μm 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) × (2–)5–10 μm l/w (\( \bar{x} \) = 8.2 × 5.4 µm, 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) × 7–9(–10) μm l/w (\( \bar{x} \) = 14.4 × 8.2 µm, 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-α = 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).
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 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)
Lembosia brigadeirensis Firmino, A.R. Machado & O.L. Pereira, sp. nov.
MycoBank number: MB822355; Facesoffungi number: FoF04108, 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 barsa = 200 µm, b–f = 10 µm
Etymology: Name derived from the mountain range where the fungus was collected, Serra do Brigadeiro.
Holotype: VIC 44208
Epiphyllous on Epidendrum sp. Sexual morphColonies 4–6 mm diam., amphigenous, circular to irregular, single to confluent, dark brown, black. Hyphae 4–5 μm diam., straight to flexuous, irregularly branched, brown, septate, hyphal cells cylindrical, smooth. Appressoria 5.5–7.5 × 6–8 μm, 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 × 140–230 μm, 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 μm wide, cylindrical, filiform, septate, unbranched, hyaline. Asci 30–47.5 × 15–22 μm, 8-spored, bitunicate, fissitunicate, subclavate to cylindrical, disposed as an upright palisade layer. Ascospores 17.5–19.5 × 5.5–8 μm, 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