Fungal Diversity (2020) 104:1–266
https://doi.org/10.1007/s13225-020-00461-7
Fungal diversity notes 1277–1386: taxonomic and phylogenetic
contributions to fungal taxa
Hai‑Sheng Yuan1,2 · Xu Lu1,2 · Yu‑Cheng Dai3 · Kevin D. Hyde4,5,6,7,8 · Yu‑He Kan1,2 · Ivana Kušan9 · Shuang‑Hui He3 ·
Ning‑Guo Liu5,10 · V. Venkateswara Sarma11 · Chang‑Lin Zhao12 · Bao‑Kai Cui3 · Nousheen Yousaf13 · Guangyu Sun14 ·
Shu‑Yan Liu15,31 · Fang Wu3 · Chuan‑Gen Lin5,6,7 · Monika C. Dayarathne5,27 · Tatiana Baptista Gibertoni16 ·
Lucas B. Conceição17 · Roberto Garibay‑Orijel18 · Margarita Villegas‑Ríos19 · Rodolfo Salas‑Lizana19 ·
Tie‑Zheng Wei20 · Jun‑Zhi Qiu21 · Ze‑Fen Yu22 · Rungtiwa Phookamsak4,5,8 · Ming Zeng4,5 · Soumitra Paloi23 ·
Dan‑Feng Bao5,24,25 · Pranami D. Abeywickrama5,6,26 · De‑Ping Wei4,5,7,25 · Jing Yang5 · Ishara S. Manawasinghe5,26 ·
Dulanjalee Harishchandra5,6,26 · Rashika S. Brahmanage5,6,26 · Nimali I. de Silva4,8,28,29 · Danushka S. Tennakoon4,5,6 ·
Anuruddha Karunarathna5,30 · Yusufjon Gafforov1,32,33 · Dhandevi Pem5 · Sheng‑Nan Zhang5,30 ·
André L. C. M. de Azevedo Santiago34 · Jadson Diogo Pereira Bezerra35 · Bálint Dima36 · Krishnendu Acharya23 ·
Julieta Alvarez‑Manjarrez18,58 · Ali H. Bahkali37,38 · Vinod K. Bhatt39 · Tor Erik Brandrud40 · Timur S. Bulgakov41 ·
E. Camporesi42,43,44 · Ting Cao1,2 · Yu‑Xi Chen21 · Yuan‑Yuan Chen45 · Bandarupalli Devadatha11 ·
Abdallah M. Elgorban37,38 · Long‑Fei Fan3 · Xing Du22 · Liu Gao14 · Camila Melo Gonçalves46 · Luis F. P. Gusmão17 ·
Naruemon Huanraluek5 · Margita Jadan9 · Ruvishika S. Jayawardena5 · Abdul Nasir Khalid47 · Ewald Langer33 ·
Diogo X. Lima34 · Nelson Correia de Lima‑Júnior48 · Carla Rejane Sousa de Lira16 ·
Jian‑Kui (Jack) Liu49 · Shun Liu3 · Saisamorn Lumyong28,29,50 · Zong‑Long Luo24 · Neven Matočec9 · M. Niranjan11 ·
José Ribamar Costa Oliveira‑Filho16 · Viktor Papp51 · Eduardo Pérez‑Pazos19,52 · Alan J. L. Phillips53 ·
Peng‑Lei Qiu15 · Yihua Ren14 · Rafael F. Castañeda Ruiz54 · Kamal C. Semwal55 · Karl Soop56 · Carlos A. F. de Souza34 ·
Cristina Maria Souza‑Motta46 · Li‑Hua Sun57 · Meng‑Le Xie31 · Yi‑Jian Yao20 · Qi Zhao4 · Li‑Wei Zhou1
Received: 10 January 2020 / Accepted: 28 August 2020 / Published online: 31 October 2020
© MUSHROOM RESEARCH FOUNDATION 2020
Abstract
This is the twelfth contribution to the Fungal Diversity Notes series on fungal taxonomy, based on materials collected from many
countries which were examined and described using the methods of morphology, anatomy, and strain culture, combined with DNA
sequence analyses. 110 taxa are described and illustrated, including five new genera, 92 new species, eight new combinations and
other taxonomic contributions (one new sequenced species, one new host and three new records) which are accommodated in
40 families and 1 incertae sedis in Dothideomycetes. The new genera are Amyloceraceomyces, Catenuliconidia, Hansenopezia,
Ionopezia and Magnopulchromyces. The new species are Amyloceraceomyces angustisporus, Amylocorticium ellipsosporum,
Arthrinium sorghi, Catenuliconidia uniseptata, Clavulina sphaeropedunculata, Colletotrichum parthenocissicola, Coniothyrium
triseptatum, Cortinarius indorusseus, C. paurigarhwalensis, C. sinensis, C. subsanguineus, C. xiaojinensis, Diaporthe pimpinellae, Dictyosporella guizhouensis, Diplodia torilicola, Fuscoporia marquesiana, F. semiarida, Hansenopezia decora, Helicoarctatus thailandicus, Hirsutella hongheensis, Humidicutis brunneovinacea, Lentaria gossypina, L. variabilis, Lycoperdon lahorense,
L. pseudocurtisii, Magnopulchromyces scorpiophorus, Moelleriella gracilispora, Neodevriesia manglicola, Neodidymelliopsis
salvia, N. urticae, Neoroussoella magnoliae, Neottiella gigaspora, Ophiosphaerella chiangraiensis, Phaeotremella yunnanensis,
Podosphaera yulii, Rigidoporus juniperinus, Rhodofomitopsis pseudofeei, Russula benghalensis, Scleroramularia vermispora,
Scytinopogon minisporus, Sporormurispora paulsenii, Thaxteriellopsis obliqus, Tomentella asiae-orientalis, T. atrobadia, T.
atrocastanea, T. aureomarginata, T. brevis, T. brunneoflava, T. brunneogrisea, T. capitatocystidiata, T. changbaiensis, T. citrinocystidiata, T. coffeae, T. conclusa, T. cystidiata, T. dimidiata, T. duplexa, T. efibulata, T. efibulis, T. farinosa, T. flavidobadia,
T. fuscocrustosa, T. fuscofarinosa, T. fuscogranulosa, T. fuscopelliculosa, T. globospora, T. gloeocystidiata, T. griseocastanea, T.
griseofusca, T. griseomarginata, T. inconspicua, T. incrustata, T. interrupta, T. liaoningensis, T. longiaculeifera, T. longiechinuli, T.
megaspora, T. olivacea, T. olivaceobrunnea, T. pallidobrunnea, T. pallidomarginata, T. parvispora, T. pertenuis, T. qingyuanensis,
T. segregata, T. separata, T. stipitata, T. storea, Trichoderma ceratophylletum, Tyromyces minutulus, Umbelopsis heterosporus and
Extended author information available on the last page of the article
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Fungal Diversity (2020) 104:1–266
Xylolentia reniformis. The new combinations are Antrodiella descendena, Chloridium macrocladum, Hansenopezia retrocurvata,
Rhodofomitopsis monomitica, Rh. oleracea, Fuscoporia licnoides, F. scruposa and Ionopezia gerardii. A new sequenced species
(Graphis supracola), one new host (Aplosporella prunicola) and three new geographical records (Golovinomyces monardae,
Paradictyoarthrinium diffractum and Prosthemium betulinum), are reported.
Keywords 110 taxa · Agaricomycetes · Ascomycota · Basidiomycota · Mucoromycota · New combinations · New genera ·
New species · Phylogeny · Taxonomy
Table of contents
Mucoromycota
Umbelopsidales Spatafora, Stajich & Bonito
Umbelopsidaceae W. Gams & W. Mey
1277. Umbelopsis heterosporus C.A. de Souza, D.X. Lima
& A.L. Santiago, sp. nov.
Ascomycota
Dothideomycetes
Botryosphaeriales C.L. Schoch et al.
Aplosporellaceae Slippers, Boissin & Crous
1278. Aplosporella prunicola Damm & Crous, Fungal
Diversity 27(1): 39 (2007), a new host record
Botryosphaeriaceae Theiss. & H. Syd.
1279. Diplodia torilicola Harishchandra, Camporesi & K.D.
Hyde, sp. nov.
Capnodiales Woron
Neodevriesiaceae Quaedvl. & Crous, in Quaedvlieg et al.,
Persoonia 33: 24 (2014)
1280. Neodevriesia manglicola Devadatha, V.V. Sarma &
E.B.G. Jones, sp. nov.
Pleosporales Luttr. ex M.E. Barr
Coniothyriaceae W.B. Cooke
1281. Coniothyrium triseptatum Dayarathne, Thyagaraja
& K.D. Hyde, sp. nov.
Didymellaceae Gruyter, Aveskamp & Verkley
1282. Neodidymelliopsis salviae Brahmanage, Camporesi
& K.D. Hyde, sp. nov.
1283. Neodidymelliopsis urticae Manawas., Camporesi &
K.D. Hyde, sp. nov.
Lophiostomataceae Sacc.
1284. Magnopulchromyces L.B. Conc., Gusmão & R.F.
Castañeda, gen. nov.
1285. Magnopulchromyces scorpiophorus L.B. Conc.,
Gusmão & R.F. Castañeda, sp. nov.
13
Paradictyoarthriniaceae Doilom, Jian K. Liu & K.D. Hyde
1286. Paradictyoarthrinium diffractum Matsush., Matsush.
Mycol. Mem. 9:18 (1996), a new host and geographical
record from India
Phaeosphaeriaceae M.E. Barr
1287. Ophiosphaerella chiangraiensis Phookamsak & K.D.
Hyde, sp. nov.
Pleomassariaceae M.E. Barr
1288. Prosthemium betulinum Kunze, in Kunze & Schmidt,
Mykologische Hefte (Leipzig) 1: 18 (1817), a new record
from Italy
Roussoellaceae J.K. Liu, R. Phookamsak, D.Q. Dai & K.D.
Hyde1289. Neoroussoella magnoliae N.I. de Silva & K.D.
Hyde, sp. nov.
Sporormiaceae Munk
1290. Sporormurispora paulsenii D.Pem, Gafforov & K.D.
Hyde, sp. nov.
Tubeufiales Boonmee & K.D. Hyde
Tubeufiaceae M.E. Barr
1291. Helicoarctatus thailandicus D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, sp. nov.
1292. Thaxteriellopsis obliqus M. Niranjan and V.V. Sarma,
sp. nov.
Incertae sedis in Dothideomycetes
1293. Scleroramularia vermispora G.Y. Sun & L. Gao, sp.
nov.
Lecanoromycetes
Graphidales Bessey
Graphidaceae Dumort.
1294. Graphis supracola A.W. Archer, Aust. Syst. Bot.
14(2): 267 (2001), a new sequenced species from Thailand
Leotiomycetes
Erysiphales Gwynne-Vaughan & Barnes
Fungal Diversity (2020) 104:1–266
Erysiphaceae Tul. & C. Tul.
1295. Podosphaera yulii S.-Y. Liu & P.-L. Qiu, sp. nov.
1296. Golovinomyces monardae (G.S. Nagy) M. Scholler,
U. Braun & Anke Schmidt, in Scholler, et al., Mycol. Progr.
15(no. 56): 4 (2016), a new record from China
Pezizomycetes
Pezizales J. Schröt.
Pezizaceae Dumort.
1297. Ionopezia Matočec, I. Kušan & Jadan, gen. nov.
1298. Ionopezia gerardii (Cooke) Matočec, I. Kušan &
Jadan, comb. nov.
1299. Hansenopezia Matočec, I. Kušan & Jadan, gen. nov.
1300. Hansenopezia retrocurvata (K. Hansen & Sandal)
Matočec, I. Kušan & Jadan, comb. nov.
1301. Hansenopezia decora Matočec, I. Kušan & Jadan,
sp. nov.
Pyronemataceae Corda
1302. Neottiella gigaspora M. Zeng, Q. Zhao & K.D. Hyde,
sp. nov.
3
1308. Hirsutella hongheensis D.P. Wei & K.D. Hyde, sp.
nov.
Glomerellales Chadef. ex Réblová et al.
Glomerellaceae Locq. ex Seifert & W. Gams
1309. Colletotrichum parthenocissicola Jayawardena, Bulgakov, Huanraleuk & K.D. Hyde, sp. nov.
Junewangiaceae J.W. Xia & X.G. Zhang
1310. Dictyosporella guizhouensis J. Yang & K.D. Hyde,
sp. nov.
Rhamphoriaceae Réblová
1311. Xylolentia reniformis C.G. Lin, K.D. Hyde & Jian
K. Liu, sp. nov.
Xylariales Nannf.
Xylariaceae Tul. & C. Tul.
1312. Catenuliconidia N.G. Liu & K.D. Hyde, gen. nov.
1313. Catenuliconidia uniseptata N.G. Liu & K.D. Hyde,
sp. nov.
Sordariomycetes
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Basidiomycota
Agaricomycetes
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor &
M.E. Barr
1303. Arthrinium sorghi J.D.P. Bezerra, C.M Gonçalves &
C.M. Souza-Motta, sp. nov.
Agaricales Underw.
Agaricaceae Chevall.
1314. Lycoperdon lahorense Yousaf & Khalid, sp. nov.
1315. Lycoperdon pseudocurtisii Yousaf & Khalid, sp. nov.
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández
Chaetosphaeriaceae Réblová, M.E. Barr & Samuels
1304. Chloridium macrocladum (Sacc.) Karun., Maharachch., C.H. Kuo & K.D. Hyde, comb. nov.
Cortinariaceae R. Heim ex Pouzar
1316. Cortinarius indorusseus Dima, Semwal, V.K. Bhatt
& Brandrud, sp. nov.
1317. Cortinarius paurigarhwalensis Semwal, Dima &
Soop, sp. nov.
1318. Cortinarius sinensis L.H. Sun, T.Z. Wei & Y.J. Yao,
sp. nov.
1319. Cortinarius subsanguineus T.Z. Wei, M.L. Xie &
Y.J. Yao, sp. nov.
1320. Cortinarius xiaojinensis T.Z. Wei, M.L. Xie & Y.J.
Yao, sp. nov.
Diaporthales Nannf.
Diaporthaceae Höhn. ex Wehm.
1305. Diaporthe pimpinellae Abeywickrama, Camporesi,
Dissanayake & K.D. Hyde, sp. nov.
Hypocreales Lindau
Clavicipitaceae Kirk, Cannon, Minter & Stalpers
1306. Moelleriella gracilispora Jun Z. Qiu & Y.X. Chen,
sp. nov.
Hypocreaceae De Not.
1307. Trichoderma ceratophylletum Z.F. Yu & X. Du, sp.
nov.
Ophiocordycipitaceae G.H. Sung, J.M. Sung, Hywel-Jones
& Spatafora, in Sung, Hywel-Jones, Sung, Luangsa-ard,
Shrestha & Spatafora
Hygrophoraceae Lotsy
Humidicuteae Padamsee & Lodge, Fungal Diversity 64: 38
(2014)
1321. Humidicutis brunneovinacea R. Garibay-Orijel, sp.
nov.
Amylocorticiales K.H. Larss., Manfr. Binder & Hibbett
Amylocorticiaceae Jülich
1322. Amyloceraceomyces S.H. He, gen. nov.
1323. Amyloceraceomyces angustisporus S.H. He, sp. nov.
1324. Amylocorticium ellipsosporum S.H. He, sp. nov.
13
4
Cantharellales Gäum
Clavulinaceae Donk
1325. Clavulina sphaeropedunculata E. Pérez-Pazos, M.
Villegas & R. Garibay-Orijel, sp. nov.
Gomphales Jülich
Lentariaceae Jülich
1326. Lentaria gossypina R. Salas-Lizana, M. Villegas &
E. Pérez-Pazos, sp. nov.
1327. Lentaria variabilis M. Villegas, R. Garibay-Orijel &
N. Matías-Ferrer, sp. nov.
Hymenochaetales Oberw.
Hymenochaetaceae Donk
1328. Fuscoporia licnoides (Mont.) Oliveira-Filho & Gibertoni, comb. nov.
1329. Fuscoporia marquesiana Gibertoni & C.R.S. de Lira,
sp. nov.
1330. Fuscoporia scruposa (Mont.) Gibertoni & OliveiraFilho, comb. nov.
1331. Fuscoporia semiarida Lima-Júnior, C.R.S. de Lira &
Gibertoni, sp. nov.
1332. Rigidoporus juniperinus Gafforov, L.W. Zhou, E.
Langer, sp. nov.
Polyporales Gäum.
Fomitopsidaceae Jülich
1333. Rhodofomitopsis pseudofeei B.K. Cui & Shun Liu,
sp. nov.
1334. Rhodofomitopsis monomitica (Yuan Y. Chen) B.K.
Cui, Yuan Y. Chen & Shun Liu, comb. nov.
1335. Rhodofomitopsis oleracea (R.W. Davidson & Lombard) B.K. Cui, Yuan Y. Chen & Shun Liu, comb. nov.
Polyporaceae Fr. ex Corda
1336. Antrodiella descendena (Corner) C.L. Zhao & Y.C.
Dai, comb. nov.
1337. Tyromyces minutulus Y.C. Dai & C.L. Zhao, sp. nov.
Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C. David
Russulaceae Lotsy
1338. Russula benghalensis S. Paloi & K. Acharya, sp. nov.
Thelephorales Corner ex Oberw.
Thelephoraceae Chevall.
1339. Tomentella asiae-orientalis H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1340. Tomentella atrobadia H.S. Yuan & Y.C. Dai, sp. nov.
1341. Tomentella atrocastanea H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1342. Tomentella aureomarginata H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
13
Fungal Diversity (2020) 104:1–266
1343. Tomentella brevis H.S. Yuan, X. Lu & Y.C. Dai, sp.
nov.
1344. Tomentella brunneoflava H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1345. Tomentella brunneogrisea H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1346. Tomentella capitatocystidiata H.S. Yuan, X. Lu &
Y.C. Dai, sp. nov.
1347. Tomentella changbaiensis H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1348. Tomentella citrinocystidiata H.S. Yuan & Y.C. Dai,
sp. nov.
1349. Tomentella coffeae H.S. Yuan & Y.C. Dai, sp. nov.
1350. Tomentella conclusa H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1351. Tomentella cystidiata H.S. Yuan & Y.C. Dai, sp. nov.
1352. Tomentella dimidiata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1353. Tomentella duplexa H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1354. Tomentella efibulata H.S. Yuan & Y.C. Dai, sp. nov.
1355. Tomentella efibulis H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1356. Tomentella farinosa H.S. Yuan & Y.C. Dai, sp. nov.
1357. Tomentella flavidobadia H.S. Yuan & Y.C. Dai, sp.
nov.
1358. Tomentella fuscocrustosa H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1359. Tomentella fuscofarinosa H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1360. Tomentella fuscogranulosa H.S. Yuan & Y.C. Dai,
sp. nov.
1361. Tomentella fuscopelliculosa H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1362. Tomentella globospora H.S. Yuan & Y.C. Dai, sp.
nov.
1363. Tomentella gloeocystidiata H.S. Yuan & Y.C. Dai,
sp. nov.
1364. Tomentella griseocastanea H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1365. Tomentella griseofusca H.S. Yuan & Y.C. Dai, sp.
nov.
1366. Tomentella griseomarginata H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1367. Tomentella inconspicua H.S. Yuan & Y.C. Dai, sp.
nov.
1368. Tomentella incrustata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1369. Tomentella interrupta H.S. Yuan & Y.C. Dai, sp. nov.
1370. Tomentella liaoningensis H.S. Yuan & Y.C. Dai, sp.
nov.
1371. Tomentella longiaculeifera H.S. Yuan & Y.C. Dai,
sp. nov.
Fungal Diversity (2020) 104:1–266
1372. Tomentella longiechinuli H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1373. Tomentella megaspora H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1374. Tomentella olivacea H.S. Yuan & Y.C. Dai, sp. nov.
1375. Tomentella olivaceobrunnea H.S. Yuan, X. Lu &
Y.C. Dai, sp. nov.
1376. Tomentella pallidobrunnea H.S. Yuan, X. Lu & Y.C.
Dai, sp. nov.
1377. Tomentella pallidomarginata H.S. Yuan, X. Lu &
Y.C. Dai, sp. nov.
1378. Tomentella parvispora H.S. Yuan & Y.C. Dai, sp.
nov.
1379. Tomentella pertenuis H.S. Yuan & Y.C. Dai, sp. nov.
1380. Tomentella qingyuanensis H.S. Yuan & Y.C. Dai,
sp. nov.
1381. Tomentella segregata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1382. Tomentella separata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1383. Tomentella stipitata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
1384. Tomentella storea H.S. Yuan & Y.C. Dai, sp. nov.
Trechisporales K.H. Larsson
Hydnodontaceae Jülich
1385. Scytinopogon minisporus J. Alvarez-Manjarrez, M.
Villegas & R. Garibay-Orijel, sp. nov.
Tremellomycetes
Tremellales Fr.
Phaeotremellaceae A.M. Yurkov & Boekhout
1386. Phaeotremella yunnanensis L.F. Fan, F. Wu & Y.C.
Dai, sp. nov.
5
Fungi have varied forms, including unicellular (yeast),
multicellular (e.g. molds) and macrofungi (e.g. mushrooms).
According to the estimation by different researchers, there
are 1,500,000 to 6,000,000 fungal species worldwide
(Hawksworth 1991, 2012; Blackwell 2011; Taylor et al.
2014). However, only 120,000 species have been described
and accepted (Hawksworth and Lücking 2017), and most
were from Europe and North America. Many regions outside
of Europe, such as East Asia, South America and Africa
have, until recently, received little attention, despite the high
levels of biodiversity (Hyde et al. 2018).
In the last two decades (2000–2020), contributions to
fungal taxonomy, in countries such as China and Thailand
have increased rapidly (Dai et al. 2015; Hyde et al. 2018).
With the wide application of molecular techniques in fungal taxonomy, numerous new taxa have been resolved from
species complexes (Hyde et al. 2014; Nilsson et al. 2014),
which were previously difficult to determine using classical methods. In recent years, high-throughput sequencing
technology has made it possible to detect, trace and discover
unculturable taxa from environmental samples, such as soil,
water and air, and this has promoted the discovery of many
more undescribed, but faceless taxa (Hibbett 2016; Hongsanan et al. 2018; Tedersoo et al. 2018).
The Fungal Diversity series (Hyde et al. 2019, 2020a, b)
publishes notes on new fungal taxa. To date, more than 1,200
fungal taxa have been introduced, including new taxa, new
sequenced isolates and other taxonomic contributions, based
on morphological characteristics and phylogenetic analyses.
This is the 12th paper in the Fungal Diversity Notes series
with 110 taxa from contributions made world-wide.
Materials and methods
Specimens, isolates and identification
Introduction
Fungi are widely distributed in ecosystems and play a
key role in nutrient cycling, maintaining biodiversity and
enhancing forest regeneration (Zechmeister-Boltenstern
et al. 2015; Willis 2018). Fungi have developed various
tropisms with plants and animals, such as endophytism,
biotrophism, hemi-biotrophism, necrotrophism and saprotrophism (de Silva et al. 2016). They have established close
relationships with various organisms, such as plants, animals, insects, protists (algae) and bacteria in the biosphere
(Landeweert et al. 2001; Grube and Berg 2009; Torruella
et al. 2015). Diverse hosts and substrates in terrestrial and
aquatic ecosystems provide important substrates for the
growth and reproduction of fungi (Hyde and Jones 1988;
Dai et al. 2017; Luo et al. 2019).
The studied specimens were collected from many countries
(Australia, Bosnia and Herzegovina, Brazil, Cameroon,
China, Croatia, Italy, India, Malaysia, Mexico, Pakistan,
Russia, Thailand, Uzbekistan, Zambia and Zimbabwe)
occurring in leaf litter on humid soil, on fallen branches
and rotten wood debris of angiosperms and gymnosperms,
and with various host plants. Isolates were cultured on cornmeal dextrose agar (CMD), potato dextrose agar (PDA) and
synthetic low nutrient agar (SNA) (Silva et al. 2000). The
specimens and ex-type living cultures were deposited in
various herbaria (AMH, BJFC, CAL, CCLAMIC, CGMCC,
CNF, CUH, E, F, FAMU, FCME, GZCC, HKAS, HMAS,
HMJAU, HMUABO, HUEFS, IFP, JZBH, KUMCC, LAH,
MEXU, MFLU, MFLUCC, NFCCI, PUFNI, TASM, URM
and YMF), which are listed under each taxon description.
13
6
The microscopic observation procedure follows Baral
(1992), Dai (2010), Hyde et al. (2016) and Wanasinghe et al.
(2017a) with some minor amendments. Macro-morphological characteristics of the specimens were examined using a
Nikon SMZ 645, a Zeiss Axioskop 40 FL and other types of
stereo microscopes. Living or dried fruitbodies of ascomycetes and basidiomycetes were tested in Congo Red, Cotton
Blue, hydrochloric acid (HCl), Lugol’s solution, Melzer’s
reagent, brilliant Cresyl Blue, protocetraric acid, 5% KOH,
or distilled water. Sections were cut with a razor blade and
studied with a Nikon Eclipse 80i and other different types of
phase-contrast microscopes; micro-morphological characteristics including ascomata sections, peridium structures, asci
and ascospores in ascomycetes, and ab- and hymenial surfaces, hyphal systems, basidia, cystidia, and basidiospores in
basidiomycetes. Photographs of the fruiting structures were
taken with a digital camera mounted on the microscope.
Measurements for ascospores, basidiospores and conidia
were taken from at least ten representatives and the means
for length and width are given. The surface morphology
of the specimens was observed with a scanning electron
microscope. Special color terms follow Watling et al. (1969),
Rayner (1970), Munsell (1976) and Kornerup and Wanscher
(1981). Index Fungorum and Faces of Fungi numbers are
provided as required (Index Fungorum 2020; Jayasiri et al.
2015).
The following abbreviations are used in the text:
CB = acyanophilous in Cotton Blue, KOH = 5% KOH,
IKI– = neither amyloid nor dextrinoid in Melzer’s reagent,
L = mean spore length (arithmetic average of all spores in
lateral views), W = mean spore width (arithmetic average
of all spores in lateral views), Q = variation in the ratios of
L/W between specimens studied, n = number of measured
structures from specimens, x = mean spore length × mean
spore width.
DNA extraction, PCR and sequencing
For most ascomycetous fungal samples, total genomic DNA
was extracted from fresh fungal mycelium grown on appropriate media at room temperature, while for basidiomycetes,
dried herbarium specimens were used. Genomic DNA was
extracted with the CTAB-based method (Góes-Neto et al.
2005), E.Z.N.A. SP Fungal DNA Mini Kit (Omega Bio-Tek,
Norcross, GA, USA), Thermo Scientific Phire Plant Direct
PCR Kit (Thermo Fisher Scientific, USA) or other fungal
DNA extraction kits following the manufacturer’s instructions. Polymerase chain reaction (PCR) was used to amplify
partial gene regions with primers shown in Table 1. The
ACT, CHS, GAPDH, ITS, LSU, mtSSU, SSU, TEF1, RPB1,
RPB2 and β-TUB partial sequences were amplified by PCR.
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Fungal Diversity (2020) 104:1–266
The sequences were provided by the Technological Platform
of Genomics, Gene Expression of the Bioscience Centre
(CB) (Recife, UFPE), Beijing Genomics Institute (BGI)
and other commercial sequencing providers depending on
the geographical areas where the studies were carried out.
Phylogenetic analyses
The newly obtained sequences were assembled and manually
modified with the software DNAMAN8 (Lynnon Biosoft,
Quebec, America). Sequence data were verified using
BLASTN in GenBank to ensure that no erroneous sequences
were used in further analyses.
The newly generated sequences reported in this paper
have been deposited in the GenBank database and they
were used to establish phylogenetic relationships with other
sequences retrieved from GenBank (https://www.ncbi.nlm.
nih.gov/) and UNITE (https://unite.ut.ee/) database. BioEdit
sequence alignment editor (Hall 2005), ClustalX (Larkin
et al. 2007), MEGA 7 (Kumar et al. 2016), MAFFT: multiple sequence alignment software version v. 7 (Katoh and
Standley 2013) and CIPRES (https://www.phylo.org) were
used for sequence alignment. The alignment formats were
converted to NEXUS and PHYLIP formats by ClustalX
and EasyCodeML v1.0 (Gao and Chen 2016). Phylogenetic
analyses were performed by Maximum Parsimony (MP),
Maximum Likelihood (RAxML) and Bayesian analysis. All
characters were weighted and gaps were treated as missing data. Maximum parsimony analyses (PAUP* version v.
4.0b10) were used (Swofford 2002) and a bootstrap (BT)
analysis with 1000 replicates (Gaget et al. 2017). Descriptive
tree statistics i.e. tree length (TL), consistency index (CI),
retention index (RI), rescaled consistency index (RC), and
homoplasy index (HI) were calculated for all trees generated
under different optimality criteria. Bayesian analysis was
carried out using MrBayes 3.2.4 (Cannatella 2015) implementing the Markov chain Monte Carlo (MCMC) technique
and parameters predetermined with the nucleotide substitution models as estimated by MrModelTest v. 2.3 (Posada and
Crandall 1998; Nylander 2004). Four simultaneous Markov
chains were run starting from random trees and keeping
one tree every 100th generation until the average standard
deviation of split frequencies was below 0.01. The value of
burn-in was set to discard 25% of trees when calculating the
posterior probabilities. The sequence dataset was subjected
to maximum likelihood phylogenetic analyses in the RAxMLGUI v. 1.3 (Silvestro and Michalak 2012; Stamatakis 2014;
Hundsdoerfer and Kitching 2017) with 1000 rapid bootstrap
replicates, and the GTR or other substitution models for the
nucleotide partitions and the default setting for binary (indel)
data were used.
Fungal Diversity (2020) 104:1–266
7
Table 1 Details of genes/loci with PCR primers and PCR profiles
Gene/loci
PCR primers (forward/reverse)
PCR conditions
References for primer
ACT
CHS
GAPDH
HIS
EF1-α
ACT512F/ACT783R
CHS79F/CHS345R
GDF/GDR
H3-1a/ H3-1b
983-F/2218-R
Carbone and Kohn (1999)
Carbone and Kohn (1999)
Templeton et al. (1992)
Glass and Donaldson (1995)
Rehner (2001)
ITS
LSU
ITS1F /ITS4
ITS5/ITS4
SSU1318-Tom/LSU-Tom4
LROR/LR5
95 °C: 40 s, 58 °C: 30 s,c 72 °C: 1 min (40 cycles)f
95 °C: 30 s, 59 °C: 30 s,d 72 °C: 45 s (35 cycles)h
95 °C: 30 s, 54 °C: 30 s, d 72 °C: 45 s (35 cycles)h
95 °C: 3 min, 58 °C: 30 s, 72 °C: 1 min (35 cycles)
94 °C: 30 min, 64 °C: 1 min, 72 °C: 1 min (10 cycles)b
94 °C: 30 min, 54 °C: 1 min, 72 °C: 1 min (35 cycles)g
94 °C: 30 s, 52 °C: 50 s,a 72 °C: 1 min (35 cycles)e
94 °C: 30 s, 52 °C: 50 s,a 72 °C: 1 min (35 cycles)e
95 °C: 30 s, 62 °C: 30 s,c 72 °C: 20 s (35 cycles)h
94 °C: 30 s, 55 °C: 50 s,a 72 °C: 1 min (35 cycles)e
mtSSU
RPB1
LROR/LR7
LROR/LR3
mrSSU1/mrSSU3R
RPB1-Af/RPB1-Cr
RPB2
SSU
TEF1
β-TUB
a
fRPB2-5f/fRPB2-7cR
NS1/NS4
EF1-983F/EF1-2218R
Btub2Fd/Btub4Rd
Bt2a/Bt2b
BT1/BT2
94 °C: 30 s, 47.2 °C: 50 s,a 72 °C: 1 min (35 cycles)e
94 °C: 30 s, 58.9 °C: 50 s,a 72 °C: 1 min (35 cycles)e
94 °C: 30 s, 55 °C: 50 s,a 72 °C: 1 min (35 cycles) d
95 °C: 30 s, 4 °C: 30 s,c
72 °C: 1 min (30 cycles)e
95 °C: 1 min, 52 °C: 2 min,a 72 °C: 1.30 s (35 cycles)e
94 °C: 30 s, 55 °C: 50 s,a 72 °C: 1 min (35 cycles)d
94 °C: 30 s, 52 °C: 50 s,a 72 °C: 1 min (35 cycles)e
95 °C: 30 s, 53 °C: 30 s, d 72 °C: 45 s (35 cycles)h
Liu et al. (1999)
White et al. (1990)
Rehner (2001)
Woudenberg et al. (2009)
Glass and Donaldson (1995)
Carbone and Kohn (1999)
Initiation step of 94 °C: 3 min
b
Initial step at 94 °C: 2 min
c
Initiation step of 95 °C: 5 min
d
Initiation step of 95 °C: 3 min
e
Final elongation step of 72 °C: 10 min and final hold at 4 °C
f
White et al. (1990)
White et al. (1990)
Taylor and McCormick (2008)
Rehner and Samuels (1994)
Vilgalys and Hester (1990)
Vilgalys and Hester (1990)
Vilgalys and Hester (1990)
Zoller et al. (1999)
Stiller and Hall (\)
Final elongation step of 72 °C: 5 min and final hold at 4 °C
g
Final elongation: 72 °C: 3 min and final hold at 4 °C
h
Final elongation step of 72 °C: 1 min and final hold at 4 °C
Taxonomy
unknown (Hoffmann et al. 2013; Wang et al. 2013; Spatafora
et al. 2016).
Mucoromycota
Umbelopsidales Spatafora, Stajich & Bonito
Notes: After an extensive study involving genomic scale
data, Spatafora et al. (2016) proposed the order Umbelopsidales Spatafora & Stajich, comprising Umbelopsidaceae
W. Gams & W. Mey. and Umbelopsis Amos & H.L. Barnett.
Species belonging to the Umbelopsidales are characterized
by their asexual reproduction, which occurs through the formation of multi-spored or uni-spored sporangia and chlamydospores. Sporangia are produced at the end of the sporangiophores, arranged in cymose or verticillate branches,
and are typically pigmented in red or ocher tones. Sporangiospores are pigmented and may vary in shape between
globose, ellipsoid or polyhedral. The columellae are usually
very small or absent and have often been described as inconspicuous. The sexual reproduction of the Umbelopsidales is
Umbelopsidaceae W. Gams & W. Mey., Mycol. Res. 107(3):
348 (2003)
Notes: The taxonomy of Umbelopsidaceae has been
extensively discussed in recent decades, and almost half of
the genus Umbelopsis (the only genus in Umbelopsidaceae)
taxa were initially described as Mortierella Coem. [e.g. M.
isabellina (Oudem.) W. Gams, M. vinacea Dixon-Stew., M.
nana (Linnem.) Arx, M. ramanniana var. autotrophica E.H.
Evans, M roseonana W. Gams & Gleeson and M. ovata H.Y.
Yip], while some were described as Mucor Fresen.[e.g. M.
ramannianus Möller = Mortierella ramanniana (Möller)
Linnem.] (Wang et al. 2015). According to Sugiyama et al.
(2003), two morphological criteria were essential for the
transfer of specimens from Mortierella to Umbelopsis:
1—the small but distinct columellae formed in Umbelopsis did not correspond to the absent or extremely reduced
13
8
columellae of other species of Mortierella; 2—the absence
of zygospores in Umbelopsis. At the time, Umbelopsis taxa
were classified into the sections Mortierella pusilla Oudem.
and M. isabellina, the subgenus Micromucor Malchevsk. and
the genera Micromucor and Umbelopsis. Meyer and Gams
(2003) included all the taxa of Micromucor in Umbelopsis,
based on restriction fragment length polymorphism (RFLP)
analysis, including sequences from the rDNA ITS-1 region.
Umbelopsis Amos & H.L. Barnett, Mycologia 58(5): 807
(1966)
Notes: The genus Umbelopsis, erected by Amos and
Barnett (1966) to accommodate U. versiformis (initially
positioned within the Deuteromycetes Sacc.), comprises
17 species (Yip 1986; Meyer and Gams 2003; Sugiyama
et al. 2003; Mahoney et al. 2004; Wang et al. 2014, 2015;
Crous et al. 2017; Wanasinghe et al. 2018). Members of
Umbelopsis produce velutinous colonies and sporangia with
colors ranging from ocher to red and pink, except U. nana,
which produces hyaline sporangia (Sugiyama et al. 2003).
The morphological classification of Umbelopsis specimens
is based mainly on colony color, branching pattern of the
sporangiophores, pigment production capacity, the size and
shape of sporangia, columellae and sporangiospores, as well
as the types of chlamydospores formed (Sugiyama et al.
2003; Wang et al. 2015). The delimitation of species of this
genus has also been based on physiological and genetic criteria (Meyer and Gams 2003; Sugiyama et al. 2003; Ogawa
et al. 2011; Wang et al. 2015; Spatafora et al. 2016; Crous
et al. 2017; Wanasinghe et al. 2018).
Umbelopsis heterosporus C.A. de Souza, D.X. Lima & A.L.
Santiago, sp. nov.
Index Fungorum number: IF556384; Facesoffungi number: FoF 06082; Fig. 1
Etymology: Referring to the variation in the shape and
size of the sporangiospores.
Holotype: URM 7882.
Colonies velvet, zoned, first pinkish then turning pale
brownish, presenting cytoplasmic oil droplets, exhibiting
low and slow growth (7.5 cm diam and 1–2 mm in height)
after seven days on PDA at 25 °C. Reverse cream to buff with
irregular margins. Mycelium exhibits randomly distributed
globose, subglobose and doliform swellings, 10–22 μm diam,
some presenting dilated rhizoid-like hyphae. No odour. Sporangiophores erect, some slightly curved, smooth-walled, hyaline, arising from the aerial mycelia and from a swollen portion, simple, cymose or occasionally sympodially branched,
2.5–5 μm diam and 135–340 μm long. Up to five septa may
be formed below the columella. Sporangia first pinkish then
becoming brownish vinaceous, globose, subglobose 13–30
μm diam, smooth-walled, evanescent, some leaving a collar.
Columellae hyaline, smooth-walled, globose, subglobose,
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Fungal Diversity (2020) 104:1–266
5–10 μm diam, subglobose to applanate, 2.5–3.5 × 2.5–6
μm, subglobose with a flattened end, 2.5–7.5 × 2.5–10 μm,
some irregular in shape, 3.5–8 μm. Some columellae inconspicuous, up to 1 μm diam. Sporangiospores variable in
shape and size, hyaline, smooth and thick-walled, ellipsoid
(1.5–)2.5–4 × 4.5–7.5(–9) μm, some cylindrical 2.5–4 × 4.5–8
μm, reniform, 2.5–4 × 3.5–7.5 μm, angular, 2.5–6 μm diam,
some irregular in shape, 3.5–5 × 5–11 μm. Chlamydospores
present in the aerial mycelium, globose, subglobose and ovoid
17–60 μm diam. Zygosporangia not observed.
Culture characteristics: On PDA. At 5 °C-no growth.
At 10 °C-limited growth (1.5 cm diam after 7 days); poor
sporulation. At 15 °C-slow growth (3 cm diam after 7 days);
poor sporulation. At 20 °C-better growth than at 15 °C (5.5
cm diam in 7 days); excellent sporulation. Mostly sporangiophores with simple branches. At 25 °C-better growth (7.5 cm
diam in 7 days); excellent sporulation. At 30 °C-no growth.
The growth of Umbelopsis on MEA was slightly slower than
on PDA at all the temperatures tested.
Material examined: BRAZIL, Brejo da Madre de Deus,
Pernambuco State (8º12′41.5" S, 36º23′73" W), in soil samples, 6 November 2015, leg. C.A.F de Souza (URM 7882,
holotype; URM 8082, ex-type).
GenBank numbers: ITS: MK804504, MK804505; LSU:
MK809511, MK809512.
Notes: According to our phylogenetic and morphophysiological analyses, Umbelopsis heterosporus exhibits wellsupported genetic datasets and morphological characteristics
that differentiate it from other species in the genus. It is morphologically distinguished from other species as it produces
sporangiospores and columellae that vary in shape and size,
including some that are bizarre in shape. The concatenated
phylogenetic tree (ITS and LSU rDNA sequences) of Umbelopsis revealed a close phylogenetic relationship between
U. heterosporus and U. gibberispora M. Sugiy., Tokum.
& W. Gams. (Fig. 2). Morphologically, the branching pattern of the sporangiophores described in U. gibberispora,
with several sympodial branches that arise in succession,
is distinguished from the simple, cymose or occasionally
sympodially branched sporangiophores observed in U. heterosporus. Additionally, colonies of U. heterosporus are
at first pinkish then brownish vinaceous, in contrast to the
colonies of U. gibberispora which are colorless to white
(Sugiyama et al. 2003). U. heterosporus produces ellipsoid,
cylindrical, reniform, and angular sporangiospores, some of
which are bizarre in shape and thick-walled, which differ
from the ellipsoid and one side thick-walled (hump-shaped)
sporangiospores observed in U. gibberispora. Moreover,
U. heterosporus produces globose, subglobose, subglobose
to applanate, subglobose with a flattened end and irregular
columellae, as well as some that are inconspicuous, differing
from the subglobose to flattened columellae displayed in U.
gibberispora (Sugiyama et al. 2003).
Fungal Diversity (2020) 104:1–266
Fig. 1 Microscopic structures of Umbelopsis heterosporus (URM
7882, holotype). a Colony surface after seven days at 25 ºC on PDA.
b Simple sporangiophore with globose sporangium. c, d Branched
9
sporangiophore with sporangia. e Simple sporangiophore with inconspicuous columella and collarette (arrow). f–g Simple sporangiophore
with columella. h Sporangiospores. Scale bars: 25 μm
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10
Fungal Diversity (2020) 104:1–266
Fig. 2 Phylogram generated from Bayesian inference analysis based
on combined LSU and ITS sequence data from 16 representative members of Umbelopsis. Mortierella verticillata (CBS 22058)
was used as outgroup. Branches with later Bayesian probabilities
(PP) ≥ 0.95 and bootstrap support values for maximum likelihood
(ML) analysis ≥ 70 where placed above or below nodes. Ex-type
strains are in bold. The newly generated sequences are indicated in
blue
Ascomycota
Dothideomycetes
Jayawardena et al. 2019; Philips et al. 2019). However, due
to the lack of morphological and ecological features that can
be used for segregation, Liu et al. (2016) expressed some
doubts about the inclusion of this many families in Botryosphaeriales. Taking morphology, phylogeny and evolutionary divergence times into account, Phillips et al. (2019)
retained only six families in the Botryosphaeriales. Species
in this order are cosmopolitan in nature and have a wide host
range (Phillips et al. 2019).
Botryosphaeriales C.L. Schoch, Crous & Shoemaker
Notes: Botryosphaeriales was erected by Schoch et al.
(2006) to accommodate the single family Botryosphaeriaceae. With the increased availability of molecular data,
eight more families have been included in this order (Minnis
et al. 2012; Slippers et al. 2013; Wyka and Broders 2016;
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Fungal Diversity (2020) 104:1–266
Aplosporellaceae Slippers, Boissin & Crous, Stud. Mycol.
76(1): 41 (2013)
Notes: Aplosporellaceae was erected by Slippers et al.
(2013) to accommodate two genera, Aplosporella Speg. and
Bagnisiella Speg. (Hyde et al. 2013; Ekanayaka et al. 2016).
Another new genus, Alanomyces was added by Sharma
et al. (2017), therefore three genera are currently accepted
in Aplosporellaceae (Sharma et al. 2017; Wijayawardene
et al. 2018; Index Fungorum 2020).
Aplosporella Speg., Anal. Soc. Cient. Argent. 10(4): 157
(1880)
Notes: Aplosporella is a species-rich genus comprising
339 epithets (Index Fungorum 2020). Aplosporella was
established by Spegazzini (1880), with A. chlorostroma
Speg. as the type species, and now it comprises six species (Hyde et al. 2013; Slippers et al. 2013; Ekanayaka et al.
2016; Sharma et al. 2017). Species of Aplosporella are circumscribed by having multilocular pycnidial conidiomata
embedded in stromatic tissues opening by a communal ostiole. The conidia are aseptate, ellipsoid to subcylindrical, initially hyaline, thin-walled and smooth, becoming pigmented,
thick-walled and spinulose (Sutton 1980; Hyde et al. 2011;
Liu et al. 2012; Ekanayaka et al. 2016; Du et al. 2017). Aplosporella species have been recorded from both temperate
and tropical countries (i.e. Australia, China, Namibia, South
Africa, Thailand) (Ekanayaka et al. 2016; Dou et al. 2017;
Du et al. 2017; Sharma et al. 2017).
Aplosporella prunicola Damm & Crous, Fungal Diversity
27(1): 39 (2007)
Index Fungorum number: IF504373; Facesoffungi number: FoF 04955; Fig. 3
Holotype: SOUTH AFRICA, Limpopo Province, Modimolle, from bark of small dead tree of Prunus persica var.
nucipersica, 30 August 2004, U. Damm, CBS-H 19848, culture ex-type CBS 121167 = STE-U 6326; Limpopo Province,
Modimolle, from the same specimen of P. persica var. nucipersica, STE-U 6327.
Saprobic on dead twig of Ficus septica Burm.f. Sexual
morph: Undetermined. Asexual morph: Conidiomata
500–1000 × 200–375 μm ( x̄ = 625 × 265 μm, n = 5), pycnidial, solitary, scattered, gregarious, immersed to semiimmersed, erumpent, globose to subglobose, coriaceous,
uni-loculate, dark brown to black, ostiolate. Peridium
30–40 μm wide, well developed, composed of several layers, inner two layers composed of thin-walled, hyaline, cells
of textura angularis, outer three layers composed of dark
brown, cells of textura angularis, becoming hyaline towards
the inner region. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 5–6 × 1–2 μm ( x̄ = 5.5 × 1.3
μm, n = 30), holoblastic, hyaline, cylindrical to doliiform,
smooth-walled. Conidia 12–16 × 6–9 μm ( x̄ = 14.5 × 7.8 μm,
11
n = 30), aseptate, broadly ellipsoid to subcylindrical, with
rounded ends, initially hyaline, becoming brown to dark
brown, smooth-walled.
Culture characteristics: Colonies on PDA reaching 25
mm diam after 2 weeks at 20–25 °C, colonies medium
sparse, circular, flat, surface slightly rough with edge entire,
margin well-defined, cottony to fairly fluffy with sparse
aspects, colony from above: white to cream at the margin
and the centre; reverse, white to grey margin and the centre;
mycelium white to grey with tufting; not producing pigments in PDA.
Material examined: CHINA, Yunnan Province, Xishuangbanna, Nabanhe, dead twig of Ficus septica (Moraceae),
20 November 2015, D.S. Tennakoon, DXH 019 (MFLU
17-0775, ibid. HKAS96335), living culture, MFLUCC
16-0791.
Host and habitat: Ficus septica (Moraceae), Prunus persica and P. nucipersica (Rosaceae).
GenBank numbers: ITS: MH974684.
Notes: As phylogenetic analysis (Fig. 4) and morphological characters examined largely overlap with those of
Aplosporella prunicola (Damm et al. 2007), we would like
to report our collection as a new host record.
Botryosphaeriaceae Theiss. & H. Syd., [as ’Botryosphaeriacae’], Annls Mycol. 16(1/2): 16 (1918)
Notes: Botryosphaeriaceae was established to include
three genera, namely; Botryosphaeria, Phaeobotryon and
Diobotryon by Theissen & Sydow (1918) with Botryosphaeria as the type genus. Currently 23 genera are accepted in
this family (Dissanayake et al. 2016). Molecular data play an
important role in defining species in this family as morphological characteristics are not sufficiently different for clear
separations (Phillips et al. 2013).
Diplodia Fr., Annales des Sciences Naturelles Botanique 1:
302 (1834)
Notes: The genus Diplodia was introduced by Montagne
in 1834 with Diplodia mutila (Fr.) as the type species. It is a
large genus with ca. 1000 species names recorded in Index
Fungorum. 31 of which are known from culture (Phillips
et al. 2013; Dissanayake et al. 2016; Gonzalez-Domínguez
et al. 2017; Yang et al. 2017). Species in this genus can
be pathogens, endophytes or saprobes on a wide range of
woody hosts with some being important pathogens causing severe damage to many important plant species (Phillips
et al. 2012).
Diplodia torilicola Harishchandra, Camporesi, A.J.L. Phillips & K.D. Hyde, sp. nov.
Index Fungorum number: IF 556364; Facesoffungi number: FoF 05980; Fig. 5
Etymology: Name reflects the host genus Torlilis.
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Fungal Diversity (2020) 104:1–266
Fig. 3 Aplosporella prunicola (MFLUCC 16-0791, new host
record). a Habit of conidiomata on twig. b Close-up of conidioma. c
Cross section of the conidioma. d Peridium. e–f Conidiogenous cells.
g–m Conidia. n Germinated spore. o Culture from above. p Culture
from below. Scale bars: c–d = 20 μm, e–n = 10 μm
Holotype: JZBH3140012.
Saprobic on dead aerial stem of Torilis arvensis. Sexual
morph: Undetermined. Asexual morph: Coelomycetous. Conidiomata on host 99–160 µm diam ( x̄ = 128 µm,
n = 10) pycnidial, solitary, scattered, globose, black, semiimmersed to immersed. Ostiole single, centrally located.
Conidiomata wall 30–40 µm wide ( x̄ = 34 μm, n = 10) at
both sides, multi-layered, brown to black cells of textura
angularis. Conidiophores not observed. Conidiogenous
cells cylindrical, thin-walled, hyaline 16–25 μm high × 3–9
μm wide ( x̄ = 16.8 × 4.6 μm, n = 20) producing a single
conidium at the apex. Conidia 17–22 μm long × 8–11 μm
wide ( x̄ = 21 × 8.8 μm, n = 40), initially hyaline soon becoming pigmented, dark brown when mature, aseptate, ellipsoid
to ovoid with central guttules.
Culture characteristics: Colonies on PDA attaining 90
mm diam within 3 days at 25 °C, with white moderately
aerial mycelium and white in reverse. Mycelia becoming
dense, cottony and black with age.
Material examined: ITALY, near Castrocaro Terme
in Forlì-Cesena Province, on dead aerial stem of
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13
Fig. 4 Phylogram generated from maximum likelihood analysis of
combined ITS and TEF1-α sequence data from species of Aplosporellaceae. Bootstrap support values for maximumlikelihood (ML) and
maximum parsimony (MP) ≥ 65% and Bayesian posterior probabili-
ties (PP) ≥ 0.90 are given above each branch respectively. The new
isolate is in blue. Ex-type strains are in bold. The tree is rooted with
Saccharata capensis (CBS 122693) and S. proteae (CBS 115206)
Torilis arvensis, 11 December 2017, Erio Camporesi,
JZBH3140012 (holotype).
GenBank numbers: ITS: MK625223; TEF1: MK640502.
Notes: Diplodia torilicola is most closely related to D.
crataegicola but forms a well-supported independent lineage
from it (Fig. 6). It differs by possessing larger conidia that
are ellipsoid to ovoid (21 × 8.8 µm) whereas D. crataegicola
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14
Fig. 5 Diplodia torilicola (JZB3140012, holotype). a Conidiomata
on the host tissue. b Section of the pycnidial wall. c Cross section of
conidiomata on host. d–e Conidiogenous cells. f–k Conidia. l Lower
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view of 7-day old culture. m Upper view of 7-day old culture. Scale
bars: a = 1000 μm, b–e = 20 μm, f–k = 10 μm
Fungal Diversity (2020) 104:1–266
15
Fig. 6 Phylogram generated from maximum likelihood analysis of combined ITS and TEF1 sequence data. Bootstrap support values ≥ 50% are
given above each branch respectively
has smaller (14 × 9 µm) globose to subglobose conidia (Ariyawansa et al. 2015). This is the first report of a Diplodia
species on Torilis arvensis (Farr and Rossman 2019).
Capnodiales Woron.
Notes: The order Capnodiales includes 15 families.
Neodevriesiaceae Quaedvl. & Crous, in Quaedvlieg, Binder,
Groenewald, Summerell, Carnegie, Burgess & Crous, Persoonia 33: 24 (2014)
Notes: Neodevriesiaceae was established and placed in
Capnodiales by Quaedvlieg et al. (2014) to accommodate a
monotypic genus Neodevriesia. It includes the species that
are intolerant to heat and lack chlamydospore formation.
Later, the boundaries of Neodevriesia were extended to
include taxa similar to Devriesia in the Neodevriesiaceae
(Crous et al. 2015). Two genera Devriesia and Neodevriesia
are accepted in this family. Members of this family are foliicolous, saprobes and plant pathogens and are reported from
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16
different habitats such as terrestrial, freshwater and marine
(Quaedvlieg et al. 2014).
Neodevriesia Quaedvl. & Crous, in Quaedvlieg et al., Persoonia 33: 24 (2014)
Notes: The genus Neodevriesia was firstly treated as a
member of the Teratosphaeriaceae (Ruibal et al. 2011).
Later Quaedvlieg et al. (2014) placed it in a new family Neodevriesiaceae typified by Neodevriesia hilliana.
Neodevriesia is characterized by a sexual morph with
immersed or superficial pseudothecial globose ascomata,
8-spored, bitunicate, thick-walled, obovoid to broadly ellipsoid, short pedicellate, straight or slightly curved, apically
rounded asci, and ascospores that are ellipsoid, hyaline to
dark brown, and an asexual morph with medium brown and
unbranched conidiophores, rarely septate conidia, short and
mostly unbranched conidial chains and an absence of chlamydospores (Quaedvlieg et al. 2014; Crous et al. 2015). This
genus includes 25 accepted species.
Neodevriesia manglicola Devadatha, V.V. Sarma & E.B.G.
Jones, sp. nov.
Index Fungorum number: IF556225; Facesoffungi number: FoF 07057; Fig. 7
Etymology: In reference to the mangrove habitat where
the fungus was collected.
Holotype: AMH-10019.
Saprobic on decaying wood of Rhizophora mucronata. Ascomata 250–350 µm high, 200–500 µm wide
( x̄ = 285 × 297 µm, n = 20), globose or subglobose, superficial to semi-immersed, carbonaceous, short papillate
and ostiolate. Ostiole 45–100 µm long, 50–60 µm diam
( x̄ = 74 × 55 µm, n = 5), periphysate. Peridium two layered,
30–70 µm ( x̄ = 51 µm, n = 5), thickened, outer layer of small
pseudoparenchymatous cells, brown, inner layer of hyaline
polygonal cells. Hamathecium composed of cellular, hyphallike, septate pseudoparaphyses. Asci 60–100 × 15–25 µm
( x̄ = 76 × 18 µm, n = 15), 8-spored, bitunicate, thick-walled,
clavate to cylindrical, short pedicellate, apically rounded.
Ascospores 8–17 × 3–10 µm ( x̄ = 14 × 8 µm, n = 25), overlapping uniseriate, ellipsoid, hyaline to dark brown, muriformed appearance due to longitudinal pseudosepta with oil
globules, 0–3-transverse septa and 1 longitudinal septum in
each cell, fully matured ascospores becoming brown with 3
dark brown banded septa, lacking mucilaginous sheaths or
appendages.
Material examined: INDIA, Tamil Nadu, Parangipettai mangroves (11.59°N, 79.5°E), on decaying wood of
Rhizophora mucronata (Rhizophoraceae), 23 April 2018,
B. Devadatha, AMH-10019 (holotype), living culture,
NFCCI-4382.
GenBank numbers: ITS: MN061371; LSU: MN061356.
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Culture characteristics: Ascospores germinating on 50%
sea water agar within 24 h, with germ tubes developed from
both end cells and sides of ascospores. Colonies on MEA
slow growing, reaching 10–20 mm diam, after 15 days of
incubation at 25 °C, surface pale brown to dark brown on
the surface, reverse brown, velvety, irregular and umbonate.
Notes: Our phylogeny of LSU and ITS sequences including taxa from Neodevriesia, Devriesia and other related
genera revealed that N. manglicola is sister to N. queenslandica with significant support (ML 78% and 1.00 PP), and
then nested with N. shakazului in a monophyletic clade with
strong support (ML 100%, MP 99% and 1.00 PP) (Fig. 8).
Recently two new species, N. cladophorae and N. grateloupiae, were reported from marine environments (Wang
et al. 2017). These two species are phylogenetically nested
distant to N. manglicola, and are known to produce only
asexual morphs. N. manglicola is the first report of a species
having a sexual morph in the genus though it remained nonsporulating in artifical media. We introduce N. manglicola
as a new species in Neodevriesia.
Pleosporales Luttr. ex M.E. Barr
Notes: See Zhang et al. (2009, 2012) and Hyde et al.
(2013).
Coniothyriaceae W.B. Cooke, Revta Biol., Lisb. 12: 289
(1983)
Notes: Cooke established Coniothyriaceae in 1983 and it
was considered as a synonym of Leptosphaeriaceae by Kirk
et al. (2008). However, in recent studies (Hyde et al. 2013;
Quaedvlieg et al. 2013; Wijayawardene et al. 2016, 2018,
2020) Coniothyriaceae is accepted as a distinct family in
the order Pleosporales. This family comprises four genera,
Coniothyrium, Hazslinszkyomyces, Ochrocladosporium and
Staurosphaeria (Wijayawardene et al. 2018, 2020).
Coniothyrium Corda, Icon. Fung. (Prague) 4: 38. 1840
Notes: Coniothyrium and Coniothyrium-like species are
polyphyletic within the Pleosporales (Hyde et al. 2013; Verkley et al. 2014; Pem et al. 2020). Coniothyrium sensu stricto
was placed within Coniothyriaceae, Pleosporales (Verkley
et al. 2014; Wijayawardene et al. 2016, 2017a; Thambugala
et al. 2017).
Coniothyrium triseptatum Dayarathne, Thyagaraja & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556425; Facesoffungi number: FoF 06036; Fig. 9
Etymology: Epithet refers to the septation of the
ascospores.
Holotype: MFLU 19-0758.
Saprobic on bark of unidentified plant, forming black,
elongated, raised structures, ascostromata opening through
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Fig. 7 Neodevriesia manglicola (AMH-10019, holotype). a–b Ascomata superficial to semi-immersed on decaying wood. c–d Vertical
sections of ascomata. e Peridium. f Filamentous pseudoparaphyses.
g–l Immature and mature asci. m–q Immature and mature ascospores.
r Germinating ascospores. Scale bars: c–d = 100 μm, e–i = 10 μm,
j = 50 μm, k–r = 10 μm
slits along the length. Sexual morph: Ascostromata
0.15–0.25 × 0.1–0.15 mm, 0.1–0.15 mm high, solitary,
immersed, fusiform. Ascomata 200–280 × 200–380 μm, 3–4
perithecial, perithecia immersed within the ascostromata,
subglobose or globose with a flattened base, dark brown
to black, with centrally located short ostiole. Peridium
5.5–8 μm wide, thick- to thin-walled, of unequal thickness, poorly developed at the base and merging with host
tissue, composed of several layers of outer brown to inner
hyaline, pseudoparenchymatous cells of textura prismatica.
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Fig. 8 Phylogram generated
from maximum parsimony
analysis based on LSU and ITS
gene regions. Bootstrap support
values for maximum likelihood
(ML, green), maximum parsimony (MP, blue) ≥ 70% and the
values of Bayesian posterior
probabilities (PP, purple) ≥ 0.95
are given above each branch,
respectively. The new isolate is
in blue. The tree is rooted with
Passalora zambiae CBS112971
Hamathecium of dense, 1–2 μm wide, filamentous, septate, cellular pseudoparaphyses, not constricted at the septum. Asci 100–150 × 6–10 μm ( x = 130 × 8 μm, n = 15),
8-spored, bitunicate, fissitunicate, cylindric-clavate, with
broad flat pedicel, apex rounded. Ascospores 15–18 × 4–6
μm ( x = 16 × 5 μm, n = 20), overlapping uni-seriate, hyaline,
fusiform, with rounded to acute ends, 3-septate, slightly constricted at the middle septum, middle cells are wider than
13
two apical cells, smooth-walled, guttulate. Asexual morph:
Undetermined.
Material examined: CHINA, Yunnan Province, Shangri
La, 27°55′54.9″ N, 099°34′39.0″ E, alt. 4045 m, on bark of
an unidentified host, 12 September 2018, Vinodhini Thyagaraja, (MFLU 19-0758, holotype).
GenBank numbers: ITS: MK791297; LSU: MK787306;
RPB2: MK791298.
Fungal Diversity (2020) 104:1–266
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Fig. 9 Coniothyrium triseptatum (MFLU 19-0758, holotype). a–c Ascostromata. d Section of ascomata. e Section through peridium. f Section
through neck region. g–i Asci. j Pseudoparaphyses. k–l Ascospores. Scale bars: d = 100 μm, c, g–j = 20 μm, f, k, l = 10 μm
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Fig. 10 Phylogram generated
from maximum likelihood
analysis based on a combined
LSU, ITS and RPB2 sequence
dataset of selected taxa. Tree
was rooted with Phaeosphaeria oryzae (CBS 110110).
Bayesian posterior probabilities
(PP) ≥ 0.80 and maximum
likelihood bootstrap (ML) values ≥ 65% are given above the
nodes. The scale bar indicates
0.03 changes. The ex-type strain
is in bold and new isolate is in
blue
Notes: The Paraphaeosphaeria-like sexual morph was
firstly recorded in Coniothyrium by Quaedvlieg et al. (2013).
Subsequently, Thambugala et al. (2017) introduced the asexual morph C. chiangmaiense Goonas., Thambugala & K.D.
Hyde which can be clearly distinguished from C. triseptata
by having uniseptate ascospores (Thambugala et al. 2017),
while C. triseptatum has 3 septa. Our phylogenetic analyses
with concatenated LSU, ITS and RPB2 sequence data confirmed its stability within Coniothyriaceae with high statistical support (0.94 PP) basal to C. sidae (CBS 135108)
(Fig. 10). However, the asexual morph of C. sidae possesses
(3–)5-septate ascospores and are longer than those of C.
triseptata (20–24 μm). Base pair differences of C. sidae and
C. triseptatum for ITS are 68 bp out of 452 bp without gaps
(15%) and 31 bp out of 300 bp without gaps (10%) for RPB2.
Didymellaceae Gruyter, Aveskamp & Verkley, Mycol. Res.
113(4): 516 (2009)
Notes: Didymellaceae was introduced for the ‘‘Didymella
clade’’ to accommodate the type species Didymella exigua,
Phoma and Phoma-like genera (Jayasiri et al. 2017). There
are 27 genera presently in Didymellaceae based on morphology and phylogeny (Hyde et al. 2013; Wijayawardene et al.
2014, 2018).
Neodidymelliopsis Qian Chen & L. Cai, Stud. Mycol. 82:
207 (2015)
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Notes: The genus Neodidymelliopsis was erected by Chen
et al. (2015a, b), with Neodidymelliopsis cannabis (G. Winter) Q. Chen & L. Cai as the type species (Chen et al. 2015a,
b; Hyde et al. 2016; Thambugala et al. 2016).
Neodidymelliopsis salviae Brahmanage, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF556376; Facesoffungi number: FoF 06125; Fig. 11
Etymology: Name reflects the host genus.
Holotype: JZBH3470001.
Saprobic on dead and dying twigs and branches of Salvia
officinalis. Sexual morph: Undetermined. Asexual morph:
Coelomycetous. Conidiomata 120–375 µm diam ( x = 240
µm, n = 5), pycnidial, solitary, scattered, globose, black,
semi-immersed to immersed. Pycnidial wall 28–45 μm wide,
pseudoparenchymatous, 2–3-layered, composed of oblong
to isodiametric cells. Conidiogenous cells phialidic, ampulliform, hyaline. Conidia 6.3–9.3 × 3–4 µm ( x = 7.8 × 3.6 µm,
n = 40), oblong to ellipsoid, initially hyaline becoming light
brown when mature, aseptate occasionally with larger 1-septate conidia, smooth-walled.
Material examined: ITALY, Carpena – Forlì (Province of
Forlì-Cesena), on Salvia officinalis (Lamiaceae), 19 March
2018, Camporesi Erio IT 3785 (JZBH3470001, holotype).
GenBank numbers: ITS: MK651827; LSU: MK651828;
RPB2: MK736714.
Fungal Diversity (2020) 104:1–266
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Fig. 11 Neodidymelliopsis salviae (JZBH3470001, holotype).
a Submerged conidiomata on
host. b Pycnidial wall on host.
c Section through conidiomata.
d, e Conidiogenous cells and
conidial attachments. f Conidia.
Scale bars: c = 100, b = 20 µm,
d, e = 10, f = 5 µm
Notes: Neodidymelliopsis salvia can be morphologically assigned to Neodidymelliopsis by having pycnidial,
conidiomata that are globose, phialidic, hyaline, smooth,
ampulliform conidiogenous cells and oblong to ellipsoid, hyaline to pale brown, usually aseptate or occasionally 1-septate conidia (Jayasiri et al. 2017). Phylogenetic
analyses based on a combined LSU, ITS, RPB2 and β-TUB
sequence data showed that N. salvia and N. sambuci formed
a well-supported (85% ML and 1.00 PP) lineage within
Neodidymelliopsis (Fig. 12), but morphologically N. sambuci can be distinguished by smaller conidia (4–7 × 2–3.5
µm, Hyde et al. 2019). Based on both morphology and
phylogeny, we introduce N. salvia as a new taxon within
Neodidymelliopsis.
Neodidymelliopsis urticae Manawas., Camporesi & K.D.
Hyde, sp. nov.
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Fig. 12 Maximum likelihood tree based on a combined LSU, ITS,
RPB2 and β-TUB sequence dataset. 38 strains of Neodidymelliopsis
are included with Xenodidymella applanata (CBS 205.63) as the outgroup taxon. Maximum likelihood bootstrap (ML) values ≥ 65% and
Bayesian posterior probabilities (PP) ≥ 0.80% are given above the
nodes. The scale bar indicates 0.006 changes. The ex-type strains are
in bold and new isolates are in blue
Index Fungorum number: IF556376; Facesoffungi number: FoF 06125; Fig. 13
Etymology: Name reflects the host genus.
Holotype: MFLU18-0877.
Saprobic or necrotrophic on dead and dying twigs and
branches of Urtica dioica (L.). Sexual morph: Undetermined. Asexual morph: Coelomycetous. Conidiomata on
host 78–286 µm diam ( x = 157 µm, n = 15), pycnidial, solitary, scattered, globose, black, semi-immersed to immersed;
Conidiophores not observed. Conidiogenous cells not
observed in host. Conidiogenous cells in culture hyaline,
flask-shaped, ampulliform. Conidia on host 4–9 × 1–3 µm
( x = 5 × 2 µm, n = 40), oblong to ellipsoid, hyaline, aseptate,
smooth-walled; in culture, 4–7 × 1–3 µm ( x = 5.5 × 2.6 µm,
n = 40). Conidial exudates were not observed.
Culture characteristics: Colonies on PDA reach 65 mm
diam after 7 days at 25 °C, with circular, entire edge, raised
olivaceous grey aerial mycelium, surface floccose to woolly.
Material examined: ITALY, Carpena—Forlì (Province of
Forlì-Cesena), on Urtica dioica L. (Urticaceae), 6 March
2018, Camporesi Erio, MFLU 18-0877 (holotype), ex-type
living culture, MFLUCC 19-0218.
GenBank numbers: ITS: MK651827; LSU: MK651828;
RPB2: MK736714.
Notes: Neodidymelliopsis urticae was isolated from dead
and dying twigs and branches of Urtica dioica in ForlìCesena, Italy. The characteristics of the spores and colony
(Fig. 13) match the concept of Neodidymelliopsis. Phylogenetic analysis based on a combined ITS, LSU, RPB2
and β-TUB sequence data-set confirms that the present
taxon belongs in Neodidymelliopsis within Didymellaceae
(Fig. 12). N. urticae developed an independent clade in the
genus with moderate support (67% ML and 1.00 PP) and is
closely related to N. salviae and N. sambuci. Morphologically, N. urticae can be distinguished from N. salviae and N.
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23
Fig. 13 Neodidymelliopsis urticae (MFLU 18-0177, holotype). a
Material examined. b Submerged conidiomata on host. c Pycnidial
wall on host. d–e Conidia on host. f–g Developing pycnidial wall on
culture. h Conidiogenous cell. i-l Conidia on host. m Septate mycelia.
n Upper side of colony on PDA. o Reverse view of 7 days old culture
on PDA. Scale bars: a–c = 100 µm, d–m = 10 µm
sambuci by its smaller conidia (7.8 × 3.6 µm in N. salviae,
6 × 2 µm in N. sambuci, Hyde et al. 2019).
Notes: The latest treatments are by Thambugala et al.
(2014) with updates by Hyde et al. (2017) and Wanasinghe
et al. (2017).
Lophiostomataceae Sacc.[as ’Lophiostomaceae’], Syll.
Fung. (Abellini) 2: 672 (1883)
Magnopulchromyces L.B. Conc., Gusmão & R.F.
Castañeda, gen. nov.
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24
Index Fungorum number: IF556378; Facesoffungi number: FoF 06109.
Etymology: From the Latin: Magno- great, maximum, + pulchre beautifully, excellently, + -myces from the
Greek, meaning fungus.
Type species: Magnopulchromyces scorpiophorus L.B.
Conc., Gusmão & R.F. Castañeda
Saprobic on decaying leaves of unidentified plant.
Asexual fungi, Conidiophores macronematous, mononematous, scorpioid growth, septate, brown. Conidiogenous
cells monoblastic intercalary, pale brown or greyish brown.
Conidial secession schizolytic. Conidia acropleurogenous,
complex, lenticular-staurospore, somewhat stellate or irregular staurospore, variegated, multicellular, composed of a
primary cell, one or several, secondary cells, several tertiary cells and several satellite cells hemispherical, straight
or slightly curved, greyish brown or very pale brown. Sexual
morph: Undetermined.
Notes: Magnopulchromyces resembles superficially the
monotypic genera Turturconchata J.L. Chen, T.L. Huang &
Tzean and Venustisporium R.F. Castañeda & Iturr. by the
multicellular, lenticular, complex conidia, with holoblastic
production and schizolytic secession (Castañeda-Ruiz and
Iturriaga 1999; Chen et al. 1999). Turturconchata is different
from Magnopulchromyces in having conidiophores single or
clustered, curved to spiral and conidia without an eccentric
protuberating pedicel. Venustisporium is distinguished by
developed sporodochia with inconspicuous prostrate conidiophores, mostly reduced to conidiogenous cells, and conidia
with a layer of uncinated cells, needle-like to horn-like projections around the periphery. Magnopulchromyces is further differentiated from Turturconchata and Venustisporium
by having a developed scorpioid growth of conidiophores
and the complex conidia.
Magnopulchromyces scorpiophorus L.B. Conc., Gusmão
& R.F. Castañeda, sp. nov.
Index Fungorum: IF556428; Facesoffungi number: FoF
06110; Fig. 14
Etymology: From the Latin: scorpiophorus-, referring to
the scorpioid unilateral, branched conidiophores.
Holotype: HUEFS 234842.
Saprobic on decaying leaves. Asexual morph: Colonies
on the natural substrate effuse, granulose, brown. Mycelium
mostly superficial, composed of branched, septate, brown
hyphae, 3–4 µm diam. Conidiophores macronematous,
mononematous, erect or prostrate, scorpioid-unilateral
branched, brown below, pale dirty brown or pale greyish
brown toward the apex, multiseptate, up to 300 µm long,
5–8 µm wide. Conidiogenous cells monoblastic, cuneiform,
sub-doliiform, intercalary and terminal, pale greyish brown
to pale brown, smooth, 5–7 × 5–8 µm. Conidial secession
schizolytic. Conidia acropleurogenous, complex, lenticular
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staurospore, somewhat stellate or irregular staurospore,
variegated, smooth, 25–35 × 24–31 µm, multicellular, composed of: (i) a more or less central primary cell, dark brown,
6.5–11 × 9–13 µm; (ii) 1–4 secondary cells brown, or dark
brown, 4.5–7 × 5–10 µm, lateral, radial or cruciate arranged
beside or surrounding the primary cells; (iii) 3–8 tertiary
cells pale brown, 4.5–6 × 6–8 µm, beside the secondary cells;
(iv) 6–13 satellite cell conical, trapezoid, mammiform or
hemispherical, straight or slightly curved, greyish brown or
very pale brown. Sexual morph: Undetermined.
Culture characteristics: Cultures growing on CMA reaching 30 mm diam after 30 days at 25 °C; moderate aerial
mycelium, circular, umbonate, regular edges, dark brown.
Material examined: BRAZIL, Bahia State, Pindobaçu,
Serra da Fumaça, 10°39′ S, 40°22′ W, on decaying leaves of
unidentified plant, 26 July 2016, L.B. Conceição, (HUEFS
234842, holotype); ex-type living culture CCLAMIC
206/16.
GenBank numbers: ITS: MG594647; LSU: MG594648;
TEF1: MG597248.
Notes: Maximum likelihood tree based on a combined
LSU, ITS and TEF1-α sequence dataset shows the phylogenetic position of Magnopulchromyces scorpiophorus
(Fig. 15).
Paradictyoarthriniaceae Doilom, Jian K. Liu & K.D. Hyde,
in Liu et al., Fungal Diversity: 10.1007/s13225-015-0324-y,
[133] (2015)
Notes: Paradictyoarthriniaceae was established as a family in the Pleosporales by Liu et al. (2015a, b, c) to accommodate Paradictyoarthrinium in Pleosporales based on its
unique morphology and distinct lineage in the phylogenetic
analysis. Members of this family are saprobes on dead wood
and occur in a wide range of habitats such as terrestrial,
aquatic and marine. They are widely distributed and reported
from South Africa and tropical countries such as Thailand
and India.
Paradictyoarthrinium Matsush., Matsush. Mycol. Mem. 9:
18 (1996)
Notes: The genus Paradictyoarthrinium, typified by P.
diffractum, was firstly reported by Matsushima (1996). This
taxon was subsequently reported by Prabhugaonkar and
Bhat (2011) from palm litter from India and later by Liu
et al. (2015a, b, c). The genus is characterized by superficial,
gregarious, black, powdery fruitbodies and macronematous
conidiophores with unevenly dictyoseptate, subglobose to
ellipsoid dark brown conidia. Paradictyoarthrinium includes
four accepted species which are supported by both morphological and molecular data (Index Fungorum 2020).
Paradictyoarthrinium diffractum Matsush., Matsush.
Mycol. Mem. 9:18 (1996) Fig. 16
Fungal Diversity (2020) 104:1–266
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Fig. 14 Magnopulchromyces
scorpiophorus (HUEFS234842,
holotype). a–b Detail of
conidiogenous cells and conidiophores. c Conidia. d Scorpioid
growth development on natural
substrate. Scale bars: 20 µm
Saprobic on decaying Avicennia marina wood. Sexual
morph: Undetermined. Asexual morph: Colonies on natural substrate superficial, black, powdery, scattered, black,
gregarious. Conidiophores 15–85 × 2–4 μm ( x = 31 × 2.6
μm, n = 5), macro- to micronematous, short, erect to slightly
curved, black, slightly constricted at the septa, arising from
hyphae. Conidiogenous cells blastic, integrated, mostly
terminal and determinate. Conidia 10–35 × 10–25 μm
( x = 22 × 18 μm, n = 30), 2 to many, occasionally unevenly
dictyoseptate, circular to irregular, dark brown to black on
maturity, verrucose, solitary or forming in branched chains,
hardly separating, variable in size and shape, mostly as tetrads on natural substrata.
Culture characteristics: Conidia germinating on 50% sea
water agar within 24 h, with germ tubes developed from all
around. Colonies on MEA moderately growing, reaching
10–25 mm diam on 7th day, after 25 days of incubation
reaching 50–65 mm at 25 °C, surface initially hyaline to pale
grey reaching dark grey to dark brown, reverse dark brown,
circular and zonate.
Material examined: INDIA, Tamil Nadu, Tiruvarur,
Muthupet mangroves on decaying intertidal wood of Avicennia marina (Acanthaceae), 28 November 2015, B. Devadatha (AMH-10161), living culture, NFCCI-4665.
GenBank numbers: ITS: MN061370; LSU: MN061354
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Fig. 15 Maximum likelihood tree based on a combined LSU, ITS and
TEF1-α sequence dataset. Numbers above branches indicate maximum likelihood bootstrap values ≥ 70%. Thickened branches indicate
Bayesian posterior probabilities ≥ 0.95. The tree is rooted with Mela-
nomma pulvis-pyrius (CBS 124080). Magnopulchromyces scorpiophorus is highlighted in yellow. Culture collection numbers are given
after taxon names
Notes: Our preliminary morphological studies and
BLAST search analyses based on LSU and ITS sequence
data showed that our taxon is 99% similar to Paradictyoarthrinium diffractum. Furthermore, the combined LSU and
ITS phylogenetic analyses revealed that it nests along with
other P. diffractum strains in a monophyletic clade with a
significant support from ML 91%, MP 86% and 1.00 PP
(Fig. 17). Morphologically our collection from natural specimen tends to produce tetrads of conidia in powdery form
and lacks conidiophores and pleomorphic conidia (Fig. 16a,
o–p). However, an isolated culture sporulated after 30 days
incubation and had conidiophores bearing two or four pleomorphic dictyoseptate, circular to irregular, dark brown to
black at maturity in branched chains, and including tetrads,
conidia (Fig. 16b–n). We do not know the exact ecological
factors that induce P. diffractum (AMH-10161) to produce
only tetrads of conidia on natural host substrata in contrast
to the isolated culture (NFCCI-4665) which tends to produce pleomorphic conidia. Our collection of P. diffractum
(NFCCI-4665) shares identical characters with P. diffractum (GUFCC-15514) reported by Prabhugaonkar and Bhat
(2011) on a dead spathe of Cocos nucifera from Goa, India.
Isaka et al. (2015) discovered two new cytotoxic hydroanthraquinones, paradictyoarthrins A (1) and B (2) from P.
diffractum (BCC 8704) collected from unidentified mangrove wood in Laem Son National Park, Ranong Province,
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Fig. 16 Paradictyoarthrinium diffractum (AMH-10161, a new
host/geographical record). a Brown to black superficial colonies on
decaying wood. b–c Colonies on MEA after 30 days. d–f, h–k Con-
27
idiophores and conidiogenous cells with conidial connections. j, l, m,
o–p Conidia. Scale bars: d–k, o = 50 μm. l,p = 10 μm
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◂Fig. 17 Phylogram generated from maximum likelihood analysis
based on combined LSU and ITS gene regions of Paradictyoarthrinium and related taxa. Bootstrap support values for maximum
likelihood (ML, green), maximum parsimony (MP, blue) ≥ 75% and
the values of Bayesian posterior probabilities (PP, purple) ≥ 0.95 are
given above each branch, respectively. The new collection is in blue.
The tree is rooted with Melanomma pulvis-pyrius CBS 124080
Thailand. Recently, Doilom et al. (2017) recorded P. diffractum on Tectonia grandis. The present record is the first
report of this species from Avicennia marina and is a new
record for marine fungi of India. Hence, the details of its
morphology and sequence data were provided in this study.
Phaeosphaeriaceae M.E. Barr, Mycologia 71(5): 948
(1979)
Notes: We follow the latest treatment and update accounts
of Phaeosphaeriaceae in Hyde et al. (2019, 2020a) and
Phookamsak et al. (2019). Eighty-three genera are accepted
in this family (Bakhshi et al. 2019; Hyde et al. 2019;
Karunarathna et al. 2019; Maharachchikumbura et al. 2019;
Phookamsak et al. 2019; Yang et al. 2019; Zhang et al. 2019;
Hongsanan et al. 2020).
Ophiosphaerella Speg., Anal. Mus. Nac. B. Aires, Ser. 3
12: 401 (1909)
Notes: We follow the latest treatment and updated
accounts of Ophiosphaerella in Phookamsak et al. (2014),
Ariyawansa et al. (2015) and Thambugala et al. (2017).
Ophiosphaerella chiangraiensis Phookamsak & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556471; Facesoffungi number: FoF 06128; Fig. 18
Etymology: The specific epithet “chiangraiensis” refers
to Chiang Rai Province, Thailand, where the holotype was
collected.
Holotype: MFLU 11-0246.
Saprobic on dead culm of grass. (Sexual morph): Ascomata 190–370 μm high (including papilla), 250–460 µm
diam, scattered, solitary, semi-immersed, visible as black,
shiny dot on the host surface, globose to subglobose, uniloculate, glabrous, ostiole central, with minute papilla, filled
with hyaline periphyses. Peridium 10–20 μm wide, of equal
thickness, composed of several layers of small, flattened
to broad, dark brown pseudoparenchymatous cells, paler
towards the inner layers, arranged in textura angularis to
textura prismatica. Hamathecium composed of numerous,
1.8–3.8 μm wide, filamentous, septate pseudoparaphyses,
anastomosing above the asci, embedded in a hyaline gelatinous matrix. Asci (125–)135–160(–180) × (6.5–)7–10 μm
( x = 152.5 × 8.6 μm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical to subcylindric-clavate, subsessile to short
pedicellate, with obtuse to furcate pedicel, apically rounded,
29
with well-developed ocular chamber, clearly seen at immature state. Ascospores (122–)140–160(–165) × 2–3(–4) μm
( x = 143.6 × 2.7 μm, n = 30), overlapping, in spiral, filiform,
tapering towards the lower cell, pale yellowish, curved,
(14–)15–16(–18)-septate, not constricted at the septa,
smooth-walled, lacking a mucilaginous sheath. Asexual
morph: Undetermined.
Culture characteristics: Colonies on PDA reaching 38–40
mm diam after 2 weeks at room temperature (20–30 °C).
Colony medium dense, circular, flattened, slightly raised,
surface smooth, with entire edge, floccose; from above,
white to cream; from below, cream to pale yellowish; not
producing pigmentation on agar medium.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Ta Sai, on dead culm of grass, 30 June 2011,
R. Phookamsak, RP0130 (MFLU 11–0246, holotype), extype living culture, MFLUCC 12–0007.
GenBank numbers: ITS: KM434272; LSU: KM434282;
SSU: KM434291; TEF1-α: KM434300; RPB2: KM434308.
Notes: Phylogenetically, strain MFLUCC 12–0007
formed a sister lineage with Ophiosphaerella agrostidis
(MFLUCC 11-0152) with full support (100% ML, 100%
MP) (Fig. 19). Comparisons of ITS and TEF1-α sequences
show that MFLUCC 12-0007 differs from O. agrostidis
(MFLUCC 11-0152) in 11/494 bp (2.2%) of ITS and 22/878
bp (2.5%) of TEF1-α. Therefore, we introduce MFLUCC
12–0007 as a new species, O. chiangraiensis. O. chiangraiensis is similar to O. agrostidis (see description of this
species in Câmara et al. 2000 and Phookamsak et al. 2014)
in having filiform, pale yellowish, (14–)15–16(–18)-septate
ascospores. However, O. chiangraiensis differs from O.
agrostidis by having slightly larger ascomata and ascospores,
and smaller asci (Câmara et al. 2000).
Pleomassariaceae M.E. Barr, Mycologia 71(5): 949 (1979)
Notes: Pleomassariaceae was established by Barr (1979)
with Pleomassaria as the type genus. The characters that
define the family are globose or depressed globose ascomata, a papillate ostiole which opens via a minute slit or
a small conical swelling in the host, cellular pseudoparaphyses, 8-spored, bitunicate, fissitunicate cylindrical or
clavate asci, ellipsoid or oblong 1-septate, multi-septate or
muriform, mostly distoseptate, brown ascospores usually
with a gelatinous sheath. Initially the genus comprised two
other genera Asteromassaria and Splanchnonema. Later,
the lichenized genus Eopyrenula was added by Barr (1993).
Pleomassaria morphologically differs from the other two
genera in producing muriform and slightly asymmetrical
ascospores with a sub-median primary septum. Later, three
species were added by Boise (1985) who reported that taxa
in the Pleomassariaceae and Massarinaceae shared similar
characters and might be closely related (Thambugala et al.
2018). Another lichenicolous genus Lichenopyrenis was
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Fungal Diversity (2020) 104:1–266
Fig. 18 Ophiosphaerella chiangraiensis (MFLU 11–0246, holotype). a Appearance of ascomata on host surface. b, c Sections
through ascomata. d, e Section through peridium. f Pseudoparaphyses
stained with Congo Red. g, h Asci. i Fissitunicate asci stained with
Congo Red. j–m Ascospores. Scale bars: a = 200 μm, b, c = 100 μm,
d = 50 μm, e–m = 20 μm
erected introduced based on perithecioid ascomata, a peridium comprising of compressed cells of textura angularis,
fissitunicate asci, wide hamathecium filaments, and 1-septate, distoseptate, pale orange-brown ascospores (Calatayud
et al. 2001). Currently, Pleomassariaceae consists of seven
genera, Beverwykella Tubaki, Lichenopyrenis Calat. et al.,
Myxocyclus Riess, Peridiothelia D. Hawksw., Prosthemium
Kunze, Pseudotrichia Kirschst. and Splanchnonema Corda
(Wijayawardene et al. 2014). The asexual morphs of Pleomassariaceae are mostly coelomycetous.
13
Prosthemium Kunze, in Kunze & Schmidt, Mykologische
Hefte (Leipzig) 1: 17 (1817)
Notes: The genus Prosthemium was introduced by Kunze
(1817) with Prosthemium betulinum as the type species. The
characters that define the genus are immersed, erumpent
Fungal Diversity (2020) 104:1–266
31
Fig. 19 Phylogram generated from maximum likelihood analysis
based on a combined ITS, LSU, SSU and TEF1-α sequence dataset
of representative genera in Phaeosphaeriaceae. Ophiobolus artemisiicola (MFLU 15-2140), O. disseminans (MFLUCC 17-1787) and
O. rossicus (MFLU 17-1639) were used as the outgroup taxa. Bootstrap support value for ML and MP ≥ 60% are given above the nodes.
Newly generated sequence is in blue. Type strains are in bold
ascomata, a dense, cellular, filiform hamathecium, broadly
cylindrical to broadly cylindro-clavate asci, muriform brown
to golden brown ascospores. The asexual morph is coelomycetous producing staurosporous brown macroconidia
as well as ellipsoid guttulate microconidia. Seven species
are recorded in Prosthemium. Species of Prosthemium can
be found worldwide especially in Europe. We illustrate the
sexual and asexual morph of P. betulinum as a new record
from Italy.
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32
Prosthemium betulinum Kunze, in Kunze & Schmidt,
Mykologische Hefte (Leipzig) 1: 18 (1817)
Facesoffungi number: FoF 06080; Figs. 20, 21
Saprobic in terrestrial habitats. Sexual morph: Ascomata
879–979 × 284–401 μm ( x = 922.9 × 339.2 μm, n = 10), solitary, scattered, or in small groups, immersed, erumpent or
depressed globose, medium to large, black, ostiolate. Ostiole
papillate, opening via a minute slit or a small conical swelling in the bark, ostiolar canal filled with a tissue of hyaline
small cells. Peridium 1–2 layered comprised of compressed
cells of textura angularis, apex comprises comparatively
larger cells than the base. Hamathecium of dense, cellular,
filiform, broad, septate pseudoparaphyses, embedded in a
gelatinous matrix. Asci 77–186 × 30–42 μm ( x = 140.1 × 36.4
μm, n = 10), 8-spored, bitunicate, fissitunicate, ellipsoid,
broadly cylindrical to broadly cylindro-clavate, with a short,
narrow, thick pedicel, rounded at the apex with a minute ocular chamber. Ascospores 52–73 × 14–16 μm ( x = 61.3 × 15.4
μm, n = 10), biseriate, partially overlapping, narrowly oblong
with broadly to narrowly rounded ends, brown to golden
brown, muriform, 5–8 transverse septa and 1–2 vertical septa
in some cells, constricted at each septum, smooth to verrucose wall with a mucilaginous sheath. Asexual morph: Pycnidia 390–359 × 301–316 μm ( x = 302.3 × 310.1 μm, n = 10),
globose to subglobose, brown to black. Conidiophores
10–14 × 4–6 μm ( x = 12.7 × 5.3 μm, n = 10), unbranched,
hyaline, septate. Macroconidia brown, staurosporous, 49–95
μm diam with two main arms and 2 or 3 smaller arms. Main
arms 22–33 × 14–19 μm ( x = 28.2 × 16.3 μm, n = 20), 4–5
transversely septate, with terminal cell hyaline to pale
brown. Smaller arms continuous or 1–3 septate, hyaline to
pale brown. Microconidia 4–5 × 2–3 μm ( x = 4.7 × 3.2 μm,
n = 10), brown to dark brown, sometimes with 1–2 guttules,
globose to ellipsoid, aseptate, thick-walled, smooth.
Culture characteristics: Colonies on MEA, 10–15 mm
diam after 7 days at 16 ºC, margin regular, slightly raised,
rough surface, entire edge, cottony or woolly, well-defined
edges with no pigmentation of the agar.
Material examined: ITALY, Via Togliatti—Forlì (Province of Forlì-Cesena [FC])—Italy, dead land branch of Betula sp. (Betulaceae), 26 March 2018, E. Camporesi (MFLU
17-0975); HKAS 97499.
GenBank numbers: LSU: MK652791; SSU: MK652791;
ITS: MK652789.
Notes: Isolate MFLUCC 17-2312 was recovered from
a dead branch of Betula sp. from Forli-Cesena Province.
The new isolate clusters with other Prosthemium betulinum
strains with moderate bootstrap support (65% ML/1.00 PP)
(Fig. 22). The ITS sequence similarity between the new
isolate and other P. betulinum strains is 100% (Jeewon and
Hyde 2016). Therefore, we introduce P. betulinum as a new
record from Italy.
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Fungal Diversity (2020) 104:1–266
Roussoellaceae J.K. Liu, R. Phookamsak, D.Q. Dai & K.D.
Hyde, Phytotaxa 181: 7 (2014)
Notes: Liu et al. (2014) erected the family to accommodate Roussoella Sacc., Neoroussoella J.K. Liu et al., and
Roussoellopsis I. Hino & Katum. Jayasiri et al. (2019) transferred Pararoussoella Wanas., E.B.G. Jones & K.D. Hyde
and Neoconiothyrium Crous to the family. The type genus
is Roussoella Sacc. This group of fungi is commonly found
on bamboo and palm hosts and is characterized by darkened,
lightly raised, somewhat linear or dome-shaped areas on the
host surface, semi-immersed or immersed ascostromata,
4–8-spored, and bitunicate asci with septate, brown to dark
brown ascospores.
Neoroussoella J.K. Liu, Phookamsak & K.D. Hyde, Phytotaxa 181: 21 (2014)
Notes: The genus Neoroussoella was established in 2014
with the type Neoroussoella bambusae Phook., Jian K. Liu
& K.D. Hyde based on an asexual morph producing hyaline
conidia with smooth walls. Two species N. bambusae and
N. lenispora Jin F. Zhang, Jian K. Liu, K.D. Hyde & Zi Y.
Liu were added by Liu et al. (2014) and Hyde et al. (2016)
respectively. Later, Jayasiri et al. (2019) added N. entade and
N. leucaenae and transferred Roussoella solani (CPC 26331)
to Neoroussoella. Karunarathna et al. (2019) added N. alishanense and Phookamsak et al. (2019) added N. heveae.
Neoroussoella magnoliae N.I. de Silva & K.D. Hyde, sp.
nov.
Index Fungorum number: IF556463; Facesoffungi number: FoF 06106; Fig. 23
Etymology: The specific epithet reflects the host Magnolia grandiflora.
Holotype: MFLU 18-1022.
Saprobic on dead twigs of Magnolia sp. Sexual
morph: Ascomata 200–300 µm high, 200–250 µm diam
( x = 245 × 230 µm, n = 10), semi-immersed on host surface, solitary, globose to subglobose, dark brown. Neck
small, elongate, located in central, covered by host tissues.
Peridium 15–20 µm wide, comprising brown textura prismatic cells. Hamathecium comprising 1–2 µm wide, cylindrical to filiform, septate, hyaline pseudoparaphyses. Asci
45–60 × 4–6 µm ( x = 56 × 5 µm, n = 25), 8 spored, bitunicate,
fissitunicate with a short pedicel. Ascospores 7–9 × 3–4 µm
( x = 8.2 × 3.2 µm, n = 30), overlapping, fusiform, 1-septate,
light brown and brown. Asexual morph: Undetermined.
Culture characteristics: Colonies on PDA reaching 25
mm diam after 1 week at 20–25 °C, colonies medium sparse,
circular, flat, surface slightly rough with edge entire, margin well-defined, cottony to fairly fluffy, colony from above:
greyish green; reverse light brown.
Material examined: CHINA, Yunnan Province, Xishuangbanna, dead twigs (attached to the tree) of Magnolia
Fungal Diversity (2020) 104:1–266
Fig. 20 Prosthemium betulinum. (MFLU 17-0975, a new geographical record). a,b Appearance of ascomata on host surface. c
Vertical section through the ascomata. d Peridium e–g Asci. h Asci
showing ocular chamber i Ascospore in Indian ink showing sheath
33
j-l Ascospores m Germinating ascospores n,o Culture characteristics on MEA (n: reverse view; o: above view) Scale bars: a = 1000
μm, b = 2000 μm, c = 400 μm, d = 40 μm, e–g = 100 μm, h–l = 20 μm,
m = 50 μm
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Fig. 21 Prosthemium betulinum. Asexual Morph (MFLU 17-0975, a new geographical record). a,b Pycnidia on culture. c–g Macroconidia.
h–m Microconidia. Scale bars: a = 1000 μm, b = 1000 μm, c = 15 μm, d–g = 20 μm, h–m = 5 μm
grandiflora (Magnoliaceae), 26 April 2017, N. I. de Silva,
NI158 (MFLU 18-1022, holotype), living culture, MFLUCC
18-0721, KUMCC 17-0190.
GenBank numbers: ITS: MK801232; LSU: MK801230;
SSU: MK801231; TEF1: MK834373.
Notes: A sexual morph of Neoroussoella was collected in
this study and phylogenetic analysis confirmed its placement
in Roussoellaceae. The asexual morph of the new species
was not observed from the culture. In our multigene phylogenetic study, N. magnoliae (MFLUCC 18-0721) clustered
with other species of Neoroussoella (Fig. 24). The new species expands the host families inhabited by Neoroussoella
with the addition of the woody plant family Magnoliaceae.
N. magnoliae differs from other Neoroussoella species
in having the shortest asci (45–60) µm and the shortest
ascospores (7–9) µm except for N. leucaenae. N. leucaenae
is morphologically similar to N. magnoliae, but phylogenetically distantly related. N. leucaenae was collected from
a decaying pod of Leucaena sp. (Fabaceae) in Thailand.
Morphological differences in ascomata, asci, ascospores,
13
condiomata and conidia among Neoroussoella species are
given in Table 2.
Sporormiaceae Munk, Dansk Botanisk Arkiv 17 (1): 450
(1957)
Notes: Sporormiaceae (Pleosporales, Dothideomycetes)
was erected by Munk (1957) and includes six genera according to the Outline of Ascomycota (Wijayawardene et al.
2018). Although most of the taxa are coprophilous, a small
number in this family are endophytes (Arenal et al. 2007;
Crous et al. 2018). The sexual morphology is characterized
by ascomata solitary or gregarious, perithecioid or cleistothecioid, immersed, erumpent or superficial, dark pigment,
globose to pyriform; peridium smooth or hairy, fairly thick,
with dark pigment; asci usually 8–spored, fissitunicate, nonamyloid, clavate, globose or cylindrical; ascospores often
partly overlapping inside the asci, uni-to triseriate, sometimes fasciculate or crowded, sometimes but rarely onecelled, usually septate and poly-celled, oval to cylindrical,
thick-walled, smooth, sometimes but rarely ornamented,
Fungal Diversity (2020) 104:1–266
35
Fig. 22 Phylogram generated from maximum likelihood analysis
based on a combined ITS, LSU and SSU sequence dataset retrieved
from the GenBank. Melanomma pulvis-pyrius and M. japonicum are used as the out-group taxa. ML bootstrap values ≥ 50% are
given as the first set of numbers and approximate likelihood-ratio
test (aLRT) ≥ 0.90 values as the second set of numbers above the
nodes. Voucher/strain numbers are given after the taxon names, the
ones from type material are indicated in bold face. Newly generated
sequence is indicated in blue. The bar length indicates the number of
nucleotide substitutions per site
dark brown. The asexual morph is rare, when present it
is coelomycetous, with Phoma-like pycnidia (Hyde et al.
2013).
Sporormurispora Wanas., Bulgakov, Gafforov & K.D.
Hyde, in Wanasinghe et al., Fungal Diversity: 10.1007/
s13225-018-0395-7, [157] (2018)
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Fungal Diversity (2020) 104:1–266
Fig. 23 Neoroussoella magnoliae (MFLU 18-1022, holotype). a–c Appearance of ascomata on host surface. d, e Vertical sections through
ascomata. f Peridium. g Pseudoparaphyses. h–j Asci. k–n Ascospores. Scale bars: d, e = 50 μm, f = 10 μm, h–j = 20 μm, k–n = 5 μm
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Fungal Diversity (2020) 104:1–266
Fig. 24 Phylogram generated from maximum likelihood analysis
based on a combined LSU, SSU, ITS, TEF1 and RPB2 sequence
dataset. Related sequences were obtained from GenBank. Lophiostoma macrostomum (HHUF 2793) was used as the out-group taxon.
37
ML bootstrap support (first set) ≥ 50% and Bayesian posterior probabilities (second set) ≥ 0.90 are given near to each branch. The ex-type
strains are in black bold. Newly generated sequence is in blue
13
38
13
Table 2 Synopsis of Neoroussoella species
Species
Ascomata (μm)
Asci (μm)
Ascospores (μm)
Neoroussoe- 230–310 high, 370–480 diam,
immersed in host epidermis,
lla alishanraised, hemispherical to subconiense
cal, coriaceous
60–80 × 4.5–6
N. bambusae Ascostromata 120–175 high,
380–450 diam, immersed under
a clypeus, or epidermis, raised,
visible as black dome-shaped or
shield-shaped
N. entadae
No sexual morph observed, only
asexual morph present
N. leucaenae 215–275 high, 175–225 diam, semiimmersed, globose to subglobose,
dark brown to black
60–80(–82)
(–85) × 5–6(–
6.5)
Decaying stems of Pennisetum pur7.5–11 × 3–4, pale brown to brown
pureum Schumach. (Poaceae)
or yellowish brown, ellipsoid to
fusiform, 2-celled, constricted at
the septum, rough-walled, longitudinally ribbed
On dead branch of Bambusa
(7.5–)8–10 × 3–4, 2-celled, con(Poaceae)
stricted at the septum, pale brown,
surrounded by a mucilaginous
sheath
–
–
Reference
Taiwan
Karunarathna et al. (2019)
Thailand
Liu et al. (2014)
On stems of Solanum mauritianum
(Solanaceae)
Dead twig of Magnolia grandiflora
(Magnoliaceae)
Jayasiri et al. (2019)
Jayasiri et al. (2019)
Guizhou, China Hyde et al. (2016)
France
Crous et al. (2016)
Yunnan, fChina This study
Fungal Diversity (2020) 104:1–266
7–9 × 3–4, fusiform, 1-septate, light
brown and brown
Locality
On decaying pod of Entada phaseo- Thailand
loides (Fabaceae)
On decaying pod of Leucaena sp.
Thailand
(Fabaceae)
50–60 × 4.5–5.5 8–9 × 2.5–3.5, hyaline to brown,
fusiform with narrow, acute ends,
two asymmetric cells with guttules, constricted at the septum
74–127 × 5.9–9 10–16 × 3.5–5, ellipsoid to fusiform, On decaying branch of
2-celled, constricted at the septum, unidentified host
pale brown to brown, guttulate
when young
Ascostromata semi-immersed to
erumpent, clypeate, globose to
subglobose, solitary to gregarious,
black, coriaceous, centrally ostiolate, visible as black, dome-shaped
spots
N. solani
No sexual morph observed, only
–
asexual morph present
N. magnoliae 200–300 high, 200–250 diam, semi- 45–60 × 4–6
immersed on host surface, solitary,
globose to subglobose, dark brown
N. lenispora
–
Host
Fungal Diversity (2020) 104:1–266
39
Fig. 25 Sporormurispora paulsenii (TASM 6149, holotype). a,b
Ascomata on host substrate. c Section of ascoma. d Peridium. e Pseudoparaphyses. f–h Asci. i–l Ascospores. m Germinating ascospores.
n,o Culture characteristics on MEA (n: Above view; o: Reverse
view). Scale bars: a = 1000 μm, b = 1000 μm, c = 100 μm, d = 20 μm,
e = 10 μm, f = 50 μm, g,h = 100 μm, i–l = 10 μm, m = 20 μm
Notes: The genus Sporormurispora was erected by
Wanasinghe et al. (2018) with S. atraphaxidis as the type
species. The latter was isolated from stems of Atraphaxis
replicata Lam in Russia. The characters that define the
genus are black, globose, uniloculate ascomata with an ostiole, a thick peridium, comprising 6–8 layers, filamentous,
branched, septate, pseudoparaphyses, cylindrical asci and
uniseriate, mostly ellipsoid, brown, muriform ascospores.
We introduce a new species of Sporormurispora based on
morphology and phylogeny.
Sporormurispora paulsenii D.Pem, Gafforov & K.D. Hyde,
sp. nov.
Index Fungorum number: IF556258; Facesoffungi number: FoF 06252; Fig. 25
Etymology: The epithet reflects the name of the host plant
‘paulsenii’.
13
40
Holotype: TASM 6149.
Saprobic in terrestrial habitats. Sexual morph: Ascomata 673–757 μm × 470–589 μm ( x = 715.5 × 529.6 μm,
n = 10), black, immersed to erumpent, solitary or in groups,
gregarious or confluent, disc-shaped, uniloculate, without
an ostiole. Peridium 43–52 μm thick, comprising 2 layers,
outer layer comprising heavily pigmented, thick-walled,
dark brown amorphous layer, middle layer comprising
thick-walled, blackish to dark brown loosely packed cells
of textura angularis, inner layer composed hyaline, flattened, thick-walled cells of textura angularis. Hamathecium
1.5–2.0 μm (n = 20), comprising numerous, filamentous,
branched, septate pseudoparaphyses. Asci 129–283 × 18–22
μm ( x = 180.4 × 18.9 μm, n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, short-pedicellate, apex rounded with
a minute ocular chamber. Ascospores 32–35 × 16–19 μm
( x = 33.9 × 18.2 μm, n = 10), uniseriate to overlapping uniseriate, muriform, mostly ellipsoid, 7–10-transversely septate,
with 2–3-longitudinal septa, slightly constricted at the middle septum, initially hyaline, becoming dark brown at maturity, rounded at the ends. Asexual morph: Undetermined.
Culture characteristics: Colonies on MEA, 10–15 mm
diam after 7 days at 16 ºC, margin regular, slightly raised,
smooth surface, entire edge, cottony or woolly, whitish,
well-defined edges with no pigmentation of the agar.
Material examined: UZBEKISTAN, Surxondaryo,
Province, Boysun district, Omonxona village, South-Western Hissar Mountains, on branches of Colutea paulsenii
(Fabaceae), 18 May 2016, Y. Gafforov, YG-S121-1 (TASM
6149, holotype); MFLU 17-0112, isotype, ex-type living
culture, MFLUCC 17-1957.
GenBank numbers: LSU: MK966143; SSU: MK963075;
RPB2: MN023029.
Notes: Sporormurispora paulsenii was collected from
Colutea paulsenii in Uzbekistan, and clusters with S.
pruni (MFLUCC 17-0803) and S. atraphaxidis (MFLUCC
17-0742) with strong bootstrap support (100% ML and
1.00 PP) (Fig. 26). These taxa are similar in having a thick
peridium, pseudoparaphyses, cylindrical asci and uniseriate,
mostly ellipsoid, brown, muriform ascospores. However, S.
paulsenii differs from S. atraphaxidis, the type species of
Sporormurispora, in having disc- shaped ascomata, a thicker
peridium (43–52 μm vs 15–25 μm), narrower hamathecium
(1.5–2.0 μm vs 1.5–3 μm) and lacks an ostiole. Therefore,
we introduce S. paulsenii as a new species.
Tubeufiales Boonmee & K.D. Hyde
Notes: The order Tubeufiales was established by Boonmee
et al. (2014) to accommodate the type family Tubeufiaceae
based on phylogenetic studies. Liu et al. (2017a) accepted
Bezerromycetaceae and Wiesneriomycetaceae in the Tubeufiales based on morphological and phylogenetic analyses.
However, Lu et al. (2018d) revised this order and confirmed
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Fungal Diversity (2020) 104:1–266
that only Tubeufiaceae was included in the Tubeufiales based
on the phylogenetic analyses of combined ITS, LSU, RPB2
and TEF1-α.
Tubeufiaceae M.E. Barr, Mycologia 71(5): 948 (1979)
Notes: Tubeufiaceae was established by Barr (1979) with
Tubeufia as the type genus. Most species of Tubeufiaceae are
widespread in tropical and temperate regions, and can live
terrestrially as saprobes on woody substrates or in aquatic
habitats (Barr 1979, 1980; Rossman 1987; Kirk et al. 2001;
Ho et al. 2002; Cai et al. 2003; Zhao et al. 2007; Boonmee et al. 2011, 2014; Hyde et al. 2016; Brahamanage et al.
2017; Chaiwan et al. 2017; Doilom et al. 2017; Lu et al.
2017a, b, c, d; Luo et al. 2017; Liu et al. 2018; Phookamsak
et al. 2018). Lu et al. (2018d) documented this family and
provided an updated phylogenetic tree of combined ITS,
LSU, RPB2 and TEF1-α sequences for the Tubeufiales. They
established 13 genera in Tubeufiaceae, and confirmed that
the Tubeufiales (including the Tubeufiaceae), comprises 42
genera based on their phylogeny-morphology analyses. We
follow the treatments of Tubeufiaceae and Tubeufiales by
Lu et al. (2018d) and present a new species, Helicoarctatus
thailandicus, based on morphological characteristics and
phylogenetic analyses.
Helicoarctatus Y.Z. Lu, J.C. Kang & K.D. Hyde, in Lu
et al., Fungal Diversity 92: 166 (2018)
Notes: Helicoarctatus was introduced by Lu et al. (2018d)
as a monotypic genus with H. aquaticus as the type species.
Helicoarctatus is characterized by setiform, unbranched,
septate conidiophores, holoblastic, mono- to poly-blastic,
denticulate conidiogenous cells and pleurogenous, helicoid,
rounded at tip, multi-septate, tightly coiled 3–4 times, guttulate, hyaline conidia. The sexual morph is unknown (Lu
et al. 2018d). We introduce a new species, Helicoarctatus
thailandicus, based on both morphology and phylogeny.
Helicoarctatus thailandicus D.F. Bao, Z.L. Luo, K.D. Hyde
& H.Y. Su, sp. nov.
Index Fungorum number: IF556259; Facesoffungi number: FoF 05831; Fig. 27
Etymology: Referring to the fungus being collected from
Thailand.
Holotype: MFLU 19-0555.
Saprobic on submerged decaying wood in a freshwater
stream. Sexual morph: Undetermined. Asexual morph:
Colonies on the substratum superficial, effuse, gregarious,
velvety, white to pale brown, shiny. Mycelium composed of
partly immersed, partly superficial, brown to dark brown,
septate, branched hyphae. Conidiophores 302–536 µm long,
6–11 μm wide ( x = 415.3 × 9.4 µm, n = 30), macronematous,
mononematous, erect, cylindrical, straight or slightly flexuous, unbranched, tapering towards apex, with red to purple
Fungal Diversity (2020) 104:1–266
41
Fig. 26 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU and RPB2 sequence data retrieved
from the GenBank. ML bootstrap values ≥ 50% are given as the first
set of numbers and approximate likelihood-ratio test (aLRT) ≥ 0.90
values as the second set of numbers above the nodes. Voucher/strain
numbers are given after the taxon names, the ones from type material
are indicated in bold face. Newly generated sequence is indicated in
blue. The bar length indicates the number of nucleotide substitutions
per site
appendage near apex, septate, brown to reddish brown,
smooth-walled. Conidiogenous cells 3–6.9 μm long, 2.5–4.5
μm wide, holoblastic, monoblastic, determinate, denticulate,
arising laterally from lower portion of the conidiophores
as tiny tooth-like protrusions, with each bearing 1 tiny
sporogenous conidiogenous loci, hyaline, smooth-walled.
Conidia 44–62 µm diam, 417–524 µm long, 3.7–6.8 μm
wide ( x = 460 × 51 µm, n = 30), solitary, helicoid, rounded at
both of ends, muriform, with small guttule, aseptate, tightly
13
42
Fig. 27 Helicoarctatus thailandicus (MFLU 19-0555, holotype).
a–c Colony on decaying wood. d–f Conidiophores with attached
conidia. g–h Conidiogenous cells with conidia. i–j Conidia. k Germi-
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nating conidium. l, m Colony on MEA from above and below. Scale
bars: d–f = 50 μm, g–k = 30 μm
Fungal Diversity (2020) 104:1–266
coiled 4 times, becoming loosely coiled in water, hyaline,
smooth-walled.
Culture characteristics: Conidia germinating on PDA
within 24 h. Colonies on MEA at room temperature reaching 2.3 cm diam in 3 weeks, mycelium pale brown to greyish
brown after 3 weeks, composed of brown to dark brown,
septate, smooth or verrucose hyphae.
Material examined: THAILAND, Khwaeng Bang Chak,
Khet Phra Khanong Krung Thep Maha Nakhon, on submerged decaying wood, October 2017, Z.L. Luo, B site
3–2–1. (MFLU 19-0555, holotype), ex-type culture,
MFLUCC 18-0332.
GenBank numbers: LSU: MK559869; SSU: MK559780;
TEF1-α: MK541685.
Notes: In our phylogenetic analyses, Helicoarctatus
thailandicus clustered with H. aquaticus but was separate
in a distinct lineage with full support (100 ML/1.00 PP)
(Fig. 28). Morphologically, H. thailandicus resembles H.
aquaticus in having unbranched, septate, and wide at bottom, tapering at apex conidiophores, holoblastic, determinate, discrete, denticulate, hyaline conidiogenous cells and
helicoid, hyaline conidia. However, H. thailandicus differs
from H. aquaticus in its conidiophores with a red to purple
appendage near the apex and aseptate conidia, rounded at
both of ends, and becoming loosely coiled in water. The
conidiophores of H. aquaticus are without an appendant and
the conidia are not rounded at the tip and do not become
loosely coiled in water.
Thaxteriellopsis Sivan., Panwar & S.J. Kaur, Kavaka 4: 39
(1977) [1976]
Notes: Thaxteriellopsis Sivan., Panwar & S.J. Kaur was
introduced as a new genus by Sivanesan et al. (1976) based
on the type species T. lignicola Sivan., Panwar & S.J. Kaur.
Two more species were added to this genus but were transferred to Chaetosphaerulina (Crane et al. 1998; Wijayawardene et al. 2017). However, the type species T. liginicola has been retained in Thaxteriellopsis by Boonmee et al.
(2011) and Lu et al. (2018d). Crane et al. (1998) suggested
that multiseptate ascospores, often with one or few vertical
septa, could be placed in Chaetosphaerulina, but they did
not mention this in the generic description (Boonmee et al.
2011). After re-examination of the holotype (IMI197065),
Boonmee et al. (2011) designated an epitype (MFLU
10-0057) and described both the sexual morph, found on a
dead piece of wood of Zizyphus mauritiana, and the asexual
morph produced in MEA culture. Thaxteriellopsis is characterized by non-ostiolate, setose, superficial, broadly sphaerical ascomata on a subiculum; bitunicate, cylindric-clavate,
short pedicellate asci and hyaline, fusiform, phragmoseptate
ascospores. It also produces a Moorella-like asexual morph
(Subramanian and Sekar 1982). At present this genus is
monotypic.
43
Thaxteriellopsis obliqus M. Niranjan and V.V. Sarma, sp.
nov.
Index Fungorum number: IF 557437; Facesoffungi number: FoF 06266; Figs. 29, 30
Etymology: With reference to an oblique septum in the
ascospores.
Holotype: AMH-10087.
Saprobic on decaying Nauclea gageana. Sexual morph:
Ascomata 140–190 × 240–345 μm, superficial, with black
subiculum, oblate to spheroid, dark brown, scattered, ostiolate, periphysate, velvety, hyphal appendages laterally surrounding ascoma, sparse, flexuous, thick-walled, brown,
septate. Necks 80 × 100 μm, depressed when dry, narrow
towards apex. Peridium up to 25 μm wide, composed of several layers of outer brown and inner hyaline cells of textura
angularis, with guttules. Hamathecium composed of 1–2
μm wide, filamentous, septate, unbranched, anastomosing
trabeculate pseudoparaphyses. Asci 67–78(–80) × 15.5–19
μm ( x = 74.5 × 17.5 μm, n = 25), 8-spored, bitunicate,
cylindric-clavate, apically rounded with an ocular chamber, narrowing towards base, short pedicellate. Ascospores
22–29 × 4.0–5.5 μm ( x = 24.8 × 4.8 μm, n = 25), hyaline, triseriate, obclavate-cylindrical, gently curved, obtuse at both
ends, apex broader than base, 6–8(–9) trans-septa, rarely
muriform with one longitudinal septum which is oblique, not
constricted at septa, mature spores becoming grey, smoothwalled. Asexual morph: Hyphomycetous, helicosporous,
Conidiophores erect, straight or somewhat bent, macronematous, mononematous, superficial, cylindrical, arising as lateral branches from hyphae, branching into whorls, septate,
slightly constricted at septa, 170–200 × 6.0–8.5 μm, dark
brown to brown, narrowing towards apex, smooth-walled.
Each conidiophore bearing 2 to 5 branches of verticels (in
whorls) along stipe at intervals (30–45 μm), each whorl having 3 to 5 brown branches. Each branch is 1–4 celled, brown
to dark brown, 3–7 μm. Terminal cell of these branches is
conidiogenous. Conidiogenous cells holoblastic, mono- to
polyblastic, integrated, sympodial, terminal, cylindrical,
3.5–6.5 × 2–2.5 μm, pale brown, denticulate, smooth-walled.
Conidia hyaline, solitary, acropleurogenous, helicoid,
12–15 × 9–11.5 μm ( x = 13.3 × 9.8 μm, n = 25), conidial
filament, coiled 1 to 1½ times, rounded, obtuse end, 3–5
septate, without constrictions, guttulate, smooth-walled,
arising on small denticles.
Cultural characteristics: Ascospores germinating on
water agar within 24 h and two germ tubes produced at end
cells. Colonies growing on MEA, cottony, sterile mycelium, initially grey-brown, become dark brown at maturity,
reaching 18 mm in 1 week at 28 °C, mycelium superficial,
raised at center, radially straight to undulated edges, septate,
unbranched, brown coloured hyphae.
Material examined: INDIA, Andaman and Nicobar
Islands, South Andaman, Chidiyatapu Reserve Forest
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Fungal Diversity (2020) 104:1–266
Fig. 28 Phylogenetic tree generated from maximum likelihood analysis based on a combined LSU, ITS and TEF1-α sequence dataset
for some selected species within the Tubeufiaceae. Botryosphaeria
dothides CBS 1154.76 and B. agaves MFLUCC 10-10051 are used
as the out-group taxa. Bootstrap support values for maximum likelihood (ML) ≥ 75% and Bayesian posterior probabilities (PP) ≥ 0.95 are
given above the nodes. Newly generated sequence is indicated in blue
and ex-type strains are in bold
(11°29′52.8" N, 92°42′38.6" E). Recorded on Nauclea gageana, 20 May 2018, M. Niranjan & V.V. Sarma (PUFNI
18,727). Herbarium submitted in Ajrekar Mycological Herbarium-AMH (AMH-10087) and NFCC-4429 (holotype)
deposited at National Fungal Culture Collection of India
(NFCCI), Pune.
GenBank numbers: ITS: MK981534; LSU: MK981533;
TEF1: MK984839.
Notes: Thaxteriellopsis obliqus produces ascospores with
one or few vertical/oblique septa similar to Chaetosphaerulina spp. (Crane et al. 1998; Lu et al. 2018d). However,
most of the Chaetosphaerulina members have multiseptate ascospores that are more than 10 septate excepting
C. eucalypti, which has 6–10 septate ascospores with few
vertical or oblique septa. T. obliqus, however, differs from
C. eucalypti by having prominent necks with periphyses,
13
Fungal Diversity (2020) 104:1–266
45
Fig. 29 Thaxteriellopsis obliqus (NFCC-4429, holotype). a Ascomata on host. b Ascomata attached by lateral hyphae. c Section through
ascoma. d Peridium. e–g Asci. g–p Ascospores. Scale bars: d = 200 µm, c = 100 µm, d = 50 µm, e–g = 20 µm, g–p = 10 µm
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Fungal Diversity (2020) 104:1–266
Fig. 30 Thaxteriellopsis obliqus (NFCC-4429, holotype). a Conidiomata on host. b Culture on MEA. c, d Conidiophores. e Conidiogenous
cells. f–i Conidia. Scale bars: c, d = 50 µm, e = 20 µm, f–i = 10 µm
narrower and smaller ascomata, smaller asci and smaller
ascospores that lack constrictions at the septa. Molecular
sequence data of Chaetosphaerulina spp. are not available in
the public domain to be included in our phylogenetic analysis for a comparison. The phylogenetic tree also shows that
13
T. obliqus branches separately from T. lignicola with good
bootstrap support of 100% in ML, 100% in MP and 1.00 in
PP (Fig. 31). T. lignicola is known to produce a Moorellalike anamorph (Subramanian and Sekar 1982). A similar
anamorph is found in T. obliqus also. T. obliqus produces
Fungal Diversity (2020) 104:1–266
ascospores that are 6–8(–9) septate without constrictions
at the septa unlike T. lignicola which produces 5-septate
ascospores that have constrictions at the septa. Furthermore,
T. obliqus differs from T. lignicola in having a central, ostiolated neck in the ascomata, and narrower ascospores with a
vertical or oblique septum. In contrast, T. lignciola produces
non-ostiolate ascomata. Hence a new species, T. obliqus, is
added to the genus.
Incertae sedis in Dothideomycetes
Notes: Species of Scleroramularia Batzer & Crous formed
a sister branch of Pleosporales in the phylogenetic tree, but
its placement was not given (Li et al. 2011).
Scleroramularia Batzer & Crous, Fungal Diversity 46: 58
(2011)
Notes: The genus was established with Scleroramularia
pomigena Batzer & Crous in 2011 as the type species and
characterized as forming black sclerotial bodies in culture
and producing chains of hyaline conidia which do not disarticulate readily (Li et al. 2011). Species of Scleroramularia
are ectophytic parasites causing sooty blotch and flyspeck
on the surface of fruits including apple, pawpaw and Japanese banana (Gleason et al. 2019). Six species have been
reported: S. abundans Batzer, Mayfield & Crous, S. asiminae Batzer, Hemnani & Crous, S. henaniensis G.Y. Sun,
H.Y. Li & Crous, S. pomigena Batzer & Crous, S. shaanxiensis G.Y. Sun, H.Y. Li & Crous and S. musae G.Y. Sun &
L. Gao (Li et al. 2011; Gao et al. 2015).
Scleroramularia vermispora G.Y. Sun & L. Gao, sp. nov.
Index Fungorum number: IF556172; Facesoffungi number: FoF 05788; Fig. 32
Etymology: Named after its conidia shape.
Holotype: HMUABO GLZJXJ6.
Ectophytic on the surface of Japanese banana (Musa basjoo) fruit. Sexual morph: Undetermined. Asexual morph
on SNA. Mycelium creeping, superficial and submerged,
consisting of hyaline, smooth, branched, septate, 1.5–2.5
μm diam hyphae. Conidiophores mostly reduced to conidiogenous cells, or with one supporting cell. Conidiogenous
cells solitary, erect, intercalary on hyphae, subcylindrical,
straight, with 1 terminal locus, 2–7 × 1.5–3 μm. Conidia
in branched chains, hyaline, smooth, straight, curved, and
usually bending into a “C” shape or vermiform if long and
thin; basal conidia mostly narrowly cylindrical, 0–3-septate, 23–57 × 1.5–2.5 μm; intercalary and terminal conidia
becoming more narrowly ellipsoid to vermiform, 0–4-septate, 11–34(–43) × 1.5–3 μm.
Culture characteristics: Colonies after 2 weeks at 25
°C on SNA spreading with sparse aerial mycelium and
47
somewhat feathery margin, reaching 15 mm diam, with
white surface.
Material examined: CHINA, Guangdong Province, Zhanjiang, on the surface of Musa basjoo fruit, October 2011,
L. Gao, W.H. Li & H.C. Chen, GLZJXJ6 (GLZJXJ6, holotype). Other specimens examined: CHINA, Guangdong:
Zhanjiang, on the surface of M. basjoo fruit, October 2011,
L. Gao, W.H. Li & H.C. Chen, GL-ZJXJ-2D, GL-ZJXJ-4B,
GL-ZJXJ-10, GL-ZJXJ-20, GL-ZJXJ-26B, GL-ZJXJ-30B,
GL-ZJXJ-34; Guangxi Province, Beihai, on the surface of
M. basjoo fruit, October 2013, L. Gao & W.Y. Zhao, GLBHBJ-6, GL-BHFJ-13, GL-BHFJ.B-18A; Guangxi, Nanning, on the surface of M. basjoo fruit, October 2013, L. Gao
& W.Y. Zhao, GL-NNFJ-10, GL-NNFJ-18, GL-NNFJ-22A,
GL-NNFJ-22B, GL-NNJJ-23A, GL-NNFJ-26B; Yunnan, on
the surface of M. basjoo fruit, October 2011, L. Gao, W.H.
Li & H.C. Chen, GL-YNFJ-3B, GL-YNFJ-5, GL-YNFJ-6,
GL-YNFJ-10A, GL-YNFJ-10B, GL-YNFJ-10C, GL-YNFJ11A, GL-YNFJ-11B.
GenBank numbers: ITS: MK454748; TEF1: MK454977.
Notes: The phylogenetic tree based on a combined ITS
and TEF1 sequence dataset shows that Scleroramularia
vermispora branches separately from the other species in
the genus with full support (Fig. 33). The conidia of S. vermispora are long and thin, usually bending into a “C” shape
or vermiform.
Lecanoromycetes
Graphidales Bessey
Notes: Nannfeldt (1932) established the order Ostropales
Nannf. to accommodate the Ostropaceae Rehm (a synonym
of Stictidaceae Fr.), a family which is comprised of nonlichenized fungi. Gilenstam (1969) suggested the lichenized
genera in the Graphidales should also be included in Ostropales sensu stricto, and this wide definition was commonly
accepted (Eriksson and Hawksworth 1993; Hawksworth
et al. 1995; Winka et al. 1998). Graphidales, proposed by
Bessey (1907), therefore, had been included in the Ostropales (Lecanoromycetes) for a long time (Staiger 2002;
Kalb et al. 2004; Hibbett et al. 2007; Lumbsch et al. 2007;
Kirk et al. 2008; Baloch et al. 2010; Rivas Plata and Lumbsch 2011; Rivas Plata et al. 2012; Lumbsch et al. 2014a,b;
Lücking et al. 2017a,b; Wijayawardene et al. 2018). However, Kraichak et al. (2018) ranked Graphidales as a separate order based on a temporal approach, and accepted five
families. In the recent outline of fungi and fungus-like taxa
(Wijayawardene et al. 2020), six families are now listed in
the Graphidales.
Graphidaceae Dumort., [as ’Graphineae’], Comment. Bot.
(Tournay): 69 (1822)
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Fungal Diversity (2020) 104:1–266
Fig. 31 Maximum parsimony tree constructed by ITS, LSU and TEF1 sequences of species in Tubeufiaceae along with Thaxteriellopsis obliqus
(NFCC-4429). Botryosphaeria dothidea CBS 1154.76 served as the out-group. Bootstrap support values for maximumlikelihood (ML), maximum parsimony (MP) and Bayesian posterior probabilities (PP) values ≥ 65% are given above the nodes. Highlighted letter ‘a’ represents the country Thailand
13
Fungal Diversity (2020) 104:1–266
Notes: Graphidaceae is the largest crustose lichenized
family of approximately 2160 species predominatly tropical, encompassing a remarkable range of morphological and
chemical diversity (Kalb et al. 2018). The family members
are easily recognized as they are characterized by crustose
thalli with elongate and often carbonized ascomata, containing parallel paraphyses, and ascospores having usually
lenticular cell lumina, which turn violet or reddish with
Lugol’s solution (Staiger et al. 2006). The work regarding
investigation of lichens in Thailand was reviewed in Kalb
et al. (2018), and a revised checklist of lichenized fungi in
Thailand, provided by Buaruang et al. (2017), contains 290
species records in 46 genera of Graphidaceae.
Graphis Adans., Familles des Plantes 2: 11 (1763)
Notes: Graphis Adans. is the largest genera in Graphidaceae, with 390 species according to Lücking et al.
(2017a). Members of Graphis usually occurr in open, light
habitats, such as roadsides or at the edge of forests (Kalb
et al. 2018). A detailed study of Graphis in Thailand was
provided in Kalb et al. (2018), and refers to a broad sense
of Graphis in Staiger (2002). According to Lücking (2009)
Graphis is characterized by having a crustose thallus with
the alga Trentepohlia as a photobiont, lirellate ascomata
with uncarbonized, partly or completely carbonized exciple, an inspersed or non-inspersed hymenium and ± hyaline, transseptate or muriform I + blue ascospores. Phylogenetically, Graphis sensu represents two distinct, distantly
related clades in Rivas Plata et al. (2011), of which Graphis
sensu stricto comprises the type species G. scripta and the
name Allographa Chevall. is proposed for the second clade.
Recently, Lücking and Kalb (2018) have instated Allographa
as a genus separate from Graphis. Allographa usually has
prominent, rather massive, mostly completely carbonized
and often striate lirellae, sometimes with pigmented pruina
and generally with larger ascospores with more numerous
septa. However, Graphis contains species with secondary
thallus compounds, and with a stronger tendency to develop
exposed discs.
Graphis supracola A.W. Archer, Aust. Syst. Bot. 14(2): 267
(2001)
Index Fungorum number: IF483622; Facesoffungi number: FoF 06021; Fig. 34
Thallus corticolous, whitish or greyish white, smooth or
slightly roughened, ecorticate, dull. Lirellae erumpent to
prominent, with lateral thalline margin, no striation, short
and stellately or irregularly branched, up to 4 mm long, libia
convergent, white pruinose, disc concealed in dry material, nearly exposed of section in water. Excipulum lateral
carbonized, black, brown in thin section. Hymenium clear.
Asci 96–112 × 21–31 µm, 8-spored, clavate. Ascospores
25–50 × 6–9 µm ( x = 35.1 × 7.6 µm, n = 34), mostly 30–40
49
µm long, overlapping or tri-seriate, hyaline, clavate to ellipsoid with rounded ends, transversely 7–11-septate, with
lens-shaped lumina and gelatinous sheath, I + yellow to blue
or violet.
Chemistry: KOH–; Thallus P + orange (Fig. 34d), protocetraric acid.
Material examined: THAILAND, Chiang Rai, on decayed
wood in a mountain roadside forest, 20 June 2018, Y.S. Liu,
L02 (MFLU 19-0013).
GenBank numbers: mtSSU: MK770352; nuLSU:
MK770351.
Notes: Graphis has 390 species and only a few have been
sequenced. According to a world-wide key to the genus in
Lücking et al. (2009) and a key for recorded Graphis species
in Thailand (Kalb et al. 2018), the new collection resembles
G. supracola (Archer 2001) as both contain protocetraric
acid (P + orange, K–). G. supracola has been reported from
Africa (Joshi et al. 2016), Australia (Archer 2001), Cambodia (Moon et al. 2013), Thailand (Poengsungnoen et al.
2010; Schumm and Aptroot 2012), USA (Seavey and Seavey
2011) and Vietnam (Joshi et al. 2013). It is very common in
Thailand but there is no available sequence data (Kalb et al.
2018). The morphological characteristics of our collection
fit those of G. supracola (Kalb et al. 2018). The new collection has larger ascospores than previously described for G.
supracola (25–40 × 6–8 μm), which is well-characterized in
having concealed disc (caesiella-morph). Herein, we provide
reference sequences for G. supracola, and a phylogenetic
analysis based on mtSSU and nuLSU sequence data of Graphis species (Fig. 35). G. supracola and G. caesiella are
closely related in the phylogenetic tree, but they form distinct lineages with high statistical support and differ mostly
in the chemistry.
Leotiomycetes
Erysiphales Gwynne-Vaughan & Barnes
Notes: The order was erected by Gwynne-Vaughan and
Barnes (1927) with Erysiphaceae Tul. & C. Tul. as the type
family and the sole member of the order. All members of
Erysiphales are exclusively obligate biotrophic ascomycetes
with superficial mycelia and ascomata lacking ostioles and
enclosing single to numerous asci.
Erysiphaceae Tul. & C. Tul., Select. Fung. Carpol. 1: 191
(1861)
Notes: The establishment of Erysiphaceae dates back
to 1861 by the brothers Tulasne. This family consists of
ca. 900 species among 18 genera (Braun and Cook 2012;
Marmolejo et al. 2018). As one of the most important plant
pathogenic groups, species of Erysiphaceae can infect ca.
10,000 species of angiosperms including many economically
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Fungal Diversity (2020) 104:1–266
Fig. 32 Scleroramularia vermispora (HMUABO GLZJXJ6, holotype). a Flyspeck signs on banana peel. b Colony on synthetic nutrient-poor agar (SNA) after 2 weeks. c vermiform conidia. d Chains of
conidia. e “C” type conidia. f Chains of conidia with branch. Scale
bars: c–f = 20 µm
important, cultivated species such as cereals, vegetables,
fruit trees, and other flowering plants (Takamatsu 2018).
genus Podosphaera which subsequently was divided into
two non-monophyletic groups, sect. Podosphaera, sect.
Sphaerotheca. Section Sphaerotheca was further divided
into two subsections, subsect. Sphaerotheca and subsect.
Magnicellulatae. In section Podosphaera 86% of hosts
and in subsection Sphaerotheca 57% of hosts belong to the
Rosaceae and the close evolutionary relationship between
Podosphaera and its rosaceous hosts has been revealed by
Takamatsu et al. (2010).
Podosphaera Kunze, Mykol. (Hefte) 2: 111 (1823)
Notes: Many molecular analyses have confirmed that the
genera Podosphaera and Sphaerotheca form a clearly single
monophyletic clade (Saenz & Taylor 1999; Mori et al. 2000;
Takamatsu et al. 2000). Therefore, Braun and Takamatsu
(2000) proposed to merge the two genera into an emended
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51
Fig. 33 Maximum parsimony
tree based on a combined ITS
and TEF1 sequence dataset of
representing Scleroramularia.
Bootstrap support values for
MP ≥ 50% are given above the
nodes. The tree is rooted with
Phyllosticta lauridiae (CBS
145559). The new isolates are
in bold
Podosphaera yulii S.-Y. Liu & P.-L. Qiu, sp. nov.
Index Fungorum number: IF556431; Facesoffungi number: FoF 06104; Fig. 36
Etymology: Refers to the name of Prof. Yu Li.
Holotype: HMJAU91796.
Mycelium amphigenous, effuse, persistent, often causing yellowish or dark brown discoloration and malformation of the hosts leaves. Hyphal appressoria indistinct
nipple-shaped, solitary. Conidiophores arising from the
upper surface of hyphal mother cells, (45–)50–105 × 9–12
μm and producing the conidia in chains with clear fibrosin
bodies. Foot cells, cylindrical, 22–51(–56) × 7.8–12.7 μm,
sometimes slightly swollen at the very base and the basal
septum occasionally raising above the junction with the
mother cell. Conidia ellipsoid or oval, with distinct fibrosin
bodies, 18.6–29.1(–36.5) × 14–21 μm with a length/width
ratio of mostly 1.2–2.1. Germ tubes of conidial germination, clavate or somewhat forked, straight or most curved,
producing in single, opposite or double at the one end or
both ends of the conidia. Chasmothecia 69–93 μm diam,
amphigenous, gregarious, including numerous yellowish
droplets, and containing one ascus each. Peridium cells of
chasmothecia, irregularly polygonal. Appendages, 6–22 per
chasmothecium, (41.7–)50–108(–148.4) μm, dark brown at
the base, becoming paler towards the apex, apices dichotomously branched, 0.5–1.7 times the chasmothecial diam.
Asci oval to subglobose, 49.3–75.5 × 46.5–65.2 μm, sessile
or with a very short stalk, terminal oculus 5.5–17(–24) μm
wide. Ascospores 8 per ascus, curved, crescent-shaped, very
few ellipsoid, 18–25 × 9.2–13.2 μm, colorless, and emerging
yellowish oil drops after heating in lactic acid.
Material examined: CHINA, Jilin Province, Changchun,
Jilin Agricultural University, on Crataegus pinnatifida, 1
June 2018, P.L. Qiu (HMJAU91796, holotype).
Additional specimens examined: On Crataegus pinnatifida – CHINA, Heilongjiang, Heihe City, 1 September 2017,
F.Y. Zhao & V. Nguyen HMJAU91798, HMJAU91799; Jilin
Province, Changchun City, Jilin Agricultural University, 5
July 2018, P.L. Qiu (HMJAU91797).
GenBank numbers: ITS: MK409979, MK409981,
MK409982, MK409980; LSU: MK734073, MK734075,
MK734076, MK734074; IGS: MK955881.
Notes: Maximum parsimony phylogenetic analysis based
on the ITS sequences demonstrated that Podosphaera yulii
formed a single clade with 100% bootstrap support that is
distant from P. clandestina, P. curvispora and P. paracurvispora (Fig. 37). The molecular analysis of 28S rDNA
sequences suggested the four species clustered with each
other but without bootstrap support (Fig. 38). The morphological characteristics and molecular analysis support
that the powdery mildew on C. pinnatifida in this study is
a new and undescribed species. P. yulii is clearly different
from P. clandestina by having mostly curved ascospores,
but a few straight, short foot-cells (up to 56 μm in P. yulii vs
35–100(–125) μm in P. clandestina) and more appendages
(6–22 per chasmothecium in P. yulii vs 5–15 in P. clandestina). P. paracurvispora and P. curvispora also have curved
ascospores. Tang et al. (2017) recorded that P. paracurvispora has foot-cells (13–)20–27(–32) μm in length which
is shorter than P. yulii with foot-cells 22–51(–56) μm, and
pale brown, curved, crescent-shaped ascospores that differ from P. yulii having curved and colorless ascospores.
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Fungal Diversity (2020) 104:1–266
Fig. 34 Graphis supracola (MFLU 19 − 0013, a new sequenced species). a − c Habitus. d. Thallus with P + orange. e − f Vertical section
through an ascoma. e Thick section, excipulum black. f Thin section,
excipulum looks brown. g Structure of excipulum in thin section.
h − j Ascus; j Ascus in Melzer’s reagent. k Hymenium, clear. l − q
Ascosproes; l − m Ascospores in water. n − p Ascospores in Lugol’s
iodine, which present yellow to blue. q Ascospore in Melzer’s reagent
which presents violet. Scale bars: a = 1 mm, b = 500 μm, c,d = 200
μm, e − f = 50 μm, g − j = 20 μm, k − q = 10 μm
The conidial germination type (Fibroidium-type, singly
formed subapically or laterally) within P. paracurvispora
is also different from P. yulii. No specific anamorphic characters of P. curvispora were described in Braun and Cook
(2012). P. yulii differs from P. curvispora by having smaller
chasmothecia (69–93 μm diam vs (70–)100–150(–170) μm,
Braun and Cook 2012).
13
Golovinomyces (U. Braun) Heluta, Biol. Zhurn. (Armenii)
41(1): 357 (1988)
Fungal Diversity (2020) 104:1–266
53
Fig. 35 Phylogram generated from maximum likelihood analysis
based on a combined mtSSU and nuLSU sequence dataset in the
tribe Graphideae (subfamily Graphidoideae). Redonographa chilensis (Tehler 8407) and R. saxiseda (Tehler 8370) in the subfamily
Redonographoideae were used as outgroup. Bootstrap values for
maximum likelihood (ML) ≥ 75% are placed above the branches.
Branches with Bayesian posterior probabilities (PP) ≥ 0.95 are in
bold. The newly generated sequence is indicated in blue
Notes: Golovinomyces had been treated as one of the three
sections of genus Erysiphe (Braun 1978). Braun (1999) proposed to raise Erysiphe sect. Golovinomyces to the generic
level with Erysiphe cichoracearum DC. as the type species,
and moved it to a newly established tribe Golovinomyceteae
based on the distinct anamorphic characters and the results
of molecular phylogenetic analyses (Braun and Takamatsu
2000). Up to 2283 species of plants from 58 families mostly
herbaceous plants such as the Asteraceae, Bolaginaceae,
Scrophulariaceae and Cucurbitaceae can be infected by
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Fungal Diversity (2020) 104:1–266
Fig. 36 Podosphaera yulii (HMJAU91796, holotype) on Crataegus
pinnatifida. a Nipple-shaped hyphal appressoria. b Conidiophore. c
Elliptical conidia. d–k Germ tubes of conidial germination. l Chasmothecia. m Peridium cells. n Ascus with 8 curved ascospores. o
Ascus after heating. p Ascospore curved and colorless. q Ascospore
with yellowish droplet after heating. r Droplet. Scale bars: a–o = 20
μm, p–r = 10 μm
species of Golovinomyces (Amano, 1986). Many studies
with morphological and molecular phylogenetic analyses
of this genus were conducted including the studies of evolutionary lineages of the G. cichoracearum complex (Cunnington et al. 2010), evolution of host-parasite relationships
of Golovinomyces (Matsuda and Takamatsu 2003; Takamatsu et al. 2013), the spliting of the complex G. biocellatus (Scholler et al. 2016), G. cynoglossi (Braun et al.
2018), G. orontii (Braun et al. 2019) and G. ambrosiae (Qiu
et al. 2020a), and the epitypification of G. tabaci (Qiu et al.
2020b). G. ambrosiae, G. spadiceus and G. circumfusus can
not be split based only on the ITS and 28S rDNA sequences
(Takamatsu et al. 2013). Recent research on the complex G.
ambrosiae conducted by Qiu et al. (2020a) illustrated that
multilocus phylogeny and further morphological examinations are necessary to split this complex.
13
Golovinomyces monardae (G.S. Nagy) M. Scholler, U.
Braun & Anke Schmidt, in Scholler et al., Mycol. Progr.
15(no. 56): 4 (2016) Fig. 39
White powdery layers effuse, on both sides of leaves,
but more evident on the upper leaf surface. Hyphal
Fungal Diversity (2020) 104:1–266
55
Fig. 37 Phylogram generated from maximum parsimony analysis based on ITS sequences of Podosphaera. Cystotheca wrightii
(AB000932) was used as outgroup. Bootstrap values for maxi-
mum parsimony ≥ 50% were shown on the respective branches. The
sequences of new taxon are shown in boldface
appressoria nipple-shaped, solitary. Conidiophores (50.3–)
62.5–110 × 9.2–13.5 μm, arising from the upper surface of
hyphal mother cells. Foot cells 31.6–81.0 × 8.8–13.6 μm,
mostly curved at the base, basal septum of the foot-cell
partly raised above the junction with the hyphal mother
cell up to 14.6 μm, foot-cell followed by 1–3 shorter cells,
forming catenescent conidia; conidia mostly ellipsoid, partly
doliiform, 24.5–40.8 × 14.4–19.4 μm, length/width ratio
1.4–2.4; germ tubes mostly short to moderately long, terminating simply or in a nipple-shaped appressorium, producing
at the top or perihilar of the conidia.
Material examined: CHINA, Jilin Province, Changchun, Jilin Agricultural University, on Monarda didyma, 22
August 2018, P.L. Qiu (HMJAU-PM91795).
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Fungal Diversity (2020) 104:1–266
Fig. 38 Phylogram generated
from maximum parsimony analysis based on LSU sequences
of Podosphaera. Cystotheca
wrightii (AB022355) was used
as outgroup. Bootstrap values
for maximum parsimony ≥ 50%
were shown on the respective
branches. The sequences of new
taxon are shown in boldface
GenBank numbers: ITS: MK414680; LSU: MK951687;
IGS: MK955882.
Notes: Golovinomyces monardae sensu stricto was split
from the G. biocellatus complex by Scholler et al. (2016),
based on the distinct morphology and phylogenetic analysis.
The species is mostly found on Lamiaceae and occasionally on Verbenaceae plants. The phylogenetic analysis based
on ITS sequences demonstrated that the powdery mildew
found on M. didyma is G. monardae which is distinct from
G. neosalviae, G. salviae and G. biocellatus (Fig. 40). The
current fungus has the characters of G. monardae in Scholler et al. (2016).
Pezizomycetes
Pezizales J. Schröt.
Notes: See Ekanayaka et al. (2018).
13
Pezizaceae Dumort., Syst. Mycol. (Lundae) 3(1): 72 (1829)
Notes: Pezizaceae was established by Dumortier (1829)
with Peziza Fr. as the type genus. A systematic overview
of the family was given by Jaklitsch et al. (2016) who recognized 32 genera in this family. Ekanayaka et al. (2018)
reviewed the families in the class Pezizomycetes and outlined
45 genera belonging to Pezizaceae, including also some taxa
synonymized by other authors. Since the first phylogenetic
studies, the rather large genus Peziza Dill. ex Fr., has been
shown to be genetically very heterogenous (Norman and
Egger 1999; Hansen et al. 2001, 2005). The species related
to the type species of the genus, P. vesiculosa Bull., were
proposed as a core group for future recircumscription of
Peziza (Hansen et al. 2002). Multigene analysis (Hansen
et al. 2005) defined 14 lineages within Pezizaceae, where
eight of them contain Peziza sensu lato representatives
Fungal Diversity (2020) 104:1–266
57
Fig. 39 Symptom of powdery moldew on Monarda didyma and
microscopic structures of Golovinomyces monardae (HMJAUPM91795, new geographical record). a White, effused powdery lay-
ers on diseased plant leaf. b Hyphal appressorium with distinct nipple-shaped. c Conidiophore. d, e conidia. f, g Geminated conidium.
Scale bars: a–g = 20 µm
spread among other genera. Six of those eight lineages correspond to genera that were adopted by Boudier (1907):
Aleuria (Fr.) Gillet (Peziza sensu stricto), Galactinia
(Cooke) Boud., Lepidotia Boud., Pachyella Boud., Plicaria Fuckel and Sarcosphaera Auersw. (Pfister 2015b). Of
these, modern phylogeny showed that only Galactinia sensu
Boudier is polyphyletic. An integrative revision of the whole
group combining non-molecular characters with phylogeny
has still not been done. Recently, some authors started to
adopt narrower taxonomical concept using several existing names (i.e. Pfister et al. 2016; Van Vooren et al. 2015,
etc.). Based on a multiple gene phylogeny and phenetic evidences, we describe two new genera for species classified
earlier as “Peziza” for which no name is available: Ionopezia
Matočec, I. Kušan & Jadan for Peziza gerardii Cooke and
Hansenopezia Matočec, I. Kušan & Jadan for Peziza retrocurvata K. Hansen & Sandal. In addition, we present a new
species Hansenopezia decora Matočec, I. Kušan & Jadan.
Ionopezia Matočec, I. Kušan & Jadan, gen. nov.
Index Fungorum number: IF556250; Facesoffungi number: FoF 07927
Etymology: Gr. ιον- (ion-) = violet and lat. pezia = sessile
fungus
Type species: Ionopezia gerardii (Cooke) Matočec, I.
Kušan & Jadan
Ascomata apothecial, solitary or in small groups, comparatively small-sized, epigeous, shortly stipitate, mostly regularly circular from the top view, cupulate when young, then
shallowly saucer-shaped to plate-shaped, hymenial surface
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Fig. 40 Maximum parsimony tree based on ITS sequences of Golovinomyces. Bootstrap support values ≥ 50% are given near nodes respectively.
The tree is rooted with Parauncinula septata (MUMH585). The new strain is in bold
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greyish violet to vividly violet, matte, not wrinkled, margin
upright at first, then flattened, beset with whitish floccose
granules on pale greyish violet ground, excipular surface
concolourous with the margin, beset with less floccose granules. On section hymenium creamy grey, subhymenium differentiated as a thin line due to dark violet pigmentation,
flesh creamy grey, not exuding latex, stipe base without
visible subiculum. Asci cylindrical to slightly ventricose,
hyaline, apex subtruncate-rounded, protruding above paraphysis tips at full maturity, 8-spored, base tapered, pleurorhynchous, arising from croziers, operculum apical, functional,
only faintly predelimited before opening, barely thicker than
surrounding wall, not lenticular, not encircled by indentation
ring, periascal amyloid mucus thin, film-like, evenly thick
throughout; in Lugol’s solution periascal mucus amyloid,
reaction of medium strength in upper ascus area, intensity
gradually decreasing towards the base. Ascospores hyaline,
fusiform to naviform, with subacute poles, slightly asymmetrical, 1-celled, smooth under immersion lens when in
water mount, unstained longitudinal striae visible in Cotton
Blue, perispore not cyanophilic or not present, thin-walled,
without sheath when spores are ejected; sporoplasm multiguttulate, lipid content maximal with 2–3 large lipid bodies
and many smaller ones, uninucleate, nucleus and nucleolus not contrasted nor stained in Lugol’s solution, de Bary
bubbles absent in anhydrous media. Paraphyses cylindrical,
apically obtuse, ± straight, not branching in the upper part,
embedded in hyaline epithecial exudate, containing nonrefractive KOH-soluble and HCl-inert coarse deep purplish
violet granules and paraphysal ectochroic plaques, apical
cell infrequently contains a few small moderately refractive
hyaline granules, otherwise content non-refractive, Lugol’s
solution provokes formation of refractive sienna purple to
brownish purple granules in paraphysis cytoplasm. Marginal
texture composed of compact perihymenial area with thinwalled, hyaline narrowly prismatic cells, encrusted with
dark purple to violet plaques, and of proximal/distal margin mostly occupied with thin-walled, hyaline cell chains,
forming hyphoid fascicles embedded in gel; texture not
cyanophilous. Subhymenium composed of textura angularis-epidermoidea, cells thin-walled and hyaline, richly beset
by non-refractive KOH-soluble and HCl-inert coarse deep
purplish violet granules and ectochroic plaques, the same
as on the top of the hymenium. Medullary excipulum with
upper part composed of textura ellipsoidea-epidermoidea,
with cells bearing scattered dark purple to violet–purple
plaques, and lower part of textura intricata-porrecta to
intricata-prismatica running ± parallel with the surface,
principally without any pigment or refractive intercellular
content; texture not cyanophilous. Ectal excipulum of two
layers: discontinuous cortex composed of cell aggregates
of textura globulosa-angularis, with individual cells globose, subangular to shortly clavate, walls hyaline thickened,
59
without any internal refractive or pigment cell inclusions,
intercellular hyaline gel matrix conspicuous; continuous
part composed of textura angularis-prismatica, cell chains
directed ± perpendicularly to the surface, cells ellipsoid,
clavate to prismatic, hyaline, thin-walled, refractive or pigment cell inclusions absent.
Ecology: Terricolous, directly on forest soil, sometimes
with mosses, on river banks and pathway edges, inhabiting
biomes from colline to arcto-alpine zone of Northern Hemisphere, fruitbodies appear from spring to autumn.
Notes: We establish the new genus Ionopezia for a single
species, Peziza gerardii Cooke. In all phylogenies done so
far (e.g. Hansen et al. 2005; this paper), P. gerardii forms a
well-supported lineage with Marcelleina Brumm., Korf &
Rifai as a sister group to the rest of the Pezizaceae (Fig. 41).
Due to the lack of amyloid reaction of the asci Marcelleina
was not considered as a member of the Pezizaceae for a long
time (e.g. Hawksworth et al. 1995; Eriksson et al. 2001)
until the first phylogenetic analyses (Hansen et al. 2001).
Our observations based on living material of ‘Peziza gerardii’ show that there are numerous unique traits that are synapomorphic either to the newly established genus Ionopezia
or that are shared with the sister genus Marcelleina. The
non-refractive deep purplish violet pigments, heavily deposited in intercellular spaces of the subhymenial layer and/or
ectochroic, adpressed as patches to a cell wall (Fig. 42g, k,
m), irreversibly soluble in KOH without any colour reaction
and inert in HCl, is a unique character shared only by the
genera Ionopezia and Marcelleina. This type of character
is mostly present as cell wall patches in the latter genus.
The same type of pigment is richly embedded also in the
epithecial amorphous layer, marginal plaques, and to a lesser
extent in the intercellular spaces of the medullar layer. The
ascospore wall and the ornamentation in Ionopezia are the
characters that distinguish it most within the Pezizaceae. The
wall is not cyanophilous and the perispore might be absent,
while the surface is finely longitudinally wrinkled (Fig. 43k).
Similar spore wall properties are found in some Morchella
species. Furthermore, ascospores in Ionopezia are fusiform
to naviculate and filled with numerous lipid bodies, a combination of characters not found in other species of Pezizaceae.
The structure of the apothecial margin is also diagnostic. It
has a sharp outline in section, and is nearly entirely composed of hyphoid fascicles (cylindrical to narrowly clavate)
embedded in abundant gel plaques. Another distinctive character attributable only to the genus Ionopezia is the appearance of refractive sienna purple to brownish purple granules
in the paraphysis cytoplasm when Lugol’s solution is added
to a tap water mount (Fig. 43h, i). A refractive, hyaline to
yellowish granules or guttules, readily visible in a tap water
mount that turn to rusty brown or brownish red in Lugol’s
solution were determined to be glycogene accumulations
by Baral (1992). The former phenomenon differs by the
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Fig. 41 Phylogram generated from maximum likelihood analysis based on a combined ITS, LSU and RPB2 sequence dataset for
Hansenopezia decora and Ionopezia gerardii and other related
sequences in Pezizaceae retrieved from the GenBank. Ascobolus
crenulatus KH.02.005 (C) is used as the outgroup taxon. ML bootstrap values ≥ 50% are given as the first set of numbers and approxi-
mate likelihood-ratio test (aLRT) ≥ 0.90 values as the second set of
numbers above the nodes. Voucher/strain numbers are given after the
taxon names, the ones from type material are indicated in bold face.
Newly generated sequences are indicated in blue. The bar length indicates the number of nucleotide substitutions per site
fact that nothing is discernable in the paraphysis cytoplasm
before Lugol’s solution is applied. There is also a difference
in colour of these two types of cell inclusions. It is also
worth mentioning that the fine structure of the ascus wall
and its cytochemistry are also aberrant in the Pezizaceae,
especially concerning the epigeous members. The operculum of Ionopezia is not at all lentiform, but is encircled only
by a slight, barely visible optical discontinuation in the wall
structure. This character is shared by Marcelleina. The periascal mucus is very thin and film-like, continuing through
the entire ascus length demonstrating weak amyloidity all
over the surface, corresponding to the IIIa type as defined
by Hansen et al. (2001). The overall cyanophobic property
of the apothecial texture is shared only by few species in the
Pezizaceae known to the authors (e.g. Hansenopezia, Peziza
simplex Dougoud & Moyne, etc.).
Peziza gerardii Cooke was established by Cooke (1875b)
who revised Gerard’s material of P. violacea W.R. Gerard
(1873, in litt.) which is a nomen illegitimum (non P. violacea Pers. 1794). Cooke’s description (Cooke 1875a, 1875b)
includes spore dimensions 32–35 × 8–9 µm which does not
agree with Cooke’s drawing (Cooke 1874–1875) of spores
inside the ascus drawn to scale. Thus, from his drawing
spores are 27–32 × 8.5–9.5 µm. The original collection of
Gerard was restudied by Pfister (1978) who gives a description of P. gerardii based both on Gerard’s collection and
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Fig. 42 Ionopezia gerardii. a-f Ascomata. g Vertical median section
through apothecium. h Perihymenium and proximal margin. i Terminal cells on extreme margin. j Medullary excipulum. k Contact zone
between subhymenium and medullary excipulum. l Terminal cells of
ectal excipulum with Lugol induced bodies. m Subhymenium. n Cell
aggregates on ectal excipulum. o Ectal excipulum. g–k, m–o In water
61
mount (*), l In Lugol’s solution (*). g In dark field, a–f, h–o In bright
field. a, e, f–o From CNF 2/10798, b, d From CNF 2/7450, c From
CNF 2/9431. Photo: N. Matočec and I. Kušan. Scale bars: a, d = 5
mm, c = 2 mm, b, e, f = 1 mm, g = 100 µm, h, n, o = 50 µm, i–m = 10
µm. Asterisk (*) denotes living material
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Fig. 43 Ionopezia gerardii. a–c Living mature asci with paraphyses.
d Ascus bases with ascogenous cells. e–g Paraphyses with epithecial
gel. h, i Paraphyses with Lugol induced granules. j, k Ascospores. l
Terminal cells on ectal excipulum. m Margin. a, b, d–g, j In water
mount (*), c, h, i In Lugol’s solution (*), k-m In Cotton Blue (†).
All from CNF 2/10798. Photo: N. Matočec and I. Kušan. Scale bars:
a–m = 10 µm. Asterisk (*) denotes living material. Cross (†) denotes
dead material
the collection of von Höhnel’s from the herbarium at FH,
giving spores as 28–32(–35) × 7–9 µm. On the other hand,
Van Vooren (2009) described two collections having a spore
range of (23–)24–32 × 9–10.2 µm and one with even smaller
spores (17–)18–21 × 8–10 µm. According to today’s perception this species varies greatly in the length of the spores.
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Below we give a description of Ionopezia gerardii based on
samples collected in Croatia and Bosnia and Herzegovina.
Ionopezia gerardii (Cooke) Matočec, I. Kušan & Jadan,
comb. nov.
Index Fungorum number: IF556252; Facesoffungi number: FoF 07862; Figs. 42, 43
Basionym: Peziza gerardii Cooke, Hedwigia 14(6): 81
(1875)
= Peziza ionella Quél., Bull. Soc. Bot. Fr. 24: 328 (1878)
[1877]
= Plicaria pedicellata Velen., Novitates Mycol.: 198
(1940)
For complete synonymy see Pfister (1978) and Van
Vooren (2009).
Ascomata apothecial, individual, circular from the top
view, larger fruitbodies can be slightly irregular from the top
view, cupulate when young, then shallowly saucer-shaped
to plate-shaped, short stipitate, 3.8–6.8 mm diam, hymenial
surface greyish violet to vividly violet, matte, not wrinkled,
margin upright at first, then flattened, beset with whitish
floccose granules on pale greyish violet ground, granules
up to 0.5 mm tall; excipulum surface concolourous with the
margin, beset with less floccose granules; on section hymenium creamy grey, subhymenium differentiated due to dark
violet pigmentation, flesh creamy grey, not exuding latex,
marginal and excipular flank granules whitish and conical,
stipe-like base 1.9–2.1 mm wide, 0.5–0.7 mm high, without pronounced subiculum. Hymenium 330–380 µm thick,
arranged as a regular palisade. Subhymenium 70–90 µm
thick, composed of textura angularis-epidermoidea, cells
5.1–15.7 µm wide, thin-walled and hyaline, but richly beset
by dark violet–purple pigment, cells often contain one to
few strongly refractive globules, due to the high concentration of pigment well discerned from the medulla. Marginal
texture differentiated into perihymenial area, extreme margin/proximal margin as a whole, distal margin and marginal
medulla: (1) perihymenial area wholly compact, composed
of narrowly prismatic cells arranged as textura porrecta
with thin and hyaline cell walls, encrusted with dark purple
to violet plaques and intercellular diffuse purplish matrix,
especially on a surface, cells 3.5–7.8 µm wide, this area is
rather unclearly distinguished from; (2) proximal margin,
composed of wider prismatic cells forming textura prismatica oriented perpendicularly to the surface with protruding individual elements, cells are thin-walled and hyaline,
with less pigments and more abundant intercellular matrix,
cells 5.6–12.2 µm wide, terminal cells often clavate, these
cell chains arising from basal; (3) marginal medulla textura
angularis with cells 6.5–16.2 µm wide, hyaline and thinwalled but fairly rich in purple-violet plaques and diffuse
pale purple pigment in intercellular spaces; (4) extreme margin formed as a single pustule, 75–130 µm wide and 18–74
63
µm high, composed entirely as proximal margin; (5) distal
margin of 2–4 pustules, 75–115 µm wide, 15–70 µm high,
composed of hyaline and thin-walled shortly prismatic cells
to ellipsoid to clavate ones, rather untidily organized, cells
12.4–29.1 × 7.2–17.4 µm, entirely without any refractive/
pigment inclusions, cell chains in distal marginal pustules
originate from basal marginal medullar textura angularis
with cells 7–11.2 µm wide in deeper areas and 8–18 µm
wide in the area immediately below the pustules, this area is
very rich in both plaque-form darker consolidated pigments
as well in diffuse ones. Medullary excipulum unclearly
differentiated into two sublayers, altogether 212–260 µm
thick, upper part 160–185 µm thick, composed of textura
ellipsoidea-epidermoidea, cells 5–22.3 µm, mostly thinwalled and hyaline, some individual cells thick-walled,
without intracellular refractive/pigment inclusions but rich
in diffuse pale violet–purple intercellular matrix, lower part
38–75 µm thick, composed of textura intricata-porrecta to
intricata-prismatica with wavy cell chains running ± parallel with the surface, hyphae 4–12.5 µm wide, cells thinwalled, hyaline, principally without any pigment/refractive
intra- or intercellular content. Ectal excipulum composed of
weakly developed discontinuous cortex and thick continuous layer; discontinuous part composed of cell aggregates of
textura globulosa-subangularis, 30–95 µm high, 47–105 µm
wide, individual cells globose, subangular to shortly clavate,
8.8–22.5 µm wide, walls hyaline, somewhat thickened, 0.7–2
µm thick, without any internal refractive/pigment inclusions
but with apparent hyaline intercellular matrix; continuous
part 130–173 µm thick, composed of textura angularis-prismatica, cell chains directed ± perpendicularly to the surface,
cells ellipsoid, clavate to prismatic, 16–86.5 × 6.4–37.8 µm,
hyaline, thin-walled, intercellular spaces with some hyaline
matrix only in outermost area, refractive/pigment inclusions
absent. In Cotton Blue overall texture not cyanophilous; in
Lugol’s solution terminal cells of ectal excipulum may contain Lugol-induced bodies; in KOH pigmented granules
and ectochroic plaques soluble; in HCl inert. Paraphyses
cylindrical, apically obtuse, ± straight, not branching in the
upper part, apical cell 28.4–110 × 4.5–7.5 µm, containing
few weakly refractive globules, embedded in subhyaline to
faintly violet, sparse and faintly developed exudate, containing also strongly refractive granules/plaques, not forming continuous layer; Lugol’s solution causes formation of
refractive sienna purple to brownish purple granules in paraphysis cytoplasm. Asci 265–345 × 16.4–18 µm, cylindrical
to slightly ventricose, broadest point is in a lowest zone of
pars sporifera, apex subtruncate-rounded, protruding above
paraphysis tips up to 22.5 µm at full maturity, pars sporifera
97–108 µm, 8-spored, base tapered, pleurorhynchous, arising
from croziers, operculum only faintly predelimited, 4.3–4.6
µm diam, 0.7–0.8 µm thick, strictly apical, flat-lenticular,
encircled by rather broad but very shallow indentation ring,
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64
1–1.1 µm wide, periascal mucus thin, film-like, evenly thick
throughout, 1–2 perimarginal asci contain at least lower 4
globose to subglobose spores; in Lugol’s solution periascal mucus amyloid, reaction of medium strength in upper
ascus area, intensity gradually decreasing towards the base.
Ascospores (20–)21.1–26–27.9(–28.5) × 7.5–8.9–11(–11.2)
µm, Q = 1.98–3.02–3.45(–3.60) (n = 200, in H2O), hyaline,
fusiform to naviform, with subacute poles, even slightly
papillate, slightly asymmetrical, 1-celled, smooth under
immersion lens when in water mount, in Cotton Blue elongated striae visible but not cyanophilic, multiguttulate, with
2–3 large lipid bodies, 3.2–6.2 µm diam, and many smaller
ones, 0.7–2.8 µm diam, uninucleate, nucleus 2.1–2.3 µm
diam, equatorially-peritunically positioned, thin-walled,
wall 0.5 µm thick, freshly ejected without sheath; in Cotton
Blue without de Bary bubbles, perispore not cyanophilic or
not present.
Habitat and phenology: Terricolous, directly on forest
soil, sometimes with mosses, on river banks and pathway
edges, inhabiting biomes from colline to arcto-alpine zone
of Northern Hemisphere, fruitbodies appear from spring to
autumn.
Known distribution: The species is known from North
America, Europe, and India.
Material examined: CROATIA, Zagreb City, Park forest
Maksimir, 45°49′12" N, 16°00′59" E, 120 m a.s.l., Pinus
sylvestris plantation, directly on forest soil, 25 May 1989,
N. Matočec (CNF 2/947); Park forest Jelenovac, 45°49′32"
N, 15°57′03" E, 170 m a.s.l., forest of Quercus petraea and
Carpinus betulus, directly on moist soil near the creek, 14
July 1990, N. Matočec, (CNF 2/1301); Park forest Maksimir,
170 m NE from Vidikovac, 45°49′33" N, 16°01′23" E, 135
m a.s.l., forest of Quercus petraea, Carpinus betulus, on
forest soil with litter, 12 October 2005, N. Matočec and I.
Kušan (CNF 2/7450); Medvednica mountain, vicinity of
Kraljičin zdenac hut, 600 m SE from Medvedgrad castle,
45°51′59" N, 15°56′49" E, 370 m a.s.l., forest of Fagus
sylvatica, Quercus petraea, Carpinus betulus, Castanea
sativa, on forest soil at the edge of a path, 14 July 2013, D.
Mrvoš and N. Matočec (CNF 2/9431); Virovitica-Podravina
County, Dumače area in Bilogora range, 9.1 km SW – S-SW
from Virovitica, 45°45′55" N, 17°19′19" E, 180 m a.s.l.,
forest of Fagus sylvatica, Tilia sp. and Acer sp., forest soil
near the creek, 15 August 1997, S. Gottstein (CNF 2/3493);
BOSNIA AND HERZEGOVINA, Sarajevo County, Igman
mountain plateau, Javornik area, 43°44′54" N, 18°14′04" E,
1480 m a.s.l., forest of Picea abies, Abies alba, Fagus sylvatica, Acer pseudoplatanus, on forest soil at path edge near
creek, 10 August 2018, N. Jukić, I. Kušan and N. Matočec
(CNF 2/10798, duplex FAMU N.J. 100818-Y9).
GenBank numbers: ITS: MK514543; LSU: MK514537;
EF1-α: MK705937; RPB2: MK673768 (CNF 2/10798).
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Hansenopezia Matočec, I. Kušan & Jadan, gen. nov.
Index Fungorum number: IF556255; Facesoffungi number: FoF 07926
Etymology: The genus is named in honour of Karen
Hansen for her great contribution to the understanding of
Pezizaceae phylogeny.
Type species: Hansenopezia retrocurvata (K. Hansen &
Sandal) Matočec, I. Kušan & Jadan, comb. nov.
Index Fungorum number: IF556256; Facesoffungi number: FoF 07937
Basionym: Peziza retrocurvata K. Hansen & Sandal,
Nord. J. Bot. 18(5): 616 (1998)
Other species included (description follows after the
generic description below): Hansenopezia decora Matočec,
I. Kušan & Jadan.
Ascomata apothecial, solitary or often in groups,
medium-sized, epigeous, centrally attached to the substrate, regularly circular to much sinuous from the top
view, cupulate when young, then plate-shaped, hymenial
surface reddish brown, lilac-violet, violet-brown, purplish
brown to dark brown, matte, margin upright then flattened
to reflexed, entire, blunt, smooth and or beset with scattered
white fibrils, whitish grey or pale brownish yellow, excipular ground concolourous with the margin, finely pubescent
or ornamented with reddish brown patches distributed to
the basal parts. Flesh fragile, homogenous, soft waxy, dry
on cut or exuding watery to milk-white latex. Hymenium
arranged as a regular palisade. Asci cylindrical, hyaline, apex
subtruncate, protruding above paraphysis tips at full maturity, 8-spored, base tapered, arising from croziers, operculum apical, very narrow, occupying ≤ 28% of living mature
ascus diam, functional, flat lentiform before opening, encircled by slight “u”-shaped indentation in the wall, periascal
amyloid mucus thin and film-like through the entire length
but almost lacking on opercular lense, while the thickest at
the opercular edge where is moderately euamyloid, forming
unclear thin annular zone, reaction gradually decreasing in
strength towards the base, with reacting area on half of the
ascus length or more. Ascospores hyaline, ellipsoid to narrowly ellipsoid with ± rounded poles, radially symmetrical,
1-celled, subglabrous or with cyanophilic verrucae, perispore cyanophilic, not loosening when heated in lactic Cotton Blue or in KOH, wall 3-layered ± thick-walled, without
sheath on ejected spores; sporoplasm multiguttulate, lipid
content nearly maximal with all guttules of approx. same
size or 2–3 large lipid bodies mixed with many smaller ones,
lipid bodies much coalesced in dead spores, uninucleate,
nucleus and nucleolus not contrasted nor stained in Lugol’s
solution, de Bary bubbles absent in anhydrous media. Paraphyses cylindrical, apically obtuse to subclavate, straight to
bent, not branching in the upper part, embedded in hyaline
epithecial exudate or only with hyaline, thin coating, apical
cell contain single or few non-refractive vacuoles entirely
Fungal Diversity (2020) 104:1–266
filled with finely dispersed minute pinkish brown particles
and/or partially with larger non-refractive granules of the
same colour, not containing vacuolar bodies, in Lugol’s
solution pigment partially turning to rusty brown colour.
Marginal texture in section displaying broad outline, textura
globulosa-angularis mixed with some cylindric cell chains
that may protrude as scattered cylindric obtuse, thin-walled
hyphoid hairs or form loose tufts; texture is not cyanophilous. Subhymenium well-discerned, composed of hyaline subgelatinous densely woven textura porrecta. Medullary and
ectal excipulum of ± homogenous, subhyaline texture, except
for the cortical layer. Upper part composed of textura globulosa-moniliformis mixed with cylindric very long, mostly
vertically oriented hyphae. Lower part composed of textura
globulosa-angularis with intermixed intricate connective
hyphae, cells smaller in size towards the surface. Cortical
layer composed of textura porrecta-intricata, discontinuous
layer when present pustulate, cells with ochraceous walls
and rusty brown patches, coiled and agglutinated together;
texture is not cyanophilous.
Ecology: Saprotroph on rich soil with litter and wood
residues of Abies alba, on very rotten stumps and trunks
of Picea abies and Fagus sylvatica, sometimes also on rich
soil outside forests, fruitbodies appear in spring or autumn
to winter.
Hansenopezia decora Matočec, I. Kušan & Jadan, sp. nov.
Index Fungorum number: IF556257; Facesoffungi number: FoF 07861; Figs. 44, 45
Etymology: Refers to the diverse ornamentation units on
the outer excipular surface.
Holotype: CNF 2/10621.
Ascomata apothecial, epigeous, cup-shaped when young,
expanding when ripe becoming plate-shaped or strongly
sinuous, with a central depression, centrally attached to the
substrate, deeply buried if the litter is thick, solitary or in
small groups, 11–55 mm diam, hymenial surface lilac-violet, matte, in the central part lighter and with pale brownish
tinge with age, constantly violet towards the margin, margin
upright to flattened, never bent outwards, entire, finely whitish pruinose, ground concolourous with excipular surface,
excipulum greyish white, finely whitish pruinose and covered with reddish brown irregular patches throughout, flesh
of homogenous soft waxy consistence, 3–3.8 mm thick at the
base and 1.1–3 mm at half way to the margin (incl. hymenium), upon cutting exuding watery to milk-white latex,
under UV light at 254 nm flesh obscurely violet-lilac, excipulum dull greyish ochre, hymenium negative, at 366 nm flesh
very dull greyish olivaceous, excipulum the same as flesh,
hymenium sepia to umber brown. Hymenium 300–345 µm
thick, arranged as a regular palisade. Subhymenium 82–123
µm thick, well-discerned from the upper medulla, composed
of densely woven hyaline subgelatinous textura porrecta,
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cells 4.6–9.6 µm wide. Marginal and submarginal area beset
with scattered hyphoid hairs or forming loose tufts, distributed up to the upper flank, usually far protruding, flexuous,
often sinuous or spirally arranged, not branched, cylindric
obtuse, 1–4 celled, 28.6–306 × 4.6–7 µm, walls hyaline and
delicate, extreme margin is formed as (1)2–5 paraphysoid
pustules, each cell fascicle is composed of 2–5 cell chain
rows, with 1–2 cells protruding, cells hyaline short prismatic, terminal cells cylindric obtuse to cylindric clavate
or even subglobose, 16.3–44.8 × 7.5–15.6 µm, containing
dotted aggregates of purplish sepia pigment. Medullary and
ectal excipulum ± homogenous, except for the cortical layer,
1780–2500 µm thick in middle flank. Upper part not sharply
delimited, composed of textura globulosa-moniliformis
mixed with cylindric very long mostly vertically oriented
lactiferous hyphae whose cytoplasm is moderately refractive
and ± homogenous, rarely colloid-granulose, globose cells
16.6–67.7 µm diam, hyphae 8.2–15.6 µm wide. Lower part
composed of textura globulosa intermixed with intricate
connective hyphae which are not lactiferous, globose cells
14.9–72.3 µm diam, intricate hyphae 4.6–9.8 µm wide. Cortical layer of ectal excipulum formed as continuous and discontinuous layers, continuous layers composed of subhyaline
textura porrecta-intricata, 38–85 µm thick, hyphae 4.5–9.3
µm wide, discontinuous layer formed of flat-granular to subconical pustules 49–280 × 30.4–165 µm, composed of subhyaline smooth-walled short celled hyphoid to cuboid cells,
9.2–41.2 × 7–10.8 µm, cells coiled and agglutinated together
and combined with scattered or clustered short prismatic
to cuboid cell(s) with ochraceous walls bearing ectochroic
rusty brown patches, these cells 10.6–27.6 × 11.2–13.5 µm.
In Lugol’s solution lactiferous hyphae strongly cadmium
yellow to rusty orange, submarginal and excipular terminal
hairs yellow-orange to rusty orange; in brilliant Cresyl Blue
subhymenium lilac (gelified), lactiferous hyphae greyish
violet to dark violaceous grey, dark purplish globules present in the cortical layer, hyphoid marginal and submarginal
hyphoid hairs with purplish violet vacuoles, cortical coiled
cell chains with dark greyish blue plaques on cyan walls; in
Cotton Blue all textures cyanophobic except for the prosenchymatous subhymenium layer with cell cytoplasm lightly
bluish, pigment in paraphyses and paraphysoid pustules as
well as terminal excipulum cell wall dark grey. Paraphyses
cylindrical, apically obtuse to subclavate, straight to slightly
bent, not branching in the upper part, apical cell 16.7–49.4(–
72.6) × 3.3–6.1(–7.1) µm, with non-refractive vacuoles completely filled with dot-shaped pinkish brown dense freely
floating pigment granules, or contain non-refrative granules
with the same pigment type; in Lugol’s solution pigment
brownish red; in Congo Red pigments greyish red; in brilliant Cresyl Blue vacuoles with pigments greyish violet. Asci
(232–)262–345 × 13.5–15.4 µm, cylindrical, hyaline, apex
subtruncate, protruding above paraphysis tips up to 40 µm
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Fig. 44 Hansenopezia decora. a-f Ascomata. g Vertical median section through excipular middle flank. h Marginal tissue. i Terminal
cells of excipular pustule. g–i In water mount (*). a–c, g, i From CNF
2/10621 (holotype), d–f From CNF 2/10618, h From CNF 2/7823.
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Fungal Diversity (2020) 104:1–266
Photo: N. Matočec and I. Kušan. Scale bars: a, b, d, e = 1 cm, c = 0.5
cm, f = 0.1 cm, g = 100 µm, h, i = 50 µm. Asterisk (*) denotes living
material
Fungal Diversity (2020) 104:1–266
67
Fig. 45 Hansenopezia decora. a Contact zone of subhymenium
and medullary excipulum. b Medullary excipulum. c Ectal excipulum with a pustule. d Living asci with ascospores. e Ascus bases
with ascogenous cells. f Paraphyses. g Amyloidity of ascal mucus. h
Ascospores. a–f, h In water mount (*), g In Lugol’s solution (*/†). a,
b phase contrast, c–h bright field. All from CNF 2/10621 (holotype).
Photo: N. Matočec and I. Kušan. Scale bars: a–c = 50 µm, d–h = 10
µm. Asterisk (*) denotes living material. Cross (†) denotes dead
material
at full maturity, pars sporifera 74.5–114 µm, 8-spored, base
pleurorhynchous, arising mostly from non-repetitive croziers, operculum narrow, 3.7–4.2 µm diam, 0.7–0.8 µm thick,
strictly apical, flat-lentiform, with pronounced “u”-shaped
indentation ring, periascal mucus nearly lacking above operculum and thickest in immediate vicinity of opercular edge;
in Lugol’s solution periascal mucus amyloid, reactive zone
strongest immediately at the opercular edge, intensity gradually decreasing towards the base; in Congo Red operculum
yellowish rosy red, indentation zone well visible due to its
discolouration, median wall layer rutile- to blood-red, outermost wall layer dark red; in brilliant Cresyl Blue periascal
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68
mucus sharply visible, wall unstained. Ascospores 13–15
.2–16.1(–16.8) × (7–)7.3–8–9.1(–9.9) µm, Q = (1.55–)1.
57–1.81–1.99(–2.15) (n = 150, in H2O), hyaline, ellipsoid
to narrowly ellipsoid, rarely oblong, radially symmetrical,
1-celled, seemingly smooth under immersion lens when in
water mount, in brilliant Cresyl Blue and Cotton Blue finely
roughened but not cyanophilic, sporoplasm multiguttulate
with almost maximal lipid content, spores when freshly
ejected with 2–3 larger lipid bodies, 3.3–4.8 µm diam, mixed
with smaller ones, 1.3–2.5 µm diam, lipid bodies in KOH
readily coalesce in one large irregular mass, uninucleate,
nucleus 2.7–3 µm diam, equatorially and eccentrically positioned, wall clearly 3-layered, 0.7–0.8 µm thick; in Lugol’s
solution without glycogene accumulations, lipid bodies coalesce immediately, nuclei not contrasted; in KOH wall stable
(not loosening); in Congo Red not stained; in brilliant Cresyl
Blue perispore dark blue, middle layer unstained, innermost
layer pale violet; in Cotton Blue perispore cyanophilic and
not loosened, smooth to very finely roughish and/or irregularly wrinkled, de Bary bubbles not formed.
Habitat and phenology: Saprotroph on rich soil with litter and wood residues of Abies alba (Pinaceae), mostly in
timber storages, in altimontane forests with Abies alba, fruitbodies appear in April and May.
Known distribution: The species is known so far only
from Gorski kotar region, Croatia.
Material examined: CROATIA, Primorje-Gorski kotar
County, Široka draga area, 4.1 km S from Mrkopalj,
45°16′45" N, 14°51′21" E, 930 m a.s.l., forest of Abies alba,
Fagus sylvatica and Acer pseudoplatanus, on very rotten
branches of Abies alba covered with mosses, 17 May 2002,
N. Matočec and D. Mrvoš (CNF 2/5619); Ravna grabovača
area near Gomirje (Vrbovsko), 3.6 km S from Vrbovsko,
45°20′29" N, 15°04′33" E, 500 m a.s.l., forest of Abies
alba and Fagus sylvatica, pebble soil mixed with wood
residues in storage of timber, 7 April 2007, N. Matočec,
(CNF 2/7823); forest Gorica near Gomirje (Vrbovsko), 3.4
km S-SE from Vrbovsko, 45°20′51" N, 15°05′57" E, 410 m
a.s.l., forest of Abies alba with some Picea abies, on soil
with litter and wood residues, 2 May 2009, N. Matočec and
D. Mrvoš (CNF 2/8168); Široka draga area, 2.5 km S-SW
from Mrkopalj, 45°17′44" N, 14°50′27" E, 900 m a.s.l., forest of Abies alba, Fagus sylvatica, Picea abies, on rich soil
beset with Abies alba litter, 26 April 2016, N. Matočec and
I. Kušan (CNF 2/9896); Široka draga area, 2.5 km S-SW
from Mrkopalj, 45°17′41" N, 14°50′30" E, 900 m a.s.l., forest of Abies alba, Fagus sylvatica, Picea abies, on rich soil
with Abies alba litter in a storage of timber, 16 May 2018, I.
Kušan and N. Matočec (CNF 2/10618); Široka draga area, 4
km S from Mrkopalj, 45°16′47" N, 14°51′06" E, 930 m a.s.l.,
forest of Abies alba, Fagus sylvatica, Acer pseudoplatanus
with some Sambucus nigra, on thick litter of Abies alba in
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a storage of timber, 16 May 2018, I. Kušan and N. Matočec
(CNF 2/10621, holotype)
GenBank numbers: ITS: MK514542; LSU: MK514536;
EF1-α: MK705936; RPB2: MK673767 (CNF 2/10621,
holotype).
Notes: A data matrix for alignment was constructed to
show a phylogenetic position of Hansenopezia Matočec, I.
Kušan & Jadan within the Pezizaceae, with a special emphasis on Peziza. Phylogenetic analysis (Fig. 41) included the
ITS, LSU and RPB2 sequences generated from the type collection of Hansenopezia decora and other related sequences
retrieved from GenBank as well as a newly sequenced collection of Ionopezia gerardii (Cooke) Matočec, I. Kušan &
Jadan (CNF 2/10798). The phylogeny based on a concatenated analysis of ITS, LSU and RPB2 nested Hansenopezia
in the Pezizaceae, embracing two species—H. retrocurvata
(K. Hansen & Sandal) Matočec, I. Kušan & Jadan and H.
decora Matočec, I. Kušan & Jadan next to the Peziza polaripapulata-Iodowynnea, Sarcopeziza sicula, P. saniosa-Terfezia-Tirmania and Peziza phyllogena–Eremiomyces clades
(Fig. 41). H. retrocurvata clusters with high support with
H. decora showing a high level of genetic similarity and is
therefore here combined in a new genus Hansenopezia. Both
species, H. decora and H. retrocurvata demonstrate a distant relationship to the Peziza core-species group clustered
around P. vesiculosa Bull. which is a type species of Peziza,
suggesting taxonomic affinity outside of this genus.
Besides molecular evidence, Hansenopezia decora and H.
retrocurvata are different from Peziza sensu stricto [a core
species group gathered around the type species, P. vesiculosa, cf. Hansen et al. (2001, 2002, 2005)], primarily by
properties of the ascus wall and ascospore cytology when in
the living state. While true members of Peziza (consisting of
three subgroups: a core-species group, a group represented
by P. subviolacea Svrček and one by P. subcitrina (Bres.)
Korf) have asci equipped with a thick lentiform, rather broad
operculum with a diameter of approximately 35–45% of the
ascus diameter, sharply encircled by a “z” indentation ring,
also present in epigean species of the broader P. succosa-P.
succosella clade (pers. data), asci of Hansenopezia have a
very narrow, flat lentiform operculum with a diameter ≤ 28%
of the ascus diameter encircled only by a slight “u” indentation ring (Fig. 45d). The former trait is considered here as
synapomorphy of Peziza sensu stricto, while the latter is
shared only by Sarcosphaera coronaria, a semi-hypogeous
species belonging to a distantly related lineage. These fine
structural traits are visible using a set of light microscopic
techniques including vital differential staining, exclusively
when asci are in living state and fully mature (cf. Baral
1992; Pfister et al. 2009). Another important differential
microscopic marker in pezizacean taxonomy, i.e. the amyloidity of the periascal mucilaginous coat of the ascus wall
was already treated by Hansen et al. (2001) using Melzer’s
Fungal Diversity (2020) 104:1–266
reagent. Her study revealed four types of amyloidity in Pezizaceae (with two variants). However, getting the insight into
the real state of this vulnerable character could be compromised or altered by application of aggressive media and/or
physical conditions (pressure, heat etc., cf. Donadini 1985;
Samuelson 1978) in the process of examination. A character
of similar vulnerability yet very important in species recognition is the ascospore apiculus in Thecotheus Boud., a sister
group of the Pezizaceae, where Melzer’s reagent proved to
be inappropriate for reliable examination of this character,
while Lugol’s solution was ascertained as recommended
because it does not disturb nor alter its natural state (Kušan
et al. 2015). The suppressing effect on hemiamyloidity by
Melzer’s reagent is elaborated by Baral (1987). Bearing in
mind all these risks and problems, Lugol’s solution is here
considered as a primary medium for examination of the periascal coat amyloidity in terms of vital taxonomy research
(Baral 1992). The periascal amyloidity obtained by vital
staining using Lugol’s solution demonstrated sharp differences, but also some similarities (contrary to Hansen et al.
2001) between members of Peziza sensu stricto (with more
than 40 spp. tested) and both species of Hansenopezia. As
described in Hansen et al. (2001), all tested species of Peziza
sensu stricto demonstrated amyloidity of type II (thick apical
ring-shaped but also cap-shaped strong amyloid mucilaginous accumulation; reaction abruptly decreased towards the
base). On the other hand, both species of the genus Hansenopezia showed a thin and film-like periascal coat extending through the entire ascus length but almost lacking on
the opercular surface. The coat is obviously thickest around
the opercular edge where it is moderately amyloid, forming a flat and delimited ring around the operculum, while
the reaction intensity gradually decreased towards the base,
with the reacting area 1/2 of the ascus length or more. The
former character state is confirmed to be synapomorphic
of the genus Peziza sensu stricto, while the latter is shared
by two other lineages (namely Sarcosphaera coronaria and
Peziza phyllogena). The nuclear membrane contrasting and
differentially staining of nucleolus by Lugol’s solution in living mature spores is the third differential cytological marker
useful in separating the two genera. All species of genus
Peziza sensu stricto were positive (nearly 40 spp. tested)
with Lugol’s solution (synapomorphy), while two species
of Hansenopezia were negative (a character shared by the
majority of other pezizacean lineages). Another distinctive
character that separates the two genera is the lipid content
of the living mature spores (freshly ejected spores from living asci are recommended for study). The ascospores of the
genus Peziza sensu stricto are either eguttulate (devoid of
lipid bodies) or biguttulate, rarely with bipolar groups of
lipid bodies, always with lipids occupying less than 50% of
the extranuclear sporoplasm. The ascospores of the genus
Hansenopezia are multiguttulate with maximal lipid content,
69
occupying nearly all the extranuclear sporoplasm. A somewhat lower but still high lipid content in hyaline ellipsoid
spores (though of different configuration) is found also in
other lineages, e.g. Sarcopeziza sicula (cf. Phookamsak
et al. 2019), Sarcosphaera coronaria, Adelphella babingtonii (cf. Pfister et al. 2009) etc. The ascospores of some
species from the broader Peziza phyllogena lineage have an
apparently high lipid content, but there are various levels
of refractive metachromatic corpuscles intermixed with the
true lipid bodies (pers. data). Also, there is a clear distinction in the cyanophilous reaction of the texture in the genus
Peziza sensu stricto and Hansenopezia (Cotton Blue in lactic
acid is applied on the sections of living material, with gentle heating or overnight staining at room temperature). All
tested species of the genus Peziza sensu stricto (ca. 30 spp.)
had at least cyanophilic septal rings in the medullary excipulum, while both species of Hansenopezia had completely
cyanophobic cell walls, a character shared only by the very
distantly related genus Ionopezia. Finally, a very important
exclusive character that separates genus Hansenopezia (synapomorphy) from all other pezizacean genera is that it has
a unique homogenous subhyaline texture composed equally
of vesicular and hyphoid cells that invariably occupy the
whole excipulum between the thin subhymenium and the
reduced cortical layer and between the marginal surface and
the apothecial centre.
Peziza retrocurvata was described by Hansen et al.
(1998) from Danish woodlands mostly dominated by Fagus
sylvatica or Sambucus nigra on calcareous soil. The apothecia were found on very rotten Picea abies stumps. Its
unique characters, such as ellipsoid, hyaline, multiguttulate
ascospores having lipid bodies occupying almost the whole
of the sporoplasm and the homogenous excipulum equally
composed of vesicular cells and cylindric cell chains, positioned this species in a separate lineage in the 3-gene phylogenetic analysis performed by Hansen et al. (2005). More
recent collections were analysed by Van Vooren et al. (2017)
who showed the relationship of P. retrocurvata to milk-exuding species such as P. succosa, P. succosella and P. michelii.
We discovered a new species very similar to P. retrocurvata
whose close relationship was confirmed by molecular analysis and by microscopic evidence obtained by the analysis of
the living material. Along with P. retrocurvata our new species (Hansenopezia decora) is in the genus Hansenopezia.
The most outstanding characters that differentiate Hansenopezia retrocurvata and H. decora are: (a) configuration of
lipid bodies in freshly ejected spores, (b) spore ornamentation, (c) ornamentation of marginal and excipular surfaces
and (d) phenology. In contrast to H. retrocurvata, the marginal and excipular surfaces of H. decora are richly decorated with three types of ornamentation: (1) extreme margin
with pustules composed of small fascicles of hyaline-walled,
broadly clavate-cylindric cell rows of paraphysoid-type
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70
whose cytoplasm is filled with vacuoles containing purplish
sepia granular freely floating pigment, (2) submarginal hyaline, hyphoid, flexuous, delicate-walled, cylindric-obtuse
hairs and (3) excipular flat pustules of irregular fascicles of
brown-walled short-celled hyphae. The excipular surface of
H. retrocurvata is glabrous to only finely pubescent due to
some protruding hyphoid hairs. The ascospore surface in H.
decora is subglabrous with non measurable and non cyanophilic elevations, while the ascospore wall in H. retrocurvata
is densely warted, flat granular, (0.3–)0.6–1.3(–2.3) µm long
and wide and ca. 0.5(0.7) µm high, and cyanophilic. The
lipid bodies in H. decora are highly diverse in size and number; there are only 2–3 large guttules 3.3–4.8 µm diam and
many smaller ones 1.3–2.5 µm diam, while all guttules in H.
retrocurvata are of similar size, ca. 1–2 µm diam. Finally,
H. decora is a strictly vernal species, apparently specialised
to a rich calcareous soil mixed with remnants of Abies alba
in altimontane areas. H. retrocurvata fruits in autumn and
winter in sites with a Mediterranean influence (Van Vooren
et al. 2017).
Pyronemataceae Corda, [as ’Pyronemaceae’], Anleit. Stud.
Mykol., Prag: 149 (1842)
Notes: This family produces hypogeous to epigeous ascomata with a variety of shapes, sessile to stalked, discoid
to cupulate, turbinate to pulvinate, folded to solid. (Perry
et al. 2007; Hansen et al. 2013; Pfister 2015a). In addition,
members of this family are distinguished by inamyloid asci
and smooth to ornamented ascospores (Pfister 2015a). More
than 65 genera are included within this family, with around
50% of genera having available molecular data (Kajevska
et al. 2013; Pfister 2015a; Ekanayaka et al. 2018). Recent
phylogenies on the family were performed by Hansen et al.
(2013) and Ekanayaka et al. (2018).
Neottiella (Cooke) Sacc., Syll. Fung. (Abellini) 8: 190
(1889)
Notes: Neottiella was introduced in 1889 and typified
by Neottiella albocincta (Berk. & M.A. Curtis) Sacc. This
genus is characterized by colored ascomata, inamyloid asci
and ornamented ellipsoid ascospores with warty or reticulate surfaces (Akata and Kaya 2013; Beug et al. 2014; Yao
and Spoone 1996). The type of ornamentation of ascospores
is an important character to identify species, especially the
warty ascospores and reticulate ascospores (Kullman 2002).
Fungal Diversity (2020) 104:1–266
Saprobic on soil. Sexual morph Apothecia epigeous,
scattered to gregarious, stipitate. Disc 0.2–0.3 cm high,
0.2–0.6 cm broad, discal to cupulate, orange, hymenium
orange, receptacle surface yellow to orange, margin conspicuous, dentate to wavy slightly. Stipe 0.2–0.4 cm long,
0.1–0.2 cm broad, cylindrical, yellowish to white. Medullary excipulum 50–200 µm broad, of textura intricate, yellow, composed of 3–7 µm broad hyphae. Ectal excipulum
60–100 µm broad, of textura angularis to globulosa, yellow, composed of 28–37 × 18–27 μm cells, with long septate hyphold hairs, 6–8 µm broad. Stipitipellis 70–160 µm
broad, of textura angularis to globulosa, yellow, comprised
of 24–35 × 16–24 μm cells, with long septate mycelial hairs,
5–8 µm broad. Paraphyses 3–4 µm broad, filiform, aseptate,
straight, or flexuous slightly at the apex, with yellow content,
apex enlarged, 4–7 µm broad, J−. Asci 359–411 × 17–22 µm,
8-spored, subcylindrical to clavate, operculate, inamyloid.
Ascospores [20/1/1, in H2O] (27.1–)27.7–31.1(–32.8) × (17
.0–)17.9–19.7(–20.5) µm (Q = 1.46–1.78, Q = 1.57 ± 0.10),
broadly ellipsoid, uniseriate, ornamented with verrucose,
uniguttulate to biguttulate. Asexual morph unknown.
Material examined: CHINA, Sichuan Province, G318,
on the ground with mosses, alt. 4215 m, 17 August 2018;
Ming Zeng, ZM38 (HKAS 104669, holotype); ibid. (MFLU
19-0523, isotype).
GenBank numbers: LSU: MK589293; TEF1-α:
MK577716.
Notes: This species is distinguished by the orange
stipitate ascomata, yellow paraphyses and broadly ellipsoid ascospores with a warty surface. The phylogenetic
tree shows a close relationship with Neottiella albocincta
(Berk. & M.A. Curtis) Sacc. (Fig. 47). Across the LSU
sequences, there are 33 notable nucleotides including
4 gap-differences among the 845 analyzed nucleotides
between N. gigaspora (HKAS 104669) and N. albocincta
(PRM 945,796). N. albocincta differs from N. gigaspora in the size of the asci (290–310 × 15–17.5 μm) and
ascospores (18.8–23.9 × 12.1–14.5 μm) (De Marchi
2017). N. vivida (Nyl.) Dennis has a paler outer surface,
smaller asci (140–175 × 17–18.5 μm) and ascospores
(22.46 ± 2.13 × 13.14 ± 1.23 μm) (Stoykov et al. 2015). N.
rutilans (Fr.) Dennis has ellipsoid to fusiform spores with
reticulate ornamentation (Kullman 2002; Akata and Kaya
2013).
Sordariomycetes
Neottiella gigaspora M. Zeng, Q. Zhao & K.D. Hyde, sp.
nov.
Index Fungorum number: IF556258; Facesoffungi number: FoF 05830; Fig. 46
Etymology: The epithet refers to the big ascospores.
Holotype: HKAS 104669.
13
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Notes: See Hongsanan et al. (2017) and Daranagama et al.
(2018) for details.
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor &
M.E. Barr, Sydowia 50 (1): 23 (1998)
Fungal Diversity (2020) 104:1–266
Fig. 46 Neottiella gigaspora (HKAS 104669, holotype). a–h
Typic mature specimens. i Vetical section of ascomata. j Stipitipellis. k Receptacle surface of pileus. l Receptacle surface of pileus in
Congo Red. m Asci and paraphyses in Melzer’s reagent. n–p Asci in
71
Congo Red (n Asci in Melzer’s reagent. o–p Asci in Congo Red). q–v
Ascospores (t Asci in Melzer’s reagent. u–v. Ascospores in Congo
Red). Scale bars: e–g = 1000 μm, h-i = 500 μm, j–p = 100 μm, q–
v = 20 μm
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Fig. 47 Maximum likelihood tree based on a combined LSU and TEF1 sequence dataset. MLBP and MPBP ≥ 70% are indicated above the
nodes. PP ≥ 0.95 are indicated above the nodes. Names in blue indicate new species. Otidea alutacea was used as outgroup taxon
Notes: Apiosporaceae was erected by Hyde et al. (1998)
to accommodate Apiospora (sexual morph of Arthrinium)
and Appendicospora. Based on molecular and morphological data, Crous and Groenewald (2013) synonymized Apiospora under Arthrinium Kunze. According to the Outline
13
of Ascomycota (Wijayawardene et al. 2018), six genera are
accepted in this family (Appendicospora, Arthrinium, Dictyoarthrinium, Endocalyx, Scyphospora, and Spegazzinia).
Members of Apiosporaceae are found as saprobes and
Fungal Diversity (2020) 104:1–266
73
Fig. 48 Arthrinium sorghi (URM 93000, holotype). a Colony on PDA after 7 d at 25 ± 1 °C. b Conidiophore and conidial bunch. c–e Conidial
bunch, conidiophores, and conidiogenous cells giving rise to conidia. f Conidia. Scale bars: b = 25 µm, c–f = 10 µm
pathogens of leaves, stems and roots, as endophytes, and
occasionally infecting humans (Senanayake et al. 2015).
Arthrinium Kunze, in Kunze & Schmidt, Mykol. Hefte 1:
9 (1817)
Notes: The genus Arthrinium was erected by Kunze
(1817) and is distributed in several ecological niches (Crous
and Groenewald 2013). Members of Arthrinium are associated with plants as endophytes, pathogens or saprobes, especially on monocotyledonous plants belonging to the families
Poaceae and Cyperaceae (Senanayake et al. 2015; Réblová
et al. 2016; Wang et al. 2018). Several Arthrinium species
are known to produce bioactive compounds used in the pharmaceutical industry (Hong et al. 2015; Shrestha et al. 2015).
Arthrinium sorghi J.D.P. Bezerra, C.M Gonçalves & C.M.
Souza-Motta, sp. nov.
Index Fungorum number: IF555795; Facesoffungi number: FoF 05762; Fig. 48
Etymology: The name refers to the host plant, Sorghum
bicolor, from which it was isolated as an endophyte.
Holotype: URM 93000.
Endophyte from leaves of Sorghum bicolor. Sexual
morph: Undetermined. Asexual morph: Hyphae hyaline,
branched, septate, 2–4 μm diam. Conidiophores erect or
ascending, simple, straight or flexuous, 28–50 × 5 μm, or
reduced to conidiogenous cells. Conidiogenous cells aggregated in clusters on hyphae, hyaline, cylindrical to subcylindrical, 3–5 × 3–4 μm. Conidia brown to dark brown,
13
74
smooth, globose, subglobose, with a longitudinal germ slit,
6–8 × 6–10 μm.
Culture characteristics: On PDA, colonies are flat, cottony, circular margin, sparse, aerial mycelium, surface and
reverse white, reaching 90 mm after 7 d, conidia are produced after 30 d. On MEA, colonies are flat, cottony, aerial
mycelium abundant, white surface, reverse yellowish, reaching 90 mm after 7 d.
Material examined: BRAZIL, Pernambuco state, Goiana
municipality, isolated as endophyte from leaves of Sorghum
bicolor, 25 August 2014, R.M.F. Silva (URM 93000, holotype); ex-type living culture, URM 7417.
GenBank numbers: ITS: MK371706; β-TUB: MK348526.
Notes: The ITS rDNA sequence of Arthrinium sorghi
demonstrated 93% identity with sequences of A. pseudosinense (CPC 21546, GenBank: KF144910) and A. ovatum
(CBS 115042, GenBank: NR_121558), and 88% identity with a sequence of A. phaeospermum (CBS 114314,
GenBank: KF144904). The β-TUB sequence also showed
87% identity with the sequence of A. phaeospermum (CBS
114314, GenBank: KF144996), 82% identity with the
sequence of A. obovatum (LC4940, GenBank: KY705166),
and 79% identity with the sequence of A. ovatum (CBS
115042, GenBank: KF144995). The β-TUB sequence was
not included in the phylogenetic analysis, but it is available
to verify its relationship to sequences from other Arthrinium
species. The phylogenetic analysis (Fig. 49) using ITS rDNA
sequences, placed A. sorghi as an unique lineage within the
genus Arthrinium. Morphologically, A. sorghi resembles A.
pseudosinense, A. ovatum and A. phaeospermum but differs from them by the culture charcteristics, conidiophores,
and conidia size, and by lacking a sexual morph which is
reported in A. pseudosinense (Crous and Groenewald 2013).
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández
Notes: Chaetosphaeriales was established by Huhndorf
et al. (2004) based on LSU phylogenetic analysis (Lu et al.
2016). Chaetosphaeriales consists of four families Chaetosphaeriaceae, Helminthosphaeriaceae, Leptosporellaceae
and Linocarpaceae with 49 genera (Wijayawardene et al.
2018).
Chaetosphaeriaceae Réblová, M.E. Barr & Samuels,
Sydowia 51(1): 56 (1999)
Notes: Chaetosphaeriaceae was introduced to accomodate Chaetosphaeria, Ascocodinaea, Melanochaela, Melanopsammella, Porosphaerella, Porosphaerellopsis and Striatosphaeria (Wei et al. 2018). The family includes 38 genera
(Wijayawardene et al. 2017a). Among them 32 hyphomycetous genera were recorded (Wijayawardene et al. 2017b).
Chloridium Link, Mag. Gesell. Naturf. Freunde, Berlin
3(1–2): 13 (1809)
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Fungal Diversity (2020) 104:1–266
Notes: The hyphomycetous genus Chloridium was typified with C. viride Link. A study by Hughes (1951) revealed
that Gonytrichum caesium, the generic type of Gonytrichum
is the asexual morph of Melanopsammella inaequalis. Furthermore, Réblová et al. (2016) noted that C. viride, the type
of Chloridium, is congeneric with M. inaequalis (Réblová
and Winka 2000; Fernández et al. 2006; Crous et al. 2012).
Hence, Réblová et al. (2016) synonymized Gonytrichum and
Melanopsammella under Chloridium. Chloridium species
are commonly found on decaying wood, soil and in aquatic
habitats (Lee and Go 2000; Wei et al. 2018).
Chloridium macrocladum (Sacc.) Karun., Maharachch.,
C.H. Kuo & K.D. Hyde, comb. nov.
≡ Gonytrichum macrocladum (Sacc.) S. Hughes, Trans.
Br. Mycol. Soc. 34(4): 565 (1952) [1951]
≡ Chloridium aseptatum M.J. Wei & H. Zhang, in Wei,
Zhang, Dong, Boonmee & Zhang, Phytotaxa 362(2): 191
(2018)
Index Fungorum number: IF557649; Facesoffungi number: FoF 06257; Fig. 50
Saprobic on culms of Pennisetum purpureum. Asexual
morph: Conidiophores crowded, solitary with single erect
main stalk, 275–290 × 5.5–7.5 μm ( x = 282.5 × 6.5 μm,
n = 10), septate, dark brown becoming light brown towards
the apex, swollen at the base and paler towards the apex; 3–4
collar-like hyphae appear underneath each node at the lower
half of the conidiophore giving rise to an irregularly curved,
succession of 2–4 phialides. Conidiogenous cells phialidic,
cylindrical to lageniform, producing conidia from multi conidiogenous loci, phialides borne around the conidiophore.
Conidia 3.6–4.3 × 1.8–2.8 μm ( x = 3.9 × 2.3 μm, n = 40),
ellipsoid to oval, subhyaline to olivaceous to light green,
slightly flattened at the end. Sexual morph: unknown.
Material examined: TAIWAN, Chia Yi Province, Kwang
Hwa, 18 March 2018, Anuruddha Karunarathna AKTW 58
(MFU 19-1003), living culture, NCYUCC 19-0367.
GenBank numbers: ITS: MK963072; LSU: MK963073.
Notes: Gonytrichum is a synonym of Melanopsammella
and later was synonymized with Chloridium, which Hyde
et al. (2020a, b) followed. In our phylogenetic analysis
(Fig. 51), the strains NCYUCC 19-0367 and CBS 875.68
show 0.98 similarity in the ITS region. Hence, we introduce
C. macrocladum NCYUCC 19-0367, a new host and geographical record. The strain NCYUCC 19-0367 is 99.8%
identical to C. aseptatum MFLU11-1051 (514/515 bp similarity). The solitary disagreement is due to a gap in C. aseptatum MFLU11-1051 occurring above the adenine–thymine
in the NCYUCC 19-0367 sequence at the 503rd position.
According to the morphology along with similarities in the
ITS sequence data, we propose C. aseptatum as a synonym
of C. macrocladum.
Fungal Diversity (2020) 104:1–266
75
Fig. 49 Bayesian inference tree based on ITS sequences representing Arthrinium. Bayesian posterior probabilities ≥ 0.93 are given near nodes.
The tree is rooted with Seiridium phylicae (CPC 19965). The new species is in blue. Hosts or substrates are indicated in green
Diaporthales Nannf.
Notes: Diaporthales is an important ascomycete order
in the subclass Diaporthomycetidae (Sodariomycetes) comprising phytopathogenic, endophytic and saprobic species
(Senanayake et al. 2018). In 1932 Nannfeldt erected Diaporthales to accommodate Höhnel’s Eu-Diaportheen group,
and since then it has undergone many taxonomic reconstructions (Wehmeyer 1975; Barr 1978; Maharachchikumbura
et al. 2015). Currently there are 14 families accepted in the
Diaporthales (Senanayake et al. 2018). With combined
taxonomic and phylogenetic results an updated classification for Diaporthales was done by Senanayake et al. (2017,
2018).
Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638 (1926)
Notes: Diaporthaceae is in Diaporthales, Sordariomycetes. This family consists of pathogenic, saprobic and
endophytic species which inhabit a wide diversity of host
substrates, including economically important trees and
crops. Species within the Diaporthaceae form solitary
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Fig. 50 Chloridium macrocladum on decaying stem of Pennisetum
purpureum culms. a Conidiophores on host. b Conidiophore. c Base
of the conidiophore. d Conidiogenus cells on conidiophore. e Coiled
terminal setae. f, g Conidiogenesis. h Conidia. i Conidia germination.
Scale bars: b–d = 50 µm, e–i = 10 µm
or aggregated, immersed to erumpent, rarely superficial,
orange, brown to black perithecial ascomata, with short or
long necks, located in stromatic tissues or substrates, with a
centrum (or hamathecium) lacking or with few paraphyses.
The asci are unitunicate with a conspicuous refractive ring
(Hawksworth et al. 1995; Rossman et al. 2007; Senanayake
et al. 2018). The ascospore morphology is diverse, ranging
from short to elongate and aseptate or septate with hyaline or
pigmented walls. The asexual morphs of Diaporthaceae are
generally coelomycetous (Rossman et al. 2007; Senanayake
et al. 2018), producing acervuli or pycnidial conidiomata,
with or without a well-developed stroma. Conidiogenesis is
phialidic or rarely annellidic and conidia are usually unicellular or 1-septate (Rossman et al. 2007).
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77
Fig. 51 Maximum likelihood
tree based on a combined
dataset of LSU and ITS partial
sequences. Bootstrap support
values for maximum likelihood
(ML) ≥ 75% and Bayesian posterior probabilities (PP) ≥ 0.98
are displayed above the nodes
respectively. The tree is rooted
with Chaetosphaeria fuegiana
CBS 114553 and C. innumera
SMH 2748. Newly generated
sequence is indicated in red.
The ex-type isolates are in bold
Diaporthe Nitschke, Pyrenomyc. Germ. 2: 240 (1870)
Notes: The genus Diaporthe is typified by Diaporthe eres
Nitschke and its species are found worldwide on a diverse
range of host plants as endophytes, pathogens and saprobes
(Udayanga et al. 2015; Dissanayake et al. 2017). Species
of Diaporthe are distinguished mainly by their molecular
phylogenies, and the best five gene regions to conduct a
multi-gene phylogenetic analysis are ITS, TEF1, ACT CAL
and HIS (van Rensburg et al. 2006; Santos et al. 2010; Dissanayake et al. 2017).
Diaporthe pimpinellae Abeywickrama, Camporesi, Dissanayake & K.D. Hyde, sp. nov.
Index Fungoru mnumber: IF556363; Facesoffunginumber: FoF 05979; Figs. 52, 53
Etymology: The specific epithet pimpinellae is based on
the host genus (Pimpinella).
Holotype: MFLU 19-0563.
Saprobic on aerial stems of Pimpinella peregrina. Sexual
morph: Not observed. Asexual morph: Coelomycetous,
Conidiomata visible as black dots up to 150–200 μm high,
150–250 μm diam, superficial, solitary, scattered on host,
oval to round, black. Peridium 18–41 μm thick, inner layer
composed of light brown to black textura angularis, outer
layer composed of dark brown to black textura angularis.
Conidiophores cylindrical, aseptate, densely aggregated,
straight or sinuous, terminal, slightly tapered towards the
apex. Conidiogenous cells 4–8 μm long × 2–4 μm wide,
phialidic, cylindrical, terminal and lateral. Alpha conidia
with 1–2 guttules per cell, 6–8 × 2–3 μm, hyaline, fusiform
or oval. Beta conidia not observed (Fig. 52).
Culture characteristics: Colonies on PDA covering entire
Petri dishes after seven days at 25 °C, white, spreading with
sparse to no aerial mycelium; surface with irregular patches
of white; surface white; reverse dirty white; conidiomata
black and white to cream conidial masses; alpha-conidia
ellipsoid to round, with an obtuse apex, tapering to an obtuse
or bluntly rounded base, beta-conidia observed (Fig. 53).
Known distribution: on dead aerial stems of Pimpinella
peregrine (Apiaceae).
Material examined: ITALY, Rocca delle Caminate, Predappio, Forlì-Cesena Province, dead aerial stems of Pimpinella peregrine (Apiaceae), 11 April 2018, Erio Camporesi, MFLU 19-0563 (holotype); ex-type living culture JZB
320131.
GenBank numbers: ITS: MK874656; β-TUB: MT373072;
HIS: MT373073; TEF1: MT373074.
Notes: Diaporthe pimpinellae, collected from Pimpinella
peregrine in Italy, formed a sister clade to D. chamaeropis
(CBS 454.81) and D. cytosporella (FAU461) with phylogenetically good support (83% ML and 1.00 PP) (Fig. 54).
Hypocreales Lindau
Notes: The order Hypocreales includes ten families
Bionectriaceae, Clavicipitaceae, Cordycipitaceae, Flammocladiaceae, Hypocreaceae, Nectriaceae, Niessliaceae,
Ophiocordycipitaceae, Stachybotryaceae and Tilachlidiaceae, which are distributed in subtropical and tropical
regions (Maharachchikumbura et al. 2015). These families
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Fig. 52 Diaporthe pimpinellae
(MFLU 19-0563, holotype). a
Appearance of fruitbodies on
host substrate. b Close-up view
of fruitbodies. c Section through
the fruitbody. d Peridium cell
wall and immature and mature
conidia attached to conidiogenous cells. e, f, g Conidia
differ amongst each other according to their fleshy and colorful perithecial ascomata, typically ostiolate perithecia, very
varied ascospores and pigment production (Rehner and
Samuels 1995).
Clavicipitaceae Kirk, Cannon, Minter & Stalpers, Dictionary of the Fungi (2008)
Notes: The Clavicipitaceae, one of the most miscellaneous families in the Hypocreales, comprises about 48 genera (Maharachchikumbura et al. 2015) involving obligate
saprobes, symbionts and parasites of insects, grasses, fungi
or sedges (Kepler et al. 2013; Schardl et al. 2013; Li et al.
2016). Clavicipitaceae, along with Cordycipitaceae and
Ophiocordycipitaceae, were distinguished by Sung et al.
(2007) based on molecular phylogeny. Most reported clavicipitaceous genera are pathogenic to whiteflies (Aleyrodidae) and/or scale insects (Coccidae), are commonly found in
the subtropics and tropics with most species belonging to the
genera Moelleriella, Hypocrella and Samuelsia (Chaverri
et al. 2008).
Moelleriella Chaverri, Liu & Hodge, Studies in Mycology
60: 66 (2008)
Notes: The genus Moelleriella was established by Bresadola (1896), with M. phyllogena as the type species. Chaverri
et al. (2008) differentiated Moelleriella from both Samuelsia and Hypocrella based on its disarticulated ascospores
in the asci while the latter two genera have filiform or
13
long-fusiform ascospores even after discharge. Species of
Moelleriella produce an Aschersonia-like asexual morph
that is characterized with acervuli or pycnidia on a brightly
colored stroma, filiform multi-septate ascospores disarticulated at the septa within the ascus and fusoid conidia
(Mongkolsamrit et al. 2014). In China, there are currently
only three species accepted in Moelleriella (Qiu et al. 2009;
Li et al. 2016).
Moelleriella gracilispora Jun Z. Qiu & Y.X. Chen, sp. nov.
Index Fungorum number: IF556162; Facesoffungi number: FoF 00010; Fig. 55
Etymology: Named after the slender spores.
Holotype: HMAS 247788.
Parasitic on whitefly nymph (Hemiptera), forming whitish to pale brown stromata on the underside of leaves. Sexual
morph: Undetermined. Asexual morph: Coelomycetous.
Stromata flattened pulvinate to discoid up to 3.2 mm diam
and 0.6 mm high with a whitish tomentum, no reaction to
3% KOH, upper surface roughened, opaque, without hypothallus. Conidiomata pycnidial, yellow to orange, solitary or
gregarious, completely embedded and on top part of stroma,
numerous pycnidia per stroma, flask-shaped, multi-loculate,
with locules, 166 (83–236) × 93 (53–136) μm, ostiolate.
Ostioles subcylindrical, single, well-developed, centrally
located. Wall of conidiomata 18–31 μm wide, composed of
relatively thick-walled, pale brown to hyaline cells. Paraphyses 66 (36–103) × 0.6(0.4–0.8) μm, filiform, tapering at
Fungal Diversity (2020) 104:1–266
79
Fig. 53 Asexual morph of Diaporthe pimpinellae (MFLU 19-0563, holotype) (from culture). a Upper view of colony on PDA. b Reverse view
of colony on PDA. c, d Pycnida forming on culture. e Spore mass. f Conidia attached to the coidiogenous cell. g Beta conida. h Alpha conidia
the apices, arising from the hymenium of the conidioma.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 6–12 × 0.8–1.3 μm, hyaline, phialidic, cylindrical, broad at base, discrete, thick- and smooth-walled.
Conidia fusoid, sometimes narrowly fusiform, unicellular,
straight or slightly curved, 11.8(11–12) × 1.4(1–2) μm, l/w
8.4(7–10), accumulating in a slimy pale brownish mass
above the conidioma.
Culture characteristics: Conidia germinating within 24
h on PDA. Colonies on PDA at 25 °C; compact, whitish,
attaining ca. 1 cm diam after 2 wks. Optimal temperature
20–26 °C, with no growth below 5 °C or above 35 °C.
Stromatic colonies pale yellow, forming moderately compact stromata. Conidial masses yellowish orange to orange,
abundant, appearing on surface of stromatic colonies after
ca. 2 wk.
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80
Fig. 54 Phylogenetic tree generated by maximum likelihood analysis of combined ITS, TEF1, β-TUB, CAL and HIS sequence data of
Diaporthe species. Tree was rooted with Diaporthella corylina (CBS
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121124). Bootstrap support values (ML) ≥ 60% and Bayesian posterior probabilities values (PP) ≥ 0.9 are shown near the nodes respectively. The newly generated sequence is indicated in blue
Fungal Diversity (2020) 104:1–266
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Fig. 54 (continued)
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Fig. 54 (continued)
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Fig. 55 Moelleriella gracilispora (CGMCC3.18989, holotype). a
Habitat. b Stroma on the underside of leaves. c Flank of stroma. d,
e Frontage of stroma. f Colonies on PDA media. g, h Longitudinal
83
section of stroma. i Section of pycnidium. j Conidiogenous cells. k
Conidia. Scale bars: b = 1 cm, c–e = 0.2 mm, f = 0.5 cm, g, h = 100
µm, i = 20 µm, j, k = 10 µm
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Fig. 56 Phylogenetic position of Moelleriella gracilispora
(CGMCC3.18989, holotype) inferred from a combined dataset of
LSU, RPB1 and TEF-1α sequences. Bootstrap support values for ML
and MP ≥ 50% are given near nodes respectively. The tree is rooted
with Balansia henningsiana (AEG96-27a) and Epichloe elymi (C.
Schardl 760). The new species is indicated in red
Material examined: CHINA, Fujian Province, Wuyishan City, Wu Yi Mountain, alt. 1000 m, on infected
whitefly nymphs (Hemiptera), Junzhi Qiu, 30 July 2017,
WYFC2017-03, WYFC2017-03 (HMAS 247788, holotype); HMAS 247,789, ex-type living cultures, CGMCC
3.18989, CGMCC 3.18990.
GenBank numbers: LSU: KC964202; TEF-1α:
KC964191; RPB1: KC964179.
Notes: We were unable to find a teleomorph of this species in our survey of entomopathogenic fungal diversity
throughout the year. Phylogenetic analysis based on LSU,
RPB1 and TEF-1α sequence data showed that Moelleriella
gracilispora is phylogenetically distinct from M. zhongdongii, M. mollii and M. ochracea (Fig. 56). M. gracilispora
morphologically resembles M. libera, M. evansii and M.
zhongdongii in having pulvinate stroma, a yellow mass of
extruded spores and fusiform conidia. However, the conidia
and paraphyses of M. gracilispora are somewhat narrower
(11.2–12.3 × 1.3–1.6 μm and 0.6 μm respectively). In contrast, the anamorph of M. libera (= Aschersonia aleyrodis)
produces conidia measuring 10–16 × 1.5–2 μm, with paraphyses 1–1.5 μm wide; M. evansii has conidia measuring
10–16 × 1.5–2 μm, and is without paraphyses; the anamorph
of M. zhongdongii (= A. incrassata) has conidia measuring 10–18 × 1.5–2 μm, with paraphyses sometimes present.
The Chinese specimens were most similar to Aschersonia
andropogonis (= M. ochracea) in having pulvinate stroma,
yellowish orange to orange spore masses and fusiform
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conidia. However, M. gracilispora differs from A. andropogonis in the absence of a hypothallus and size of conidia. M.
gracilispora produces longer and narrower conidia measuring 11.2–12.3 × 1.3–1.6 μm in comparison with 8–14 × 1.5–2
μm in A. andropogonis.
Hypocreaceae De Not., G. Bot. Ital 2(1): 48 (1844)
Notes: Hypocreaceae was erected by De Not (1844), with
Hypocrea as the type genus, and divided into six subfamilies by Lindau (1897). Seaver (1909a, b, 1910a, b, 1911)
divided Hypocreales into two families (Nectriaceae and
Hypocreaceae) depending on their perithecial and stromatic
characters and this concept was accepted by Kreisel in1969,
along with another five families in the Hypocreales based
on their bright-colored, fleshy ascomata, presence of apical paraphyses and production of phialides on free conidiophores. Rossman et al. (1999) agreed and included 12 genera
in Hypocreaceae. Hypocreaceae contains 13 teleomorph and
seven anamorph genera (Jaklitsch et al. 2014). Maharachchikumbura et al. (2015) accepted 18 genera in the family.
Trichoderma Pers., Neues Mag. Bot. 1: 92 (1794)
Notes: The genus Trichoderma was established in 1794
with four species (Samuels 1996). The holomorph typified
generic name Trichoderma is preferred over the sexual name
Hypocrea (Rossman et al. 2013; Bissett et al. 2015). Over
320 species have been described worldwide. The first record
of Trichoderma in China was from rotten wood in Sichuan
Province (Patouillard 1895). More recently, 43 new Trichoderma species were reported from China in a large-scale
survey of Trichoderma (Chen and Zhuang 2017a, b, c; Qin
and Zhuang 2017; Zhu et al. 2017a, b).
Trichoderma ceratophylletum Z.F. Yu & X. Du, sp. nov.
Index Fungorum number: IF555895; Facesoffungi number: FoF 05674; Fig. 57
Etymology: Refers to type strain isolated from Ceratophyllum demersum.
Holotype: YMF1.04621.
Saprobic on submerged stem, Sexual morph: Undetermined. Hyphae radiate and branched, dense and abundant on
PDA and SNA, relatively sparse on CMD, pustules formed
frequently on SNA and CMD, compact, abundant, asymmetrical to hemispherical, usually 0.5–1.5 mm, aggregated
to ca. 3 mm diam. Conidiophores tree-like, comprising a
straight main axis, often terminating in 3–5 phialides and
paired branches, the primary branches arising at right angle
from the main axis, terminating in verticils of 3–5 divergent
phialides, occasionally unilateral, the distance between two
neighbor branches is 9.5–32.3 μm (n = 15). The bases of
branches are usually thick, about 2.8–6.1 μm wide (n = 30).
Phialides stout, ampulliform to globose, crowded arrangement, (3.7–)4.1–8.4(–9.7) × 2.3–4.1 μm (n = 40), length/
85
width ratio (1.0–)1.2–2.8(–3.2). Conidia ovoid, sometimes ellipsoid, smooth-walled, at the base usually slightly
narrower, greenish in mass, 2.5–3.9 × 1.9–2.9 (–3.2) μm
(n = 30), length/width ratio 1.0–1.7.
Culture characteristics: On CMD after 72 h colony radius
56–62 mm, mycelium covers the plate after 96 h at 25 °C.
Colonies circular, translucent, margin distinct, aerial hyphae
sparse and arachnoid, pustules formed after 4 days. On PDA
and SNA, mycelium covers the plate after 72 h at 25 °C.
Colonies on PDA distinctly zonate, the zone around central
part of colony forms an irregular green ring, aerial hyphae
abundant, compact, cottony to downy. Colonies on SNA are
similar to PDA, but the aerial hyphae is relatively scarcer
than on PDA, pustules formed after one day, turning green
after 2 days, distribute on outer layer.
Material examined: CHINA, Guizhou Province, Qing
Zhen, Hong Feng Lake, within stem submerged in lake, 20
June 2014, YMF1.04621 (holotype).
GenBank numbers: ITS: MK327581; RPB2: MK327580;
TEF1: MK327579.
Notes: Trichoderma ceratophylletum phylogenetically
forms a single branch (Fig. 58) and is related to T. pleuroti
and T. pleuroticola. T. pleuroti differs from T. ceratophylletum by the characteristics of the branches, which are usually Gliocladium-like, with the phialides on the branches
often in crowded whorls and not divergent, whereas the
branches of T. ceratophylletum are tree-like, and the phialides are commonly divergent in whorls of 3–5. The conidia
of T. pleuroti are narrower than those of T. ceratophylletum
(2.8–4.2 × 1.6–2.2 vs 2.5–3.9 × 1.9–2.9 μm, Yu et al. 2006).
The conidiophores of T. pleuroticola tend to be regularly
verticillate, which distinguishs it from the obviously paired
branches and tree-like conidiophores of T. ceratophylletum.
The phialides of T. pleuroticola are sometimes lageniform
and longer (5.5–11.0 vs 4.1–8.4 μm) than those of T. ceratophylletum, which are commonly ampulliform to globose.
The conidia of T. pleuroticola are often larger than those of
T. ceratophylletum (2.9–4.5 × 2.4–3.5 vs 2.5–3.9 × 1.9–2.9
μm, Yu et al. 2006).
Ophiocordycipitaceae G.H. Sung, J.M. Sung, Hywel-Jones
& Spatafora, in Sung et al., Stud. Mycol. 57: 35 (2007)
Notes: Ophiocordycipitaceae was established by Sung
et al. (2007) with Ophiocordyceps as the type genus. Sung
et al. (2007) listed two teleomorphic genera (Elaphocordyceps and Ophiocordyceps) and nine anamorphic genera (Haptocillium, Harposporium, Hirsutella, Hymenostilbe,
Paecilomyces-like, Paraisaria, Syngliocladium, Tolypocladium and Verticillium-like) under Ophiocordycipitaceae.
Quandt et al. (2014) accepted six genera (Ophiocordyceps,
Tolypocladium, Purpureocillium, Harposporium, Drechmeria and Polycephalomyces) in Ophiocordycipitaceae
based on the phylogenetic analyses of a concatenated
13
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Fig. 57 Cultures and anamorph of Trichoderma ceratophylletum (YMF1.04621, holotype). a on CMD at 25 °C, 3 days. b, c Cultures (b on
PDA, 3 days. c on SNA, 3 days) at 25 °C. d–g Conidiophores and phialides (SNA, 3 d). h, I Conidia (PDA, 20 d). Scale bars: d–i 10 μm
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87
Fig. 58 Phylogenetic tree based
on Bayesian analysis of the
combined TEF1, RPB2, ITS
sequences. Nectria berolinensis
was used as outgroup in blue.
Bayesian posterior probabilities ≥ 0.90 are shown at the
nodes. The scale bar shows the
expected changes per site. The
new species are in boldface
five-gene dataset (SSU, LSU, TEF1, RPB1 and RPB2)
Maharachchikumbura et al. (2016). and Wijayawardene
et al. (2018) followed Quandt’s treatment and the former
author reviewed Ophiocordycipitaceae providing an updated
generic descriptions for those accepted genera. Hyde et al.
(2020a, b) expanded Ophiocordycipitaceae to include four
additional genera Hymenostilbe, Hirsutella, Paraisaria and
Perennicordyceps. Currently, Ophiocordycipitaceae comprises ten genera (Hyde et al. 2020a, b). New species combinations have been made for Polycephalomyces (Kepler
et al. 2013), Tolypocladium (Quandt et al. 2014), Drechmeria, Harposporium, Ophiocordyceps and Purpureocillium
(Spatafora et al. 2015). Morphologically, Ophiocordycipitaceae species are characterized by producing darkly pigmented stromata that are pliant to wiry, or fibrous to tough
in texture, rarely fleshy, often with aperithecial apices or
lateral pads, perithecia that are superficial to immersed, ordinal or oblique in arrangement, asci that are cylindrical with
thickened ascus at apex and ascospores that are cylindrical,
multiseptate, disarticulating into part-spores or non-disarticulating. Species of Ophiocordycipitaceae are pathogens of
arthropods, Elaphomyces, protozoans, rotifers, nematodes,
humans and animals (Sung et al. 2007; Maharachchikumbura et al. 2016).
Hirsutella Pat., Revue Mycol., Toulouse 14(no. 54): 67
(1892)
Notes: Hirsutella was erected by Patouillard (1892) and
typified with Hirsutella entomophila. This genus is characterized by the production of synnematous or mononematous
conidiophores, basally inflated phialides that narrow into
one or more slender needle-like necks and globose, oval,
fusiform, lemon-like to acicular hyaline conidia that are confined in a mucus droplet (Liang 1990; Gams and Zare 2003;
Sung et al. 2007; Qu et al. 2018; Hyde et al. 2020a, b). It is
worth mentioning that the phialidic conidial morphology
of Hirsutella species was mainly divided into five types,
correlating with phylogenetic information (Qu et al. 2018).
The concatenated TEF1, ITS and LSU sequences allows
for deeper understanding of the interspecific relationships
in Hirsutella (Qu et al. 2017, 2018). The sexual morph of
Hirsutella species has been linked to Ophiocordyceps (Sung
et al. 2007; Spatafora et al. 2015).
Hirsutella hongheensis D.P. Wei & K.D. Hyde, sp. nov.
13
88
Index Fungorum number IF556251; Facesoffungi number: FoF 05827; Fig. 59
Etymology: The specific epithet refers to the name of collection site.
Holotype: HKAS102451.
Parasitic on an unknown insect. Asexual morph: Mycelial subiculum bright yellow in superficial layer, becoming
white towards inner layer, cottony, dense, completely covering the host. Mycelium 1.5–2.9 µm ( x = 2.2 µm, n = 50)
wide, hyaline, thin-walled, smooth-walled, septate. The
hyphal cells usually become thick-walled and swollen, then
developing into chlamydospores. Hirsutella phialides-like
cells 7.6–14.3 × 3.1–4.6 µm ( x = 9.7 × 3.9 µm, n = 10) forming from inflated hyphae, subglose, ellipsoid, broadly cyclindrical, hyaline, slightly guttulate, bearing a needle-like neck
(2.1–3.9 × 0.4–1.1, x = 3 × 0.7 µm, n = 10). Chlamydospores
4.2–12.7 × 3.1–17.6 µm ( x = 7.7 × 4.8 µm, n = 100), subglobose, thick-walled, smooth-walled, hyaline, with yellow guttules, typically adhering in chains.
Material examined: CHINA, Yunnan Province,
Honghe County, Jiayin Villige, Amushan protection
area, On unknown insect, 23 October 2018, Deping Wei,
(HKAS102511, holotype).
GenBank numbers: ITS: MN017176; LSU: MN017175;
SSU: MN017177; TEF1: MN733824.
Notes: Attempts were made to cultivate the inner mycelium of the subiculum on PDA medium but growth was not
observed. Therefore, the morphological observation and the
sequencing of ITS, LSU, SSU and TEF1 gene regions were
conducted based on the dried specimen. Phylogenetically,
our collection has close affinity with Hirsutella versicolor
(ARSEF 1037) and is restricted to the H. guyana clade with
significant support (100% ML, 100% MP and 100% PP,
Fig. 60). H. versicolor was initially described from a leafhopper host collected in Nuwara, Sri-Lanka (Petch 1932).
The lectotype of H. versicolor was designated by Minter
& Brady (1980), with a hand-drawing of the phialides and
conidia attached. This reference species has a greyish to
orange-yellow byssoid mycelial subiculum radiating on
the leaf-hopper host, conoid or flask-shaped phialides that
narrow into 2–3 slender necks and narrow-cymbiform or
narrow-oval hyaline conidia (Petch 1932). Not one of these
characters was observed in our collections.
Glomerellales Chadef. ex Réblová et al.
Notes: Chadefaud (1960) proposed Glomerellales without a Latin diagnosis. This order was validly published
by Réblová et al. (2011) and comprises Australiascaceae,
Glomerellaceae, Malaysiascaceae, Plectosphaerellaceae
and Reticulascaceae (Maharachchikumbura et al. 2016;
Tibpromma et al. 2018).
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Glomerellaceae Locq. ex Seifert & W. Gams, in Zhang
et al., Mycologia 98(6): 1083 (2007) [2006]
Notes: Glomerellaceae was invalidly published by Locquin (1984) and was validated in Zhang et al. (2006). Glomerellaceae is a monotypic family.
Colletotrichum Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze
Deutschl.) 3(12): 41 (1831)
Notes: This genus was established by Corda (1831), to
accommodate C. lineola Corda (Hyde et al. 2009). The
genus Colletotrichum comprises mainly pathogens, as well
as endophytes and saprobes (Cannon et al. 2012; Hyde et al.
2014; Jayawardena et al. 2016).
Colletotrichum parthenocissicola Jayawardena, Bulgakov,
Huanraleuk & K.D. Hyde, sp. nov.
Index Fungorum number: IF556430; Facesoffungi number: FoF 05834; Fig. 61
Etymology: The specific epithet parthenocissicola is
named after the host genus Parthenocissus from which the
taxon was collected.
Holotype: MFLU 16-1557.
Saprobic or weakly pathogenic on dying and dead twigs
and petioles of Parthenocissus quinquefolia (L.) Planch
(Vitaceae). Sexual morph: Undetermined. Asexual morph:
Conidiomata 263–620 μm ( x = 437 μm, n = 10) diam, black,
acervulus, oval, solitary, gregarious. Setae straight or ± bent,
abundant, brown, becoming paler towards the apex, opaque,
smooth-walled, septate, 2–4 septate, 58–172 μm long, base
cylindrical, 6.5–8.0 μm diam, tip rounded. Conidiophores
simple, to 20 μm long, hyaline to pale brown, smoothwalled. Conidiogenous cells 8–14 × 2–5 μm ( x = 9.3 × 2.5
μm, n = 20), hyaline, smooth-walled, cylindrical to slighty
inflated, opening 0.5–1 μm diam, collarette or periclinal thickening rarely observed. Conidia 18–24 × 2–5 μm
( x = 20.3 × 3.8 μm, n = 40), L/W = 5.3, hyaline, smooth or
verruculose, aseptate, curved, both sides gradually tapering
towards the round to slightly acute apex and truncate base.
Appressoria not observed.
Culture characteristics: Colonies on PDA flat with entire
margin, aerial mycelium sparse and short, pale olivaceous
grey, colony surface buff, some sectors dark grey-olivaceous
to dark olivaceous grey, iron grey acervuli can be observed
mainly on the edge of the colony. Reverse olivaceous green,
concentric rings can be clearly observed, reaching 65–70
mm in 7 d at 28 °C.
Material examined: RUSSIA, Rostov region, Shakhty
City, private garden, dying and dead twigs and leafstalks
of Parthenocissus quinquefolia (L.) Planch. (Vitaceae), 5
March 2016, Timur S. Bulgakov (T-1263, MFLU 16-1557,
holotype), ex-type living culture MFLUCC 17-1098.
Fungal Diversity (2020) 104:1–266
Fig. 59 Hirsutella hongheensis (AMS12, holotype). a, b.Yellow
mycelial subiculum associated with host. c Enlargement of superficial mycelium. d–h Chlamydospores forming from hyphae indicated
89
with black arrows. i Chlamydospores adhering in chain. k–m Phialides-like cells with short necks. n–p Chlamydospores. Scale bars:
b = 1000 µm, c = 500 µm, d, e = 20 µm, f–k = 15 µm, l-p = 5 µm
13
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Fungal Diversity (2020) 104:1–266
Fig. 60 ML phylogenetic tree based on a combined TEF1, ITS and LSU dataset. The tree is rooted to Cordyceps gunnii (ARSEF 6828). Bootstrap support values (ML) ≥ 50% are indicated near the nodes. The generated isolate is in bold
GenBank numbers: ITS: MK629452; GAPDH:
MK639362; CHS: MK639356; ACT: MK639358; β-TUB:
MK639360.
Notes: The Colletotrichum dematium species complex is
mainly characterized by species with curved conidia (Damm
et al. 2009; Jayawardena et al. 2016). C. parthenocissicola
falls within the C. dematium species complex clade and
13
forms a separate branch close to C. insertae (Fig. 62). C.
quinquefoliae and C. insertae were found on Parthenocissus species in Russia (Li et al. 2015; Hyde et al. 2016). C.
parthenocissicola differs from C. insertae in having long
setae and conidia (C. insertae L/W = 5.0). The new taxon
differs from C. quinquefoliae by having short setae and
longer conidia (C. quinquefoliae L/W = 3.0). The cultural
Fungal Diversity (2020) 104:1–266
91
Fig. 61 Colletotrichum parthenocissicola (MFLU 16-1557,
holotype). a Conidiomata on
host. b Setae. c Conidiogenous
cells. d Conidia e Upper view
of the 7 d old culture on PDA. f
Reverse view of the 7 d old culture on PDA. Scale bars: b = 50
µm, c–d = 5 µm
characteristics of these three species cannot be compared as
both C. quinquefoliae and C. inserate lack cultures and were
described based on sequences. A BLASTn search of NCBI
GenBank with the ITS sequence of MFLUCC 17-1098
showed 99–100% similarity to several Colletotrichum species with curved conidia. The closest match in a BLASTn
search in GenBank with the GAPDH sequence of MFLUCC
17-1098 showed 99–100% similarity with the species of the
C. dematium complex. The CHS sequence showed 99% similarity to C. sonchicola (1 bp difference); the ACT sequence
showed a 99% similarity to C. menispermi (2 bp difference),
a 98% similarity to C. dematium (3 bp difference) and to
C. lineola (3 bp differences) and the β-TUB sequences also
showed a 99% similarity to C. dematium (1 bp difference).
The newly described taxon differs from its sister taxon C.
insertae in 4 bp in ITS, 10 bp in GAPDH, 8 bp in CHS, 11
13
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Fungal Diversity (2020) 104:1–266
Fig. 62 One of the 100 most parsimonious trees obtained from a heuristic search of the combined ITS, GAPDH, CHS, ACT and β-TUB
sequence data of Colletotrichum dematium species complex. The tree
is rooted with C. nigrum (CBS 169.49). Bootstrap support values
(MP and ML) ≥ 50% and Bayesian posterior probabilities ≥ 0.70 are
indicated above the nodes. The ex-type strains are in bold. New species in this study is in blue
bp in ACT and 47 bp in β-TUB. C. parthenocissicola differs
from C. quinquefoliae in 2 bp in ITS, 73 bp in CHS and 11
bp in ACT and 36 bp in β-TUB. Therefore, a new species is
described to accommodate this taxon. Morphological characters as well as phylogenetic evidence support this taxon to
be a new species of Colletotrichum.
Junewangiaceae J.W. Xia & X.G. Zhang, Scientific Reports
7: 7888 (2017)
Notes: Junewangiaceae was established by Xia et al.
(2017) to accommodate the asexual genus Junewangia
which includes five Acrodictys-like species (Baker et al.
2002; Xia et al. 2017; Song et al. 2018a). Dictyosporella
and Sporidesmiella were later accepted in the family based
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Fungal Diversity (2020) 104:1–266
on morphological characters and molecular analyses (Luo
et al. 2019).
Dictyosporella Abdel-Aziz, Fungal Diversity 75: 144 (2015)
Notes: Ariyawansa et al. (2015) erected genus Dictyosporella with D. aquatica Abdel-Aziz as the type species.
Three species (D. aquatica, D. hydei H.Y. Song & D.M.
Hu and D. thailandensis W. Dong, H. Zhang & K.D. Hyde)
are accepted in the genus, all reported from freshwater
habitats. Dictyosporella is a holomorphic genus with the
asexual morphs known of D. aquatica and D. hydei and the
sexual morph of D. thailandensis. The genus is characterized by immersed brown ascomata with the neck erumpent
through the host surface, unitunicate, cylindrical asci with
a J- apical ring and uniseriate, fusiform, hyaline, 3-septate ascospores with appendages at both ends (Zhang et al.
2017); macronematous or reduced conidiophores, monoblastic, globose or cylindrical conidiogenous cells and globose
to broadly cylindrical and muriform conidia (Ariyawansa
et al. 2015; Song et al. 2018b). Junewangia resembles Dictyosporella in the conidial morphology but differs in having
macronematous, brown conidiophores with percurrent elongation (Baker et al. 2002; Xia et al. 2017). Sporidesmiella
is distinguished from Junewangia and Dictyosporella by its
transversely septate conidia and polyblastic conidiogenous
cells (Kirk 1982; Luo et al. 2019).
Dictyosporella guizhouensis J. Yang & K.D. Hyde, sp. nov.
Index Fungorum number: IF556314; Facesoffungi number: FoF 05999; Fig. 63
Etymology: Referring to the collecting location in
Guizhou Province, China.
Holotype: MFLU 18-1505.
Saprobic on decaying twigs from freshwater habitats.
Asexual morph: Colonies on natural substrate sporodochial, scattered, dark brown or black. Mycelium partly
immersed, partly superficial, composed of septate, hyaline
hyphae. Conidiophores macronematous, mononematous,
compact, flexuous, simple or branched, mostly moniliform,
with globose to subglobose, ellipsoid or clavate cells, hyaline, smooth-walled, up to 52 μm long. Conidiogenous cells
monoblastic, integrated, terminal, globose, subglobose,
ellipsoid or clavate, hyaline, smooth-walled, 9–17 × 6.5–12
μm ( x = 11.5 × 9.5 μm, n = 25). Conidia acrogenous, globose, subglobose, ellipsoid or irregular, muriform, with
slightly irregular multi-euseptate, smooth, olivaceous to
mid-brown, paler at the basal cell, 15–26 × 12.5–18.5 μm
( x = 19 × 16 μm, n = 50). Sexual morph: Undetermined.
Material examined: CHINA, Guizhou Province, Anshun City, Gaodang village, 26°4.267′ N, 105°41.883′ E, on
decaying wood submerged in the Suoluo river, 19 October
2016, J. Yang, GD 34–2 (MFLU 18-1505, holotype; HKAS
102160, isotype).
93
GenBank numbers: ITS: MK593606; LSU: MK593605;
SSU: MK593611.
Notes: Dictyosporella, Junewangia and Sporidesmiella
nested within Junewangiaceae in the phylogenetic tree based
on a combined LSU and ITS sequence dataset (Fig. 64).
However, Junewangia is shown to be polyphyletic which
agrees with the result of Luo et al. (2019). D. guizhouensis
clustered as a sister taxon to D. aquatica and D. thailandensis with full support (100% ML and 1.00 PP). D. hydei
grouped with J. aquatica H.Y. Song & D.M. Hu, with full
support (100% ML and 1.00 PP) and only one nucleotide
difference, but only the LSU sequence data is available for
D. hydei. Relationships between the genera and taxa in Junewangiaceae are awaiting to be resolved.
Morphological characters of the new taxon matched
the generic concept of Dictyosporella. Septa in D. guizhouensis and D. hydei are slightly constricted while those in
D. aquatica are strongly constricted. The conidial color is
brown to black in D. aquatica, yellowish brown in D. hydei
and olivaceous green in D. guizhouensis. In addition, D.
guizhouensis has conspicuous hyaline globose to subglobose
conidiogenous cells which are not observed in D. aquatica
and D. hydei.
Rhamphoriaceae Réblová, Mycologia 11: 754 (2018)
Notes: Rhamphoriaceae was erected by Réblová and
Štěpánek (2018) and characterized by species having perithecial ascomata with a cylindrical or rostrate neck, the
absence of stromatic tissue or clypeus, similar anatomy of
two-layered ascomatal walls, cylindrical paraphyses, unitunicate asci with a distinct, non-amyloid apical annulus,
and dictyoseptate or transversely septate, hyaline or brown
ascospores. Five genera, Linkosia, Rhamphoria, Rhamphoriopsis, Rhodoveronaea and Xylolentia, are accepted in this
family.
Xylolentia Réblová, Mycologia 11: 759 (2018)
Notes: The genus Xylolentia was established by Réblová
and Štěpánek (2018) to accommodate X. brunneola Réblová.
It is characterized by brown, 1-septate ascospores, conidiogenous cells that are polyblastic with sympodially extending
rachis, and conidia that are hyaline becoming brown, and
aseptate (Réblová and Štěpánek 2018).
Xylolentia reniformis C.G. Lin, K.D. Hyde & Jian K. Liu,
sp. nov.
Index Fungorum number: IF556253; Facesoffungi number: FoF 05995; Fig. 65
Etymology: In reference to the reniform conidia.
Holotype: MFLU 19-0210.
Saprobic on decaying wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse on the natural
substrate, scattered, hairy, dark brown. Mycelium partly
13
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Fig. 63 Dictyosporella guizhouensis (MFLU 18-1505, holotype). a Colony on substrate. b Conidiophores. c–f Conidiogenous cells with
conidia. g–i Conidia. j Germinated conidium on PDA. Scale bars: b–e, j = 20 μm, f–i = 10 μm
superficial and partly immersed in the substratum. Conidiophores macronematous, mononematous, scattered, erect,
simple, straight or slightly flexuous, smooth, thick-walled,
13
septate, cylindrical, dark brown at the base, paler towards the
apex, 65–160 µm long, 3–9.5 µm thick at the base, slightly
tapering towards the conidiogenous cells just below the
Fungal Diversity (2020) 104:1–266
95
Fig. 64 Maximum likelihood tree of Diaporthomycetidae isolates
based on a combined LSU and ITS sequence dataset. Bootstrap support values (ML) ≥ 75% and Bayesian posterior probabilities ≥ 0.95
are indicated above the nodes. Branches with 100% ML BP and 1.0
PP are shown as bold branches. The tree is rooted with Leotia lubrica
(AFTOL-ID 1) and Microglossum rufum (AFTOL-ID 1292). The
new species is in blue
apex then swelling again. Conidiogenous cells polyblastic,
integrated, sympodial, terminal, clavate, subhyaline to pale
brown, 14–40 µm long, 3–4.5 µm thick in the broadest part.
Conidia aggregated in slimy masses, acropleurogenous, simple, smooth, aseptate, reniform, hyaline, 3.2–4.6 µm long,
1.8–3.0 µm thick in the broadest part.
Culture characteristics: Conidia germinating on WA
(water agar) within 48 h. Colonies on PDA circular, appearing waxy-mucoid, white-brown, reaching a diam of 7.5–9
mm in 30 days at 28 °C.
Material examined: CHINA, Guizhou Province, Qiannan Buyi Miao Autonomous Prefecture, Dushan County,
Guizhou Zilinshan National Forest Park (Shengou district),
on decaying wood, 6 July 2018, C.G. Lin, DS 1–37 (MFLU
19-0210, holotype; HKAS 105134, isotype), ex-type living
cultures GZCC 1-–0048.
GenBank numbers: ITS: MK547646; LSU: MK547648.
Notes: The phylogenetic result based on a combined LSU
and ITS sequence dataset indicated that Xylolentia reniformis is closest to Linkosia multiseptum (HKUCC 10825)
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Fig. 65 Xylolentia reniformis (MFLU 19-0210, holotype). a Host material. b Conidiophores on the host surface. c, d Conidiophores, conidiogenous cells with conidia. e, f Conidiogenous cells. g, h Conidia. Scale bars: c, d = 20 μm, e, f = 10 μm, g, h = 5 μm
and X. brunneola (PRA 13,611) (Fig. 66). Shenoy et al.
(2016) showed Linkosia is polyphyletic and phylogenetically
located in three different lineages within the Sordariomycetes, viz., Chaetosphaeriales, Hypocreales and Sordariomycetes incertae sedis. L. multiseptum has brown and distoseptate conidia and conidiophores reduced to conidiogenous
cells. Réblová and Štěpánek (2018) showed the phylogenetic
placement of L. multiseptum in the Rhamphoriaceae was
unexpected based on its morphology. Morphologically, X.
13
reniformis fits well with the concept of Xylolentia (Réblová
and Štěpánek 2018). X. reniformis differs from L. multiseptum and X. brunneola by its conidia being aseptate, reniform
and hyaline and longer conidiophores. The conidia of X.
brunneola are ellipsoid to subglobose to obovoid, aseptate,
straight, hyaline becoming pale brown. Due to the unsolved
taxonomic placement of Linkosia, as well as the morphological assignment, we hereby identify our new collection
Fungal Diversity (2020) 104:1–266
97
Fig. 66 Phylogenetic tree generated from ML analysis based on combined LSU and ITS sequence data for Rhamphoriaceae. Bootstrap
support values (MP and ML) ≥ 50% and Bayesian posterior probabili-
ties ≥ 0.95 are indicated above or below the nodes. Ex-type strains are
in bold, the new isolate is in blue. The tree is rooted with Brachysporium nigrum (M.R. 1346)
as a species of Xylolentia even though it is phylogenetically
closer to L. multiseptum (Fig. 66).
chains, ellipsoid to cylindrical, septate, septum darkened,
constricted at the septum, truncate at both ends, subhyaline
to olive green when immature, brown to dark brown when
mature.
Notes: In the phylogenetic analysis, Catenuliconidia
uniseptata formed a distinct subclade and was basal to the
Barrmaeliaceae clade without significant support (Fig. 67).
Barrmaeliaceae comprises two genera, Barrmaelia Rappaz
and Entosordaria (Sacc.) Höhn. (Voglmayr et al. 2018).
Barrmaelia has a Libertella-like asexual morph which is
only known from pure cultures (Voglmayr et al. 2018), while
the asexual morph of Entosordaria is unknown. Based on
morphology and phylogeny, we treat Catenuliconidia as a
distinct new genus in Xylariales incertae sedis.
Xylariales Nannf.
Notes: The Xylariales is a large order introduced by
Nannfeldt (1932). Barr (1990) initially assigned Xylariales in the class Hymenoascomycetes. Later, Eriksson and
Winka (1997) established the subclass Xylariomycetidae
and included Xylariales in it. Recently, different numbers
of families in Xylariales were accepted by several authors
(Maharachchikumbura et al. 2015, 2016; Senanayake et al.
2015; Hongsanan et al. 2017; Wijayawardene et al. 2018).
Xylariales genus incertae sedis
Catenuliconidia N.G. Liu & K.D. Hyde, gen. nov.
Index Fungorum: IF556464; Facesoffungi number: FoF
06116
Etymology: “Catenu-” in reference to catenulate conidia.
Type species: Catenuliconidia uniseptata N.G. Liu &
K.D. Hyde
Saprobic on decaying wood in terrestrial habitat. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies on natural substrate effuse, black, velvety. Mycelium mostly immersed, composed of septate, branched,
hyaline hyphae. Conidiophores macronematous, mononematous, in groups, erect, straight or broadly curved, brown
to dark brown, septate, unbranched, smooth, thick-walled.
Conidiogenous cells blastic, terminal, integrated, cylindrical, brown. Conidia catenate, developing in acropetal
Catenuliconidia uniseptata N.G. Liu & K.D. Hyde, sp. nov.
Index Fungorum number: IF556465; Facesoffungi number:
FoF 06117; Fig. 68
Etymology: Named after its 1-septate conidium.
Holotype: MFLU 19-0689.
Saprobic on decaying wood. Colonies on natural substrate
effuse, black, velvety. Mycelium mostly immersed, composed of septate, branched, hyaline hyphae. Conidiophores
macronematous, mononematous, in groups, erect, straight
or broadly curved, cylindrical, brown to dark brown, septate, unbranched, smooth, thick-walled. Conidiogenous cells
blastic, terminal, integrated, cylindrical, brown. Conidia
17.5–24.5 × 6.0–9.5 μm ( x = 21.4 × 8.0 μm, n = 30), catenate, developing in acropetal chains, ellipsoid to cylindrical,
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Fig. 67 Maximum likelihood tree based on analysis of a combined
LSU, ITS and RPB2 sequence dataset representing Xylariales. Bootstrap support values for ML ≥ 75% and Bayesian posterior probabilities ≥ 0.95 are given near nodes respectively. The tree is rooted with
Pseudoneurospora amorphoporcata (CBS 626.80) and Sordaria
fimicola (CBS 508.50). Ex-type or ex-epitype strains are indicated in
bold. The new taxon is indicated in blue
1-septate, septum darkened, constricted at the septum, truncate at both ends, subhyaline to olive green when immature,
brown to dark brown when mature.
Culture characteristics: Conidia germinating on water
agar within 24 h and producing germ tubes from one or
both ends. Colonies growing slowly on PDA medium, surface velvety, with entire edge, white, center fairly dense and
protuberant, middle ring flat, outer ring sparse from above;
center yellow, color decreasing from the center to periphery
from below.
Material examined: CHINA, Guizhou Province, Guiyang,
Guizhou Academy of Agricultural Sciences, on decaying
wood, 27 May 2018, N.G. Liu, NKY025 (MFLU 19-0689,
holotype); ex-type living culture, GZCC 20-0036.
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99
Fig. 67 (continued)
GenBank numbers: ITS: MK804515; LSU: MK804516;
SSU: MK804519; TEF1: MK828509; RPB2: MK828513.
Notes: Phylogenetically, Corynesporopsis belongs to
Xylariaceae, while Catenuliconidia is treated as Xylariales
genus incertae sedis. In Xylariales, Catenuliconidia uniseptata resembles Corynesporopsis species in having catenate,
obclavate to ellipsoid conidia. However, C. uniseptata has
shorter conidiophores than those of Corynesporopsis species. In addition, the conidia of C. uniseptata are 1-septate,
while those of Corynesporopsis spp. are multi-septate (Xia
et al. 2013; Kirschner 2015).
Basidiomycota
Agaricomycetes
Agaricales Underw.
Notes: The order Agaricales was introduced by Underwood (1899) and mainly includes gilled fungi along with
13
100
Fig. 68 Catenuliconidia uniseptata (MFLU 19-0689, holotype). a, b
Colonies on wood. c. Sporodochium d Conidiophore and conidium.
e–f Conidial chains. g–k Conidia. l Germinating conidium. m, n Col-
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onies on PDA medium. Scale bars: c = 30 µm, d–f = 20 µm, g–k = 5
µm, l = 10 µm
Fungal Diversity (2020) 104:1–266
some non-gilled relatives. It is represented by more than
13,000 species belonging to 413 genera of 33 families (Kirk
et al. 2008). Several mycologists developed different classification schemes for the grouping of these mushrooms
(Fries 1821; Fayod 1889; Kühner 1980). These traditional
classification systems were based on morphological, anatomical or cytological characters. The most significant work
was done by Singer (1986) in which he proposed a classification system by combining both morphological and anatomical features. This grouping was supported by molecular data (Hibbett et al. 1997; Moncalvo et al. 2000, 2002).
Recently, molecular phylogenetic studies based on multigene
DNA sequences generated six major clades of Agaricales
(Matheny et al. 2006).
Agaricaceae Chevall., Fl. Gén. Env. Paris (Paris) 1: 121
(1826)
Notes: Agaricaceae was erected by Chevallier (1826)
based on the type genus Agaricus L. Previously, this family
contained only gilled fungi. But research based on molecular data transferred many non-gilled fungal families such as
Lycoperdaceae, Nidulariaceae and Tulostomataceae to Agaricaceae. Now, it is represented by more than 1300 species
belonging to 85 genera (Kirk et al. 2008). Some gasteroid
puffball genera included in the Agaricaceae are Arachnion
Schwein., Bovista Pers., Calvatia Fr. and Lycoperdon Pers.
Lycoperdon Pers., Ann. Bot. (Usteri) 1: 4 (1794)
Notes: The genus Lycoperdon was first established by
Persoon in (1796a) who described L. perlatum as the type
species. Species are characterized primarily by subglobose
to pyriforme basidomata with a conspicuous sterile base and
dehiscence by an apical pore. The genus which has been
studied extensively by many mycologists (Hollòs 1904; Perdeck 1950; Ahmad 1952; Kreisel 1962; Demoulin 1971)
is represented by 52 species worldwide (Kirk et al. 2008;
Jeppson et al. 2012). Molecular phylogenetic studies have
widened the concept of the genus by including species of
the genera Bovistella, Handkea, Morganella, and Vascellum
(Krüger et al. 2001; Bates 2004; Larsson and Jeppson 2008;
Bates et al. 2009; Gube 2009; Jeppson et al. 2012). Larsson
and Jeppson (2008) established new limits of Lycoperdon by
proposing subgenera within Lycoperdon, viz., Apioperdon,
Bovistella, Lycoperdon, Morganella, Utraria and Vascellum.
Subgenus Vascellum corresponds to genus Vascellum
which was proposed by Smarda (Pilát 1958). He described
V. depressum (Bonord.) F. Šmarda (= V. pratense Pers.) as
the type species and segregated it from genus Lycoperdon.
Members of this group are characterized by the presence
of a diaphragm between the gleba and subgleba, reduced
or absent eucapillitium, abundant paracapillitium and asperulate basidiospores (Kreisel 1962, 1993; Smith 1974). In
101
this paper, it is treated here as subgenus Vascellum of genus
Lycoperdon following Larsson and Jeppson (2008). According to published literature and Index Fungorum (2020), the
total number of species belonging to Vascellum is 19 which
are distributed worldwide (Ponce de Leon 1970; Homrich
and Wright 1988; Kreisel 1993; Kreisel and Hausknecht
2001; Rocabado et al. 2007). The habitats of the species
range from pastures and meadows of lower altitude to grasslands of higher elevations. A few species of this group are
also involved in forming fairy rings in turf grass (Miller et al.
2011). Two new Lycoperdon species belonging to subgenus
Vascellum i.e. Lycoperdon lahorense and L. pseudocurtisii
are described, illustrated and presented here as new species. A key to the species of Lycoperdon subgenus Vascellum is provided (Smith 1974; Homrich and Wright 1988;
Kreisel and Hausknecht 2001; Bates et al. 2009; Cortez
et al. 2013). The phylogenetic tree of Lycoperdon spp. is
also given (Fig. 69).
Lycoperdon lahorense Yousaf & Khalid, sp. nov.
Index Fungorum number: IF556119; Facesoffungi number: FoF 05752; Figs. 70, 71, 72
Etymology: Named after the type locality ‘Lahore’.
Holotype: LAH100000148.
Basidiocarps globose to somewhat pyriforme, small to
medium, 8–20 mm diam × 9–25 mm in height, off white to
pale yellow above when young, pale brown to greyish brown
with age, off-white to light yellow at the base; attached to
the substratum by white, branched rhizomorphs, encrusted
with particles of soil; dehiscence by an apical pore up to 4
mm diam. Peridium double. Exoperidium pale yellowish to
greyish brown with age, not persistent, fragile, peeling off in
the form of patches, patches with pale brown to dark brown
warts, sub-flocculate, sometimes entire exoperidium in the
form of minute, prominent warts, these more above, and
lesser below, encrusted with debris. Endoperidium papery,
greyish brown to grey towards apical portion when mature,
off-white to pale yellow downwards, entirely exposed in
mature specimens. Gleba olivaceous to brown, cottony. Diaphragm distinct. Subgleba pale yellow, up to 10 mm high,
comprising only 1/3rd of the basidiocarps. Basidiospores
asperulate, globose, olivaceous, 3.5–4.3 μm diam, with a
stump of a pedicel. Eucapillitium absent. Paracapillitium
abundant, hyaline to olivaceous, septate, bulging at some
points, cytoplasmic contents abundant in hyphae, encrusted
with some amorphous material, 2.3–7.5 μm diam, thickwalled (walls up to 1 μm), with attenuate tips (up to 1 μm),
branched (dichotomous branching absent). Exoperidium
composed of hyaline to olivaceous, globose to subglobose,
thin-walled hyphal sphaerocysts. Endoperidium composed
of tightly interwoven, olivaceous to hyaline, thick-walled,
aseptate, up to 5 µm diam, branched hyphae.
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Fig. 69 Maximum likelihood tree based on ITS sequences of Lycoperdon shows its subgenera. Bootstrap support values ≥ 70% are given.
Sequences generated during this study are indicated with boxes
Material examined: PAKISTAN, Punjab, Lahore, University of the Punjab, New Campus, Botanical Garden, 5 July
2011, in groups, on ground, among grass, at 217 m (712ft.)
a.s.l., N. Yousaf, VPK4 (LAH100000148, holotype).
Additional material examined: PAKISTAN, Punjab,
Lahore, University of the Punjab, New Campus, Botanical Garden, 2 September 2012, in groups, on ground,
among grass, at 217 m (712 ft.) a.s.l., N. Yousaf, VPK6,
LAH100000149.
13
GenBank numbers: ITS: KC844917, KC844918,
MK414506.
Notes: Lycoperdon lahorense (VPK4, VPK6) is characterized by the sub-flocculate to slightly warted exoperidium, bicolored endoperidium with well-defined apical pore, distinct
diaphragm, asperulate basidiospores, absence of eucapillitial
threads and abundant paracapillitium in the gleba. Phylogenetically, L. lahorense is closely related to L. curtisii (= Vascellum curtisii). Both taxa share the characters of apical pore
Fungal Diversity (2020) 104:1–266
103
Fig. 70 Lycoperdon lahorense. a Young basidiocarps with floccose exoperidium (VPK4, holotype). b Mature basidiocarps after exoperidium
has sloughed off (VPK4, holotype). c Basidiocarps (VPK6). Scale bars: a = 5 mm, b, c = 6.5 mm
dehiscence of basidiocarps and the presence of asperulate
globose basidiospores. However, these species have morphological differences. The exoperidium in both species is
fragile and deciduous but different in morphology. In L. curtisii, when young, it is composed of prominent, convergent
fibrils in contrast to the powdery to warted appearance in L.
lahorense. The bicolored endoperidium is a unique feature
in L. lahorense. In L. curtisii the gleba contains reduced
eucapillitial threads along with abundant paracapillitium,
but true capillitium is absent in L. lahorense. The diaphragm
is another feature that differs between the species. It is very
distinct in L. lahorense and lacking in L. curtisii. Morphologically, the absence of eucapillitium, a distinct diaphragm
and globose to subglobose basidiospores make L. lahorense
similar to V. hyalinum and V. intermedium. However, the
color of the basidiocarps is reddish grey to dark brown when
young in V. hyalinum compared to off-white to pale yellow in L. lahorense. The apical pore is well-defined in our
species but it is irregular in V. hyalinum. Basidiospores are
always globose with a stump of a pedicel in L. lahorense,
whereas they are mainy subglobose with a few broadly ovoid
ones having a pedicel up to 1.5 µm in V. hyalinum. V. intermedium is another closely related taxon but the exoperidium
does not slough off in plates as in L. lahorense. In addition,
V. intermedium dehisces by an irregular-shaped ostiole in
contrast to the small apical pore of L. lahorense.
Lycoperdon pseudocurtisii Yousaf & Khalid, sp. nov.
Index Fungorum number: IF 556120; Facesoffungi number: FoF 05751; Figs. 73, 74.
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Fig. 71 Lycoperdon lahorense (VPK4, holotype). a Asperulate basidiospores. b Exoperidial elements. c Paracapillitial hyphae. d Aseptate
hyphae of subgleba. e Endoperidial hyphae. Scale bars: a = 2.8 µm, b = 10 µm, c = 14 µm, d, e = 10 µm
Etymology: Named after the resemblence with L. curtisii.
Holotype: LAH100000150.
Basidiocarps subglobose, 10–25 mm diam, × 25 mm
in height, off-white when young, turning grey with age;
attached to the substratum by a small mycelial pad, heavily
encrusted with particles of soil; dehiscence is by an apical
pore or slit, pore develops with sloughing of exoperidium,
up to 5 mm diam. Exoperidium off-white, granular, floccose
with rubbery texture, not persistent, sloughed off in mature
13
specimens, not in the form of patches or plates. Endoperidium papery, greyish brown. Gleba olivaceous, cottony.
Diaphragm distinct. Subgleba present, comprising 1/3rd of
basidiocarps. Basidiospores asperulate, globose, olivaceous,
2.8–3.8 µm diam. Eucapillitium absent. Paracapillitium
abundant, hyaline, septate, encrusted with some amorphous
material, 3.0–7.5 µm idiam, thin-walled, branched. Exoperidium composed of hyaline, subglobose to elongated,
thin-walled hyphal sphaerocysts. Endoperidium composed
Fungal Diversity (2020) 104:1–266
Fig. 72 Lycoperdon lahorense (VPK4, holotype). a Light micrograph of asperulate basidiospores. b Light micrograph of paracapillitial hyphae. Scale bars: a = 4 µm, b = 9 µm
of tightly interwoven, hyaline to light brown, thick-walled,
septae, up to 5 µm diam, branched hyphae.
Material examined: PAKISTAN, Punjab, Lahore, University of the Punjab, New Campus, 25 July, 2013, under
Grevillea robusta A.Cunn. ex R.Br., in groups, among grass,
at 217 m (712 ft.) a.s.l., N. Yousaf, VPK3 (LAH100000150,
holotype).
GenBank numbers: ITS: MK414503, MK414504,
MK414505.
Notes: Lycoperdon pseudocurtisii (VPK3) is characterized by off-white basidiocarps, up to 25 mm diam, distinct
diaphragm, asperulate, globose basidiospores (2.8–3.8
µm), absence of eucapillitium, abundant paracapillitium
and thick-walled endoperidial hyphae. Phylogenetically
L. pseudocurtisii is close to V. curtisii. Both of these taxa
have globose asperulate basidiospores but the presence of
eucapillitium and an indistinct diaphragm are features of
V. curtisii not found in L. pseudocurtisii. V. hyalinum and
V. intermedium are closely related taxa because they share
the characters of a distinct diaphragm, globose to subglobose asperulate basidiospores and the absence of a eucapillitium. The exoperidium peels away in patches or plates in
these taxa but this character is absent in L. pseudocurtisii.
Basidiospores in the latter are smaller compared to the larger
basidiospores of both taxa.
Key to Lycoperdon species of subgenus Vascellum
1 Eucapillitium present…………………………………… 2
1* Eucapillitium absent……………………………………6
2 Diaphragm distinct………………………………………3
2* Diaphragm indistinct……………………………………5
105
3 Paracapillitium > 5 µm….………………………………4
3* Paracapillitium 1–5 µm…………………V. pampeanum
4 Paracapillitium 3.5–7 µm; basidiospores globose, 3.4–4.2
µm, short pedicellate (< 1 µm); eucapillitium with rare small
pores, 3–5 µm……………………………………V. pratense
4* Paracapillitium up to 9 µm; basidiospores globose to
rarely subglobose, 4.0–4.3 × 4.3–4.7 µm; eucapillitium dextrinoid, cyanophilic, 1.8–3.6 µm ………………V. cuzcoense
5 Paracapillitium up to 6 µm; basidiospores globose, 3.2–4
µm…………………………………………………V. curtisii
5* Paracapillitium 2.2–6.0 µm; basidiospores subglobose,
4.0–5.6 × 4.0–4.8 µm…………………………V. llyodianum
6 Diaphragm less distinct to distinct………………………7
6* Diaphragm indistinct…………………………………12
7 Basidiospores subglobose to ellipsoid……………………8
7* Basidiospores globose or subglobose…………………9
8 Basidiospores subglobose to broadly ellipsoid, 3.0–4.3
µm…………………………………………V. endotephrum
8* Basidiospores ellipsoid to subglobose, 3.5–4.5 × 3.2–3.5
µm……………………………………………V. floridanum
9 Basidiospores globose…………………………………10
9* Basidiospores subglobose………………………………11
10 Basidiopores globose, 3.5–4.3 μm, with a stump of pedicel
………………………………………Lycoperdon lahorense
10* Basidiopores globose, 2.8–3.8 µm diam
……………………………………………L. pseudocurtisii
11 Basidiospores subglobose mostly, globose to ovoid sometimes, 3.8–4.5 × 3.5–3.8………………………V. hyalinum
11* Basidiospores globose to subglobose, 4.0–5.6 × (3.2–)
4.0–4.8 µm…………………………………V. intermedium
12 Basidiospores globose…………………………………13
12* Basidiospores subglobose……………………………14
13 Basidiospores 3.8–5.1 µm…………………V. delicatum
13* Basidiospores 4.0–5.0 µm……………………V. qudenii
14 Basidiospores 3.2–3.8 × 3.6–4.3 µm………V. cingulatum
14* Basidiospores subglobose to gobose, 4.0–4.8
(–5.6) × 4.0–4.8 (–5.6) µm………………………V. texense
Cortinariaceae R. Heim ex Pouzar, Česká Mykol. 37(3):
174 (1983)
Notes: For details of the family see Ariyawansa et al.
(2015), Li et al. (2016) and Hyde et al. (2016).
Cortinarius (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 627
(1821)
Notes: The ectomycorrhizal genus Cortinarius is the most
species-rich group of macrofungi with a worldwide distribution (e.g. Garnica et al. 2016). Recently, in the DNA era,
new possibilities have become available to discover new species from under-investigated parts of the world, combining
molecular, morphological and ecological data to help reveal
the true diversity of this large genus. In the course of our
studies of Cortinarius in north-western India, we discovered
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Fig. 73 Lycoperdon pseudocurtisii (VPK3, holotype). a Basidiocarps. b Asperulate basidiospores. c Paracapillitial hyphae. d Exoperidial elements. e Endoperidial hyphae. Scale bars: a = 5 mm, b = 2.5 µm, c, d, e = 10 µm
several new species-level lineages with the aid of phylogenetic analysis combined with morphological and ecological
observations. Two of them are here reported as new species,
one belongs to the sect. Percomes Melot and the other to the
sect. Rubicunduli Soop, B. Oertel & Dima (2019).
Sect. Percomes is a phlegmacioid lineage with characteristic morphology such as yellowish orange to greenish
yellow colours caused by anthraquinonoid pigments in the
basidiocarps, peculiar odours, and a red KOH reaction in
the context. They are as a rule associated with either coniferous or deciduous host trees. All described species are
known only from Europe or North America (Brandrud et al.
1990–2018; Liimatainen et al. 2014; Frøslev et al. 2017),
and the collection from India appeared to be a new species
13
based on ITS phylogeny and morphology/ecology (Figs. 75,
76). Furthermore, it is the first report of a member of this
section from Asia.
Cortinarius rubicundulus (Pers.) Fr. has been treated as
a species without close relatives within the genus Cortinarius for a long time, until C. subgemmeus Soop from New
Zealand was described (Soop 2003). Together they form an
isolated lineage in all phylogenies supported also by morphology, including the presence of remarkable cheilocystidia. This lineage was recently described as sect. Rubicunduli (Soop et al. 2019). The combined ITS-LSU phylogeny
(Fig. 77) shows that sect. Rubicunduli is a diverse group of
Cortinarius with at least twelve species worldwide including
this new species from India.
Fungal Diversity (2020) 104:1–266
Fig. 74 Lycoperdon pseudocurtisii (VPK3, holotype). a Light
micrograph of asperulate basidiospores. b Light Micrograph of paracapillitial hyphae. Scale bars: a = 6 µm, b = 8 µm
Cortinarius indorusseus Dima, Semwal, V.K. Bhatt &
Brandrud, sp. nov.
MycoBank number: MB829293; Facesoffungi number:
FoF 05731; Fig. 76
Etymology: Refers to the similarity to C. mussivus (= C.
russeus)–C. russeoides complex in sect. Percomes.
107
Holotype: KCS2470.
Pileus up to 110 mm diam, plano-convex to applanate,
with a faint umbo at centre, surface glabrous, glutinous, light
greenish yellow at margin, cinnamon brown to light orange
brown (6D6–5A4) towards centre, which oxidizes to reddish brown when mature; margin rimose with age, cuticle
half peeling. Lamellae adnate, crowded, up to 7 mm broad,
lamellulae of various lengths present, wax yellow when
young later becoming more brownish orange (5A5–5C5).
Stipe 120 × 18 mm, central, subcylindrical, slightly clavate
at base, widening at apex, pastel yellow to greenish yellow (3A4) above cortina, paler downwards, (6D5); basal
mycelium mainly whitish, mycelial strands at base yellow; universal veil forming a thin girdle/zone halfway up
the stipe, brownish to faintly vinaceous brown (5D8–5D6).
Context greenish yellow to yellowish, KOH reaction not
observed. Odour distinctly earth-like. Taste not observed.
Basidiospores: 9.5–10.5 × 5.5–6.5 µm, MV = 9.9 × 5.9 µm,
Q = 1.6–1.8, Qav = 1.67 (n = 40), amygdaloid, strongly verrucose. Spore print rusty brown. Pileipellis simplex, thick.
Habitat and distribution: Solitary, occurring in leaf litter
on humid soil, in temperate broadleaved forests dominated
mainly by Quercus leucotrichophora A. Camus, Myrica
esculenta Buch. Ham. ex. D. Don, and Rhododendron
arboreum Sm.
Material examined: INDIA, Uttarakhand, Pauri Garhwal,
Phedhkhal, 1950 m asl, 17 August 2015, K.C. Semwal, KCS
2470 (CAL, holotype).
GenBank numbers: ITS: MK372065.
Fig. 75 Phylogenetic position
of Cortinarius indorusseus
(KCS 2470, holotype) within
sect. Percomes inferred from
the ITS sequences. Bootstrap
support values (ML) ≥ 50% are
given near nodes. The tree is
rooted with members of sect.
Calochroi. The new species is
in blue
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Fig. 76 Basidiocarp and spores of Cortinarius indorusseus (KCS 2470, holotype). a–b Basidiocarp. c Spores. Scale bar: c = 10 µm
Notes: Cortinarius indorusseus is a typical member of
sect. Percomes characterized by the glutinous, (greenish)
yellow pileus surface that oxidizes to reddish brown with
age, especially at the centre, the slender, clavate stipe, the
yellowish context, the earth odour and the amygdaloid,
strongly verrucose spores. Phylogenetically, C. indorusseus
is very distinct from all member of sect. Percomes with a
difference of more than 70 substitution and indel positions
from the closest relative C. russeoides. Within the section,
C. indorusseus forms a subclade with the European C. russeoides and the North American C. superbus (Fig. 75). Macromorphologically it resembles very much C. russeoides and
C. mussivus (= C. russeus) associated with European conifers (Jeppesen et al. 2012). However, C. indorusseus differs
from these and all other macroscopically similar species in
sect. Percomes in possessing significantly smaller spores
(< 10 × 6 µm). Furthermore, the habitat is special; occurring in forests with the evergreen Quercus leucotrichophora
(Banjh oak) and Rhododendron arboretum in north-western
Himalaya, India. In the Mediterranean region, C. aurilicis
is frequent in a similar habitat with Quercus ilex. However,
this species differs in the more strongly red-brown to orange
pileus colour and the slightly marginate bulb, where the universal veil remnants remain attached at the base of the stipe.
This feature also distinguishes this species from those found
in other broad-leaved forests, such as C. nanceiensis associated with Fagus and Tilia in south-eastern Europe.
13
Cortinarius paurigarhwalensis Semwal, Dima & Soop, sp.
nov.
MycoBank number: MB829294; Facesoffungi number:
FoF 04972; Fig. 78
Etymology: The epithet refers to the type locality Pauri
Garhwal in India.
Holotype: KCS2487.
Pileus up to 125 mm diam, hemispherical then applanate,
surface dry, slightly tomentose, glabrous but centre with distinct radial wrinkles towards margin, red-brown to reddish
golden (6B7–6C7), more brownish to light brown (7D8) at
centre, margin undulate when mature. Lamellae adnate to
subdecurrent, crowded, up to 6 mm broad, lamellulae of
various lengths present, greyish brown to greyish orange
(6B5–5B5). Stipe 70 × 16 mm, cylindrical, slightly clavate to
bulbous, base up to 24 mm broad, greyish orange (6B5–6B6)
at apex, paler to whitish towards the base, but staining yellow. Context strongly canary yellow (2B7). Odour and
taste not observed. Basidiospores: 6.0–7.5 × 4.0–5.0 µm,
MV = 6.5 × 4.3 µm, Q = 1.4–1.6, Qav = 1.5 (n = 40), amygdaloid to broadly amygdaloid, very finely verrucose, almost
smooth. Cheilo- and pleurocystidia present, 20–40 × 3–7 µm
(n = 10), variable in shape, clavate, subclavate to ventricose,
sometimes subcapitate and lageniform. Spore print hazel
brown (6E8).
Habitat and distribution: Solitary, occurring in leaf litter
of Quercus leucotrichophora and Rhododendron arboreum.
Material examined: INDIA, Uttarakhand, Pauri Garhwal,
Phedhkhal, Ulkhagarhi, 2070 m asl, 30°9′36" N, 78°50′53"
Fungal Diversity (2020) 104:1–266
109
Fig. 77 Phylogenetic position of Cortinarius paurigarhwalensis (KCS 2487, holotype) within sect. Rubicunduli inferred from ITS and LSU
sequences. Bootstrap support values (ML) ≥ 50% are given near nodes. The tree is rooted with C. bolaris. The new species is in blue
E, 31 August 2015, K.C. Semwal, KCS 2487 (CAL,
holotype).
GenBank numbers: ITS: MK372066; LSU: MK372064.
Notes: Cortinarius paurigarhwalensis is the third
described species in sect. Rubicunduli besides the European type species C. rubicundulus (Rea) A. Pearson which
grows in boreal coniferous forests and C. subgemmeus Soop
associated with Nothofagus and Leptospermum in New Zealand. The closest relative of C. paurigarhwalensis based on
LSU sequences seems to be a species occurring in Malaysia (Borneo) (marked as Cortinarius sp5 in Fig. 77), but
the relationship gained no statistical support in our analysis. Using BLASTn of ITS sequences, the closest species
are Cortinarius sp3, and C. rubicundulus differing by > 40
substitution and indel positions from C. paurigarhwalensis.
Morphologically, as well as phylogenetically, C. paurigarhwalensis undoubtedly belongs to sect. Rubicunduli sharing
characters with the related species such as the orange-brown
basidiocarps, the yellowing context, the small, almost nonverrucose spores and presence of conspicuous lamellar cystidia, which is a rare feature in the genus Cortinarius.
Cortinarius sinensis L.H. Sun, T.Z. Wei & Y.J. Yao, sp. nov.
Fungal Name number: FN 570647; Facesoffungi number:
FoF 06101; Fig. 79a–b
Etymology: From the Latin for ‘Chinese’, referring to the
species collected from China.
Holotype: HMAS 99105.
Basidiocarps gregarious. Pileus 7–11 diam, up to 16 cm
diam, first hemispherical, then convex, finally convex-applanate to applanate, sometimes with depressed center; margin decurrent to straight; surface greyish brown to purplish
brown when young and sometimes with olive tint, reddish
brown with purplish tint (Brick Red to Garnet Brown) to
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Fig. 78 Basidiocarp and microscopic structures of Cortinarius paurigarhwalensis (KCS 2487, holotype). a–c Basidiocarp. d–e Spores and cystidia. Scale bar: c–e = 10 µm, d = 5 µm
purple brown (Deep Livid Brown) when mature, margin paling to greyish brown (Buckthorn Brown), yellowish brown
(Ochraceous-Tawny) to brown (Tawny to Sanford’s Brown),
mostly with dark purplish brown squamules or speckles
(Dark Purple Drab) from universal veil, fibrillose-striate,
smooth, shiny, viscid when moist. Lamellae adnate to emarginate-adnate, 5–17 mm wide; yellowish (Pale OchraceousSalmon) to brownish (Colonial Buff) when young, then
yellowish brown (Ochraceous-Tawny) to brown with rust
tint (Tawny), crowded, with lamellulae. Stipe 5–9 × 1.5–2.5
cm, central, cylindrical with a marginate-bulbous base up
to 3.5 cm across; surface pale brown (Light OchraceousBuff), yellowish brown (Yellow Ocher) to brown (Tawny)
and mostly whitish at upper part, brown (Tawny), reddish
13
brown (Brazil Brown) to purplish brown (Garnet Brown)
at base, rough, attached rust brown fibrils of universal veil;
solid to soft. Cortina almost whitish to purplish when young
and greyish purple when dry, finally rust brown with purple
tint. Universal veil covering whole of basidiocarp, purplish
when young, strongly purple when mature (especially in
exsiccata). Context up to 2.5 cm thick at center of pileus,
whitish, fleshy. Smell not distinctive. Taste mild. Spore print
rust brown. Chemical reaction with 20% KOH dark reddish brown. Fluorescence reaction under ultraviolet light
indistinct. Basidiospores (8.5–)9–12.5(–14) × (5.5–)6–7(–7.
5) μm, Q = 1.4–1.8 (x̅ = 1.6); subglobose, yellow–brown, distinctly verrucose. Basidia 26–32 × 6–8.5 μm, clavate, thinwalled, mostly subhyaline, with four sterigmata. Lamella
Fungal Diversity (2020) 104:1–266
111
Fig. 79 New Cortinarius species from China. a, b. C. sinensis
(HMAS 99105, holotype). a Basidiocarps. b Basidiospores. c, d C.
subsanguineus (HMAS 250503, holotype). c Basidiocarps. d Basid-
iospores. e, f. C. xiaojinensis (HMAS 274355, holotype). e Basidiocarps. f Basidiospores. Scale bar: a,c,e = 2 cm, b,d,f = 10 µm
margin heterogeneous, with sterile cells, 20–30 × 5–7 μm,
clavate, subhyaline, thin-walled. Cheilocystidia and pleurocystidia none. Pileipellis of brown hyphae, 2–6 μm diam,
thin walled. Clamp connections present in all tissues.
Material examined: CHINA, Qinghai Province, Qilian County, Zhamashi Town, on ground in coniferous forest with Picea sp., alt. 2950 m, 21 August 2004, Qing-Bin
Wang 444 (HMAS99105, holotype), Qing-Bin Wang 453
(HMAS99070), Qing-Bin Wang 454–1 (HMAS96980),
Qing-Bin Wang 454 (HMAS96985), Liang-Dong Guo
& Ying Zhang 624 (HMAS99446), Hua-An Wen 04,312
(HMAS138964), Hua-An Wen & Mao-Xin Zhou 4298
(HMAS144904); Babao Town, on ground in coniferous forest, 20 August 2004, Qing-Bin Wang 416 (HMAS96987),
Qing-Bin Wang 416-1 (HMAS96988); Datong County,
Dongxia Forest Station, alt. 3000 m, on ground in coniferous
forest, 17 August 2004, Qing-Bin Wang 356 (HMAS99090),
Qing-Bin Wang 356–1 (HMAS99102); Gansu Province,
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112
Tianzhu County, Huangyangchuan Town, Nanchong Forest Protection Station, on ground in coniferous forest with
Picea crassifolia, 37°15′38″ N, 103°3′58″ E, alt. 2900 m,
25 August 2016, Li-Hua Sun, Q61, HMAS279889, Li-Hua
Sun, Q62, HMAS279890, Li-Hua Sun, Q63, HMAS279891,
Li-Hua Sun, Q66, HMAS279893, Li-Hua Sun, Q78,
HMAS279900, Li-Hua Sun, Q80, HMAS279902, Li-Hua
Sun, Q83, HMAS279905, Li-Hua Sun, Q84, HMAS279906,
Li-Hua Sun, Q85, HMAS279907; Ningxia Autonomous
Region, Helanshan Mountains, August 2007, Hua-An Wen
(HMAS187061); The Inner Mongolia Autonomous Region,
Alxa Left Banner, Helan Mountains, August 2016, Li-Hua
Sun 89, HMAS279888.
GenBank numbers: ITS: MK411466, MK411462,
MK411473, MK411471, MK411472, MK411468,
MK411469, MK411464, MK411470, MK411467,
MK411463, MK411452, MK41145, MK411454,
MK411455, MK411456, MK411457, MK411458,
MK411459, MK411460, MK411465, MK411461.
Notes: Cortinarius sinensis is a species of subgenus
Phlegmacium (Fr.) Trog because of its visicid pileus and
distinguished from other species in the subgenus by its purplish pileus and universal veil. Phylogenetically, C. sinensis formed a clade with C. cupreorufus Brandrud and C.
pseudocupreorufus Niskanen, Liimat. & Ammirati (Fig. 80),
but the ITS sequences of C. sinensis deviate from that of
these two species in the phylogenic tree by at least 11 substitutions and indel positions. Furthermore, C. cupreorufus
(Breitenbach & Kränzlin 2000; Gutiérrez et al. 2006) and C.
pseudocupreorufus (Smith 1944; Liimatainen et al. 2014)
do not have purple colours. Morphologically, the species is
similar to C. rufoolivaceus (Pers.) Fr., which also has a lilac
to purplish-tinged pileus (Breitenbach and Kränzlin 2000;
Bidaud et al. 2004; Soop 2018), however, compared with
this species the new taxon has a much stronger purple coloration especially of the universal veil. C. sinensis is an edible
species and is consumed by people in the Helan Mountain
area, Ningxia Autonomous Region.
Cortinarius subsanguineus T.Z. Wei, M.L. Xie & Y.J. Yao,
sp. nov.
Fungal Name number: FN 570648; Facesoffungi number:
FoF 06102; Fig. 79c–d
Etymology: Referring to the morphological similarity to
C. sanguineus.
Holotype: HMAS 250503.
Basidiocarps gregarious. Pileus 2.5–6 cm diam, firstly
hemispherical, then convex, finally convex-applanate to
applanate, slightly umbonate at center; margin decurrent to
straight; surface dull orange-red (Vinaceous-Rufous), rust
red (Dragon’s-blood Red) to red (Scarlet) to with persistent
dark red (Ox-blood Red) minute floccose squamules from
universal veil, margin paling to orange-red (Coral Red to
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Orange Rufous). Lamellae adnate to emarginate-adnate, up
to 5 mm wide; red (Scarlet) when young, rust red (Dragon’s-blood Red) when mature, close, with lamellulae. Stipe
5.0–10.0 × 0.5–0.9 cm, central, cylindrical, base thickening
to 1.5 cm diam; surface reddish orange (orange Rufous)
when young, with rust tint when mature, with rust red floccules from universal veil; rough, solid to soft. Cortina reddish at first, finally rust red from the mature basidiospores.
Context up to 4 mm thick at center, pale red to dull red,
fleshy. Smell not distinctive. Taste mild. Spore print rust
brown. Chemical reaction with 20% KOH blackish red at
all parts. Fluorescence reaction under ultraviolet light indistinct. Basidiospores (6–)6.5–8(–9) × (4–)4.5–5.5(–5.7) μm,
Q = 1.4–1.6 (x̅ = 1.5); ellipsoid, yellow–brown to brown,
minutely to moderately verrucose. Basidia 26–31 × 6–8
μm, clavate, thin-walled, mostly subhyaline, with four sterigmata. Lamella margin heterogeneous, with sterile cells,
20–26 × 5–7 μm, clavate, subhyaline, thin-walled. Cheilocystidia and pleurocystidia none. Pileipellis of subcylindrical
hyaline, 5–15 μm diam, reddish brown to rust brown. Clamp
connections present.
Material examined: CHINA, Yunnan Province, Xianggelila County, Bitahai, alt. 3700 m, on ground in mixed forest with Picea sp. and Quercus aquifolioides, 12 August
2008, Tie-Zheng Wei, Xiao-Qing Zhang & Fu-Qiang Yu
134 (HMAS250503, holotype); Xizang Autonomous
Region, Bomi County, Gawarong Tample, 29°47′44.56″ N,
95°41′53.87″ E, alt. 3680 m, on ground in mixed forest with
Picea sp. and Quercus aquifolioides, 10 September 2014,
Tie-Zheng Wei, Jian-Yun Zhuang, Xiao-Yong Liu & Hao
Huang 4949 (HMAS253491); Milin County, Nanyigou,
29°02′11.59″ N, 94°14′29.26″ E, alt. 3166 m, on ground
in mixed forest with Picea sp. and Quercus aquifolioides,
20 September 2015, Tie-Zheng Wei & Bin-Bin Li 6542
(HMAS275034).
GenBank numbers: ITS: MK411450, MK411449,
MK411451.
Notes: Cortinarius subsanguineus is a member of section
Sanguinei Kühner & Romagn. ex M.M. Moser (Subgenus
Dermocybe) and mainly characterized by its orange-red
pileus and stipe. Phylogenetically, C. subsanguineus forms a
distinct terminal lineage and shows a close relationship with
a few red species, e.g. C. harrisonii Ammirati, Niskanen &
Liimat., C. neosanguineus Ammirati, Liimat. & Niskanen, C.
puniceus P.D. Orton, C. sanguineus (Wulfen) Gray, C. sierraensis (Ammirati) Ammirati, Niskanen & Liimat. and C.
vitiosus (M.M. Moser) Niskanen, Kytöv., Liimat. & S. Laine
(Fig. 81). However, the ITS sequences of C. subsanguineus
differ from the other species in the phylogenetic tree by at
least 7 substitutions and indel positions. C. neosanguineus
(Niskanen et al. 2013), C. puniceus (Bidaud et al. 1994; Niskanen et al. 2012; Grupo Ibero-insular de Cortinariologos
2014), C. sanguineus (Bidaud et al. 1994; Breitenbach and
Fungal Diversity (2020) 104:1–266
113
Fig. 80 Phylogenetic position of Cortinarius sinensis inferred from the ITS sequences. Bootstrap support values (ML and MP) ≥ 50% and
Bayesian posterior probabilities ≥ 0.5 are given near nodes respectively. The tree is rooted with C. rapaceoides (NR_130253 and KF732407)
Kränzlin 2000; Bellù et al. 2004; Niskanen et al. 2012,
2013; Soop 2018) and C. vitiosus (Niskanen et al. 2012) do
not have orange-tinged basidiocarps, and furthermore the
basidiocarps of C. neosanguineus (Niskanen et al. 2013) and
C. puniceus (Bidaud et al. 1994; Niskanen et al. 2012; Grupo
Ibero-insular de Cortinariologos 2014) are purplish-tinged.
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114
Fungal Diversity (2020) 104:1–266
Fig. 81 Phylogenetic position of Cortinarius subsanguineus and C.
xiaojinensis inferred from the ITS sequences. Bootstrap support values (ML and MP) ≥ 50% and Bayesian posterior probabilities ≥ 0.5
are given near nodes respectively. The tree is rooted with C. tabularis
(KX302268 and KX302275)
C. harrisonii has a deep orange-red to brownish orange
pileus and deep rusty orange stipe, but its basidiospores (4–5
μm wide, Niskanen et al. 2013) are narrower than those of
C. subsanguineus. The pileus of C. sierraensis is orange
to brownish orange, but lacks red color (Niskanen et al.
2013). Two other species, C. cinnabarinus Fr. and C. phoeniceus (Vent.) Maire, are also similar to C. subsanguineus
on account of their red basidiocarps. The basidiospores of
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the new Chinese taxon are 6–9 × 4–5.7 μm and shorter than
that those of C. cinnabarinus (8–10.5 × 4.5–5.5 μm, Bidaud
et al. 1994; 7.3–10 × 4.5–5.7 μm, Breitenbach and Kränzlin
2000; 7–10 × 4.5–5.5 μm, Bellù et al. 2004), and larger than
those of C. phoeniceus (5.5–8.5 × 4–4.5 μm, Bidaud et al.
1994; 6–7 × 3.5–4.5 μm, Soop 2018).
Cortinarius xiaojinensis T.Z. Wei, M.L. Xie & Y.J. Yao,
sp. nov.
Fungal Name number: FN 570649; Facesoffungi number:
FoF 06103; Fig. 79e–f
Etymology: Referring to the species collected from Xiaojin County, Sichuan Province, China.
Holotype: HMAS 274355.
Basidiocarps gregarious. Pileus 3.2–4.5 cm diam, firstly
hemispherical, then convex, finally convex-applanate to
applanate, sometimes slightly umbonate at center; margin
decurrent to straight; surface brown (Cinnamon-Brown),
dark brown (Chestnut) to blackish brown (Bay) with dark
brown minute flocci from universal veil, margin paling to
greyish brown with distinct olive tint (Dark Citrine), slightly
hygrophanous when moist. Lamellae adnate to emarginateadnate, up to 4 mm wide; pale brown to greyish brown with
slightly olive tint (Buffy Brown) when young, brown with
slightly rust tint (Mikado Brown to Hazel) when mature,
close, with lamellulae. Stipe 5.5–7 × 0.6–1.1 cm, central,
cylindrical, base thickening to 1.3 cm diam; surface pale
greyish brown (Olive-Buff) to pale olive brown (OliveOcher), base darkening to brown, with pale brown to rust
floccules from universal veil; rough, solid to soft. Cortina
greyish at first, finally rusty brown due to the mature basidiospores being trapped in it. Context up to 3 mm thick at
center, pale brown to dull brown, fleshy. Smell not distinctive. Taste mild. Spore print rust brown. Chemical reaction with 20% KOH blackish brown at pileus and lamellae,
brown at stipe and context. Fluorescence reaction under
ultraviolet light indistinct. Basidiospores (4.7–)5–6(–6.5
) × (3.8–)4–4.5(–4.7) μm, Q = 1.2–1.4 (x̅ = 1.3); ellipsoid,
yellow–brown to brown, minutely to moderately verrucose.
Basidia 25–32 × 6–9 μm, clavate, thin-walled, mostly subhyaline, with four sterigmata. Lamella margin heterogeneous,
with sterile cells, 20–26 × 6–7 μm, clavate, subhyaline, thinwalled. Cheilocystidia and pleurocystidia none. Pileipellis
of cylindrical hyphae, 5–10 μm diam, brownish to yellowish
brown. Clamp connections present.
Material examined: CHINA, Sichuan Province, Xiaojin
County, National Scenic Area of Four Girls Mountains,
30°59′32.26″ N, 102°50′52.47″ E, alt. 3468, on ground in
mixed forest with Abies fabri and Picea sp, 17 August 2015,
TTie-Zheng Wei, Di Wang 5602 (HMAS274355, holotype),
Tie-Zheng Wei, Di Wang 5685 (HMAS274425),.
GenBank numbers: ITS: MK411447, MK411448.
115
Notes: Cortinarius xiaojinensis is a species of subgenus
Dermocybe with olive brown basidiocarps. According to the
phylogenetic analysis based on ITS sequence (Fig. 81), the
clade of C. xiaojinensis (with high support) is sister to the
clade of C. malicorius, however, the ITS sequence of C.
xiaojinensis differs from that of C. malicorius by at least 9
substitutions and indel positions. In C. malicorius the olive
tint is weak and only found on the pileus surface (Bidaud
et al. 1994; Breitenbach and Kränzlin 2000; Grupo Iberoinsular de Cortinariologos 2009) and in the context (Bidaud
et al. 1994; Breitenbach and Kränzlin 2000; Bellù et al.
2005; Grupo Ibero-insular de Cortinariologos 2009; Soop
2018), whereas all parts of the basidiome of C. xiaojinensis
have this colour.
Hygrophoraceae Lotsy, Vortr. Bot. Stammesgesch. 1: 705
(1907)
Notes: Hygrophoraceae was erected by Lotsy in 1907.
Hygrophorus, the generic type for the family, was published
by Fries in 1836. Most species in Hygrophoraceae are characterized by thick basidiomes with distant, waxy lamellae,
spores mostly smooth, hyaline and inamyloid, and basidia
five or more times the length of their spores (Singer 1986).
However, these characters are not as reliable as they once
were (Lodge et al. 2014). Matheny et al. (2006) were the first
to show strong support for a monophyletic Hygrophoraceae.
Currently, Hygrophoraceae comprises over 600 species in
26 genera and is thus one of the larger families in the Agaricales (Lodge et al. 2014).
Humidicuteae Padamsee & Lodge, Fungal Diversity 64: 38
(2014)
Notes: The tribe Humidicuteae was introduced by Lodge
et al. (2014), based on the type genus Humidicutis (Singer)
Singer. It is characterized by basidiomes brightly colored
or greyish brown and without dopa-based pigments except
for a few species of Neohygrocybe. Clamp connections are
present at the base of the basidia and basidioles are often
toruloid. The lamellar context is regular or subregular but
not interwoven, the pileus context is inamyloid and some
species have strong odours. It includes the genera Humidicutis, Gliophorus, Gloioxanthomyces, Neohygrocybe and
Porpolomopsis (Lodge et al. 2014).
Humidicutis (Singer) Singer, Sydowia 12(1–6): 225 (1959)
[1958].
Notes: Pileus convex, convex-umbonate or conic, margin
rarely and not deeply splitting; surface subhygrophanous,
moist, rarely viscid, colors usually bright; lamellae thick,
sinuate or broadly adnate, often with a decurrent tooth; odor
absent or disagreeable; carotenoid pigments usually present, encrusting pigments may also be present on cuticular
hyphae, not soluble in alkaline solutions; pileipellis hyphae
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116
parallel, prostrate, cylindric; basidia usually 5 or more times
longer than the spore length; basidiospores hyaline, thinwalled, inamyloid, not metachromatic, ellipsoid or broadly
ellipsoid, not constricted; lamellar trama subregular or regular, of hyphae < 150 μm long, rarely tapered, with rightangled septa; clamp connections absent in context and pellis,
but toruloid clamps present at the base of the basidia and/or
basidioles (Lodge et al. 2014).
Humidicutis brunneovinacea R. Garibay-Orijel, sp. nov.
MycoBank number: MB829181; Facesoffungi number:
FoF 05673; Fig. 82
Etymology: Referring to the distinctive colors of the
basidiome.
Holotype: MEXU 28225.
Basidiocarp stipitate. Pileus conical, umbonate to campanulate, with age becoming broadly campanulate, planoumbonate or umbilicate with uplifted margin, 10–40(–57)
mm diam; lightly lubricous, moist (not viscid) but rapidly
drying, hygrophanous, glabrous, finely rugulose when
young, radially fibrillose with age; margin incurved and
folding when young, with age sometimes becoming crenulate, thin, discolored and often splitting some millimeters;
dark vinaceous brown, warm blackish brown or diamine
brown (10F6–8) when young, fading towards the margin, in
age some basidiocarps stay dark vinaceous, neutral red or
bordeaux (12D–E8) while most of them become rose red,
pomegranate purple or amaranth purple (12A–C8). Context
thin (0.5–1.5 mm), watery, concolorous. Lamellae sinuate
or adnate, some with a small decurrent tooth, thin, subdistant with 1–3 series of lamellulae, broad (7–10 mm) margin
undulate, white; in age becoming distant, thick, sometimes
transvenose to intervenose with red vinaceous margin; whitish to light greyish (10B1) becoming light brown-grey or
vinaceous grey (10C–D2) or vinaceous pink (12A–B4)
with age. Stipe 40–100 × 2–10 mm, central, terete, equal or
seldom slightly enlarged near the base, straight to curved
in age, moist to dry (not viscid), fragile, glabrous above,
base with hirsute short white mycelia, without rhizomorphs;
when young apex concolorous with pileus, then becoming
spinel red or spinel pink (12A7–6) towards the base, with
age it changes first to pinkish, Persian lilac (12A5), Congo
pink or light coral red (9A5–6) and finally discolors to pale
red, shell pink, orange-pink or buff pink (7A2–4). Context
hollow, thin (1–1.5 mm), pastel red, coral pink or Congo
pink (9A4–5). Odor and taste not distinctive.
Basidiospores broadly ellipsoid to ellipsoid, inequilateral in profile, hilar appendix < 1 µm, hyaline,
smooth, non-amyloid, non-dextrinoid, white in deposit,
(6–)7–8.5(–9.5) × (4–)5–6(–7) µm ( x = 7.61 × 5.48
µm, n = 20/5), Q = (1.07–)1.25–1.45(–1.6) ( x = 1.39
µm, n = 30/5). Basidia (30–)32–40(–44) × 5–7 µm
( x = 36.7 × 5.86 µm, n = 10/5), Q = 5–7.8 ( x = 6.24 µm,
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n = 10/5), predominantly 4-spored, seldom 3-spored, clavate,
hyaline, rare medallion clamp connections and seldom toroidal; sterigmata thin, straight or incurved, (6–)7–10(–11) µm,
however in some collections smaller (6–)6–8(–9) µm, these
collections have also smaller spores 6–8 × 4–5 µm. Abundant basidioles often with toruloid clamps and bifurcate
base. Hymenial cystidia absent. Pileipellis a weak ixocutis
composed of thin-walled, hyaline, non-incrusted, cylindrical, septate, subparallel to slightly interwoven hyphae,
(3–)3.5–4.5 µm diam, clamp connections absent. Pileus
trama of subparallel to interwoven short, inflated to fusoid
hyphae 4–8 µm diam. Hymenophoral trama subregular, composed of hyaline, nongelatinous, irregular, hyphal elements
(10–)12.5–17.5(–25) × 10–12.5(–20) µm, without cylindrical
hyphae, clamp connections absent. Stipe tissue monomitic;
stipitipellis a cutis composed of parallel or slightly interwoven, thin-walled, hyaline, cylindrical septate hyphae 3.75–5
µm diam, medullary hyphae wider, (15–)17.5–20(–22.5) µm
diam, caulocystidia absent, clamp connections absent.
Habit and habitat gregarious and occasionally caespitose,
amongst moss in mountain cloud subtropical forest dominated by Cupressus or Quercus spp.
Material examined: MEXICO, Oaxaca, Santa María
Huitepec, Cempoaltepetl, Cupressus subtropical cloud forest, 6 November 2018, Garibay-Orijel 2018-1A (MEXU
28225 holotype), Garibay-Orijel 2018-1B (MEXU 28226);
Oaxaca, Santa María Huitepec, Quercus spp subtropical
cloud forest, 6 November 2018, Garibay-Orijel 2018-2
(MEXU 28227), Garibay-Orijel 2018-3 (MEXU 28228),
Garibay-Orijel 2018-4, MEXU 28229).
GenBank numbers: ITS: MK332021, MK332022.
Notes: Phylogenetically, the ITS sequences of Humidicutis brunneovinacea form a distinct terminal lineage together
with three undescribed species (Fig. 83). Morphologically,
H. brunneovinacea is unique among described species of
Humidicutis due its combination of dark brown vinaceous
fading to vinaceous and red colors (Fig. 82). There are some
rose red undescribed species of Humidicutis in the Caribbean and Central America (Cantrell et al. 2001; Lodge et al.
2014), however these lack dark brown colors and their ITS
sequences are too divergent (nucleotide similarity < 85%)
with H. brunneovinacea to be considered the same species.
Microscopically, H. brunneovinacea is distinguished by the
very scarce clamp connections in the basidia and the subregular hymenophoral trama composed of irregular hyphal
elements without cylindrical hyphae. It is noteworthy that
these four species are distributed in the Caribbean and Mesoamerican region of the Neotropics and they share reddish
tinges unique in the genus.
Amylocorticiales K.H. Larss., Manfr. Binder & Hibbett
Notes: The order Amylocorticiales is a rather small
order in the Agaricomycetes, recently described based
Fungal Diversity (2020) 104:1–266
117
Fig. 82 Humidicutis brunneovinacea (MEXU 28225, holotype). a
Basidiomes in its habitat. b Basidiomes development. c Hymenophoral trama. d Hymenia. e Spores. f Basidiole with bifurcated base. g
Basidia arising from same toruloid clamp connection. Scale bars:
c = 40 µm, d, f, g = 20 µm, e = 5 µm
mainly on molecular phylogeny (Binder et al. 2010). The
order includes about 70 species of more than ten genera,
but several sequenced species that formed distinct lineages in the phylogenetic tree are not generic types (Larsson
2007; Niemelä et al. 2007; Binder et al. 2010; Buyck et al.
2012; Hibbett et al. 2014; Song et al. 2016; Chikowski et al.
2017). Amylocortiaceae is the only family in the order, and
an intensive revision at generic and family level is needed.
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118
Fungal Diversity (2020) 104:1–266
Fig. 83 Maximum likelihood analysis of Humidicutae showing the
phylogenetic position of Humidicutis brunneovinacea inferred from
ITS and LSU sequences. Bootstrap support values for ML ≥ 60% and
Bayesian posterior probabilities ≥ 0.80 are given near nodes respectively. The tree is rooted with Chromosera citrinopallida. The holotype of H. brunneovinacea is in bold
Species in the order have diverse characters, but they all have
a monomitic hyphal system with nodose septate hyphae, and
most of them have corticioid basidiocarps with smooth, merulioid or poroid hymenophores and amyloid basidiospores
(Bernicchia and Gorjón 2010; Hibbett et al. 2014).
Basidiocarp Annual or perennial, resupinate, effuse to
slight effuse-reflected, separable, pellicular to membranaceous, stratified. Hymenophore smooth, white, cream to
pale yellow; not cracking or sparsely cracking after dry;
margin thinning out, fimbriate. Hyphal system monomitic;
generative hyphae with clamp connections. Cystidia absent.
Basidia clavate to subcylindrical, with a basal clamp connection and four sterigmata. Basidiospores narrowly cylindrical,
hyaline, thin-walled, smooth, amyloid, 7–10 × 2–2.5 µm. On
angiosperm branch. Rot type unknown.
Notes: Amyloceraceomyces is mainly characterized by
the pellicular to membranaceous, stratified basidiocarps,
a monomitic hyphal system with nodose septate hyphae,
absence of sterile organs, and cylindrical smooth thin-walled
amyloid basidiospores. Phylogenetically, Amyloceraceomyces formed a distinct lineage in Amylocorticiales (Fig. 84).
Morphologically, the genus is almost indistinguishable from
Amylocorticium which, however, has ellipsoid to allantoid
basidiospores and causes a brown rot mainly on gymnosperm wood (Pouzar 1959; Hjortstam 1980; Gilbertson
and Lindsey 1989). Amyloathelia Hjortstam and Ryvarden
and Amylocorticiellum Spirin & Zmitr. differ from the new
genus by having thick-walled basidiospores (Hjortstam and
Amylocorticiaceae Jülich, Biblthca Mycol. 85: 354 (1982)
Notes: Amylocorticiaceae was first described to accommodate Amylocorticium, but later several genera were placed
in the family based on molecular phylogeny (Larsson 2007).
At present, Amylocorticiaceae is the only family of the
Amylocorticiales, and the delimitation of it is still not clear
(Larsson 2007; Binder et al. 2010).
Amyloceraceomyces S.H. He, gen. nov.
MycoBank number: MB830032; Facesoffungi number:
FoF 06037
Etymology: Amylo: referring to the amyloidity of the
basidiospores; ceraceomyces: a corticioid genus with similar
anatomical characters.
Type species: Amyloceraceomyces angustisporus S.H.
He.
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Fungal Diversity (2020) 104:1–266
Ryvarden 1979; Zmitrovich and Spirin 2002). Ceraceomyces
Jülich has similar anatomical structures to Amyloceraceomyces, but differs in having basidiospores without reaction in
IKI (Bernicchia and Gorjón 2010). Amyloceraceomyces is
also similar to Melzericium Hauerslev by sharing smooth,
thin-walled amyloid basidiospores and the absence of cystidia, but the latter has stalked basidia and wider basidiospores (3–6 µm, Bernicchia and Gorjón 2010).
Amyloceraceomyces angustisporus S.H. He, sp. nov.
MycoBank number: MB 830033; Facesoffungi number:
FoF 06038; Fig. 85
Etymology: Referring to the narrow basidiospores.
Holotype: BJFC 021279.
Basidiocarp Annual or perennial, resupinate, effuse to
slight effuse-reflected, separable, without odour or taste
when fresh, pellicular to membranaceous, more or less
brittle with age and after drying, effuse part up to 15 cm
long, 5 cm wide, 1.5 mm thick, stratified; reflected part
indistinct and curved after drying, slightly darker than the
hymenophore suface. Hymenophore smooth, white, cream
to pale yellow, not cracking or sparsely cracking after drying; margin thining out, fimbriate, white or concolorous with
hymenophoral surface. Hyphal system monomitic; generative hyphae with clamp connections at all septa. Subiculum
distinct, thick, with a loose texture; hyphae in this layer hyaline, thin- to slightly thick-walled, smooth or slight encrusted
with small crystals, moderately branched, frequently septate,
loosely interwoven, 2–4.5 µm diam. Subhymenium with a
rather compact texture; hyphae in this layer hyaline, thinwalled, frequently branched and septate, densely interwoven,
more or less agglutinated, 2.5–5.5 µm diam. Cystidia and
cystidioles absent. Basidia clavate to subcylindrical, with
a basal clamp connection and four sterigmata, 20–32 × 4–6
µm; basidioles similar in shape to basidia, but slightly
smaller. Basidiospores narrowly cylindrical to slightly
allantoid, hyaline, thin-walled, smooth, amyloid in Melzer’s
reagent, 7–10 × (1.8–)2–2.5 µm, L = 8.52 µm, W = 2.15 µm,
Q = 3.78–4.16 (n = 60/2).
Material examined: CHINA, Yunnan Province, Binchuan
County, Jizushan Forest Park, on dead angiosperm branch,
30 August 2017, He 2844 (BJFC 021279, holotype), He
2819 (BJFC 021256) & He 2824 (BJFC 021260).
GenBank numbers: ITS: MK520873, MK520871,
MK520872; LSU: MK491338, MK491337l.
Notes: Amyloceraceomyces angustisporus is characterized by the rather long and narrow basidiospores. It is
so far the only species of the genus. In the phylogenetic
tree, the new species is distantly related to Amylocorticium
cebennense (Bourdot) Pouzar (Fig. 84), but the latter species has pure white basidiomata and shorter basidiospores
(6.5–7.5 × 1.8–2.2 µm, Bernicchia and Gorjón 2010), and
grows on Pinus spp.
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Amylocorticium Pouzar, Česká Mykol. 13(1): 11 (1959)
Notes: Amylocorticium is characterized by resupinate
basidiocarps with a smooth hymenophore, lack of cystidia,
ellipsoid to allantoid basidiospores with amyloid reactions in
Melzer’s reagent and causing a brown rot on wood (Pouzar
1959; Hjortstam 1980; Gilbertson and Lindsey 1989; Bernicchia and Gorjón 2010). At present, the genus includes
about twelve species, but the status of several species needs
to be further clarified. The few sequenced species including the type, A. subsulphureum (P. Karst.) Pouzar, formed a
moderately supported lineage in Amylocorticiales (Binder
et al. 2010; Song et al. 2016; Chikowski et al. 2017).
Amylocorticium ellipsosporum S.H. He, sp. nov.
MycoBank number: MB830034; Facesoffungi number:
FoF 06039; Fig. 86
Etymology: Referring to the ellipsoid basidiospores.
Holotype: BJFC 023898.
Basidiocarp Annual, resupinate, effuse, separable, without odour or taste when fresh, pellicular to membranaceous,
up to 10 cm long, 2 cm wide, 0.3 mm thick. Hymenophore
smooth, cream to pale orange, not cracking or sparsely
cracking after drying; margin thining out, fimbriate, white
or concolorous with the hymenophoral surface. Hyphal
system monomitic; generative hyphae with clamp connections at all septa. Subiculum distinct, with a loose texture;
hyphae in this layer hyaline, thin- to slightly thick-walled,
moderately branched, frequently septate, loosely interwoven, 2–4 µm diam. Subhymenium with a rather compact
texture; hyphae in this layer hyaline, thin-walled, frequently
branched and septate, densely interwoven, more or less
agglutinated, 2.5–4.5 µm diam. Cystidia and cystidioles
absent. Basidia clavate, with a basal clamp connection and
four sterigmata, 15–20 × 4–5 µm; basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid,
hyaline, thin-walled, smooth, amyloid in Melzer’s reagent,
3.5–4.5(–5) × 2–2.5(–3) µm, L = 3.95 µm, W = 2.22 µm,
Q = 1.78 (n = 30/1).
Material examined: CHINA, Fujian Province, Wuyishan Nature Reserve, on rotten trunk of Tsuga chinensis var.
tchekiangenesis, 17 August 2016, He 4457 (BJFC 023898,
holotype).
GenBank numbers: ITS: MK520876; LSU: MK491341
Notes: Amylocorticium ellipsosporum is characterized
by having cream-colored basidiocarps and small ellipsoid
basidiospores. In the phylogenetic tree, A. ellipsosporum
formed a strongly supported lineage, sister to A. indicum
which is here reported in China for the first time (Fig. 84).
However, A. indicum has large yellow basidiocarps and
slenderer basidiospores (4–5 × 1.7–2.2 µm, Thind and Rattan 1972). A. ellipsosporum is similar to A. mauiense Gilb.
& Hemmes by sharing small ellipsoid basidiospores, but
the latter species differs in having shorter basidia (12–14
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120
Fungal Diversity (2020) 104:1–266
Fig. 84 Phylogenetic position of Amyloceraceomyces angustisporus
and Amylocorticium ellipsosporum inferred from the ITS and LSU
sequences. Bootstrap support values (ML and MP) ≥ 50% and Bayes-
ian posterior probabilities ≥ 0.95 are given near nodes respectively.
The tree is rooted with Jaapia argillacea (KHL 11734, CBS252.74)
and J. ochroleuca (KHL 8433). The new isolates are in bold
µm) and a distribution in Hawaii and growing on Pinus spp.
(Gilbertson and Hemmes 2004).
Cantharellales Gäum.
Notes: The order Cantharellales was proposed by Gäumann (1926). It is monophyletic and closely related to the
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121
Fig. 85 Microscopic structures of Amyloceraceomyces
angustisporus (BJFC 021279,
holotype). a Basidiospores.
b Basidia and basidioles. c
Hyphae from subiculum
Sebacinales, at the base of the Agaricomycetes clade (Moncalvo et al. 2006; Hibbett et al. 2014). It includes saprotrophic, ectomycorrhizal, lichenized, and parasitic species.
Members of this order display a wide variety of basidiocarp
morphology (e.g. effused, stipitate-hydnoid, stipitate-veined,
clavate, and coralloid) with a smooth, hydnoid or poroid
hymenophore. They are all characterized by developing
stichic basidia (Hibbett et al. 2014).
Clavulinaceae Donk, Beih. Nova Hedwigia 1(4): 407 (1970)
Notes: Clavulinaceae includes the genera Clavulina,
Membranomyces, and Multiclavula. The first two genera are
ectomycorrhizal lineages and the third one is lichenized. Clavulina is characterized by its clavarioid basidiomes, but several new species have been recognized with efuse-coralloid,
cerebriform and cantharelloid basidiomes; Membranomyces
has resupinate basidiomes that grow directly on soil; Multiclavula has clavate basidiomes and establishes mutualistic
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Fungal Diversity (2020) 104:1–266
Fig. 86 Microscopic structures of Amylocorticium
ellipsosporum (BJFC 023898,
holotype). a Basidiospores.
b Basidia and basidioles. c
Hyphae from subiculum
associations with green algae forming lichens. These taxa
have a monomitic hyphal system, with cylindrical to subclavate basidia commonly with two incurved sterigmata; and
small and smooth globose to subglobose basidiospores.
13
Clavulina J. Schröt., in Cohn, Krypt.-Fl. Schlesien (Breslau)
3.1(25–32): 442 (1888) [1889]
Notes: Currently there are 87 recognized species of Clavulina. This genus is distributed in temperate and tropical
Fungal Diversity (2020) 104:1–266
ecosystems being particularly diverse in the tropics, with
21 new species being described from the Neotropics in
the last 15 years (Thacker and Henkel 2004; Henkel et al.
2005, 2011; Uehling et al. 2012a, 2012b; Wartchow 2012;
Tibpromma et al. 2017; Pérez-Pazos et al. 2019). Species
are characterized by their clavarioid basidiomes, simple
or branched, and with matte colors ranging from white, to
grey or purplish tones. Microscopically these species have
smooth globose to subglobose basidiospores, generally less
than 10 µm long; the cylindrical to sub-clavate basidia usually with two incurved sterigmata (4–8 µm long) is considered a main characteristic of the genus; and sometimes
transversal septa develop in the basidia after basidiospores
release (Corner 1950, 1970; Petersen 1988; Olariaga et al.
2009), but this character is unstable even among different
specimens of the same species.
Clavulina sphaeropedunculata E. Pérez-Pazos, M. Villegas
& R. Garibay-Orijel, sp. nov.
MycoBank number: MB826922; Facesoffungi number:
FoF 05680; Fig. 87
Etymology: sphaeropedunculata (gr. sphaîra = sphere; lat.
pedunculus = peduncle; suffix –atus = provided with) epithet
assigned due to the presence of sphaeropedunculate cystidia
on the hymenial surface, a unique character in this species.
Holotype: FCME 27663.
Basidiocarps Simple to scarcely branched, 10–84 mm
long. Stipe well-delimited, 8–37 mm long × 1–5 mm wide,
pale yellowish white (2A2–4A2), cylindric to irregularly sub flattened, surface is smooth from the base to the
apex. Branches Acrotonic branching, b.r. = 1–3(4). Scarce
branches, dichotomous, lax, cylindrical to irregularly compressed, smooth surface (some with superficial groove at
the base of the branch), pale yellowish white (2A2–4A2),
with V axils and U-shaped; hymenium pale yellowish white
(2A2–4A2) to orange-white (5A2), and some with a little
more creamy patches (4A3) (in younger stages homogeneous pale yellowish white from the base to the top), when dry
deep yellow (4A8), to apricot yellow (5B6), and surface with
a frostlike appearance, due to the presence of spores. Tips
simple, acute to subacute, some rounded (few times cristate),
lax, concolorous to branches, becoming drab (6D3) by dehydration. Consistency fibrous-fleshy. Context initially solid,
yellowish white (4A2), hollow with age. Odor sometimes
slightly sweet and imperceptible taste. Basidia fibulate, (32–
)40–70(–81.5) × (4.5–)6–7.5(–9) μm, cylindrical to subclavate, thin-walled, hyaline, few with oily content, postpartal septa scarcely observed, 2 sterigmata, (4–)4.5–9(–9.5)
μm long, curved. Basidiospores subglobose, hyaline, thinwalled, smooth (7.7–)8.8–9.9(–10.4) × (6.3–)6.8–8.4(–
8.8) μm, (L = 8.9–9.6 ± 0.2–0.5, W = 7.2–8.0 ± 0.3–0.5,
Q = 1.20–1.29 ± 0.04–0.07), obtuse hilar appendix. Cystidia spheropedunculate, globose, thin-walled, smooth,
123
31–50 × 11.5–22 μm, infrequent. Hyphal system monomitic.
Subhymenium formed by fibulate hyphae, hyaline, and thinwalled, (4–)4.5–6(–7.5) μm wide.
Habitat and distribution: Gregarious, grows on the
ground and woody debris. Develops on temperate forests,
alt. 3000–3500 m.
Material examined: MEXICO, Mexico Estado de
Mexico City, El Zarco, Insurgente Miguel Hidalgo y Costilla National Park, Ocoyoacac municipality, San Pedro
Atlapulco, 19.293° N, 99.355° W, 28 November 2015,
Pérez-Pazos-467Z (FCME 27663, holotype); 17 October
2014, Argüelles-Moyao-104 (MEXU 28224); 28 November
2015, Pérez-Pazos-466 (FCME 27664); 28 November 2015,
Pérez-Pazos-472 (FCME 27665); MEXICO, Mexico City, El
Pantano, Desierto de los Leones National Park, Cuajimalpa
district, 19.293° N, 99.323° W, 13 October 2012, PérezPazos-43 (FCME 27666). MEXICO, Estado de Mexico,
Zinacantepec, El Contadero de Matamoros, 19.22° N, 99.82°
W, 3 September 2008, Endara Agramont ECMMX176-11.
ITS (MEXU 26712); Estado de Mexico, Amanalco, Corral
de Piedra, 19.22° N, 99.97° W, 21 September 2008, Burrola
Aguilar ECMAM152 (MEXU 25781).
GenBank numbers: ITS: MH542557, MH542560,
MH542556, MH542559, KT874987, KT874988,
MH542558; LSU: MK253717, MK253716, MK253714,
MK253718, MK253715; ITS sequence from an ectomycorrhizal root tip: MH542563.
Notes: In the phylogenetic analyses, Clavulina sphaeropedunculata is highly supported (ML = 99, PP = 1) within the
temperate species clade (Fig. 88). The main character that
distinguishes C. sphaeropedunculata from other similar species is the presence of sphaeropedunculate cystidia on the
hymenial surface, a unique character among all previously
reported species of the genus. Species reported with cystidia,
generally lack clamped hyphae in the hymenial trama, which
is not the case for C. sphaeropedunculata. Some of the species with cystidia are C. alutaceo-siccescens and C. geoglossoides, both reported by Petersen (1988) from the Nothofagus forests of New Zealand. In the former the cystidia are
longer (70–100 µm), cylindrical, and occurs in small clusters; in the latter, cystidia rise to 200 µm in length and are
often septate. Corner (1950) also reported some species with
cystidia, including C. leveillei, C. pilosa and C. cristata var.
incarnata. C. leveillei has cylindrical cystidia, 80–125 µm
length; C. pilosa also has cylindric cystidia, hair-like, 6 µm
wide that project 30 µm beyond the basidia layer; C. cristata var. incarnata also has long and cylindric cystidia, subventricose, sometimes becoming immersed on the hymenia.
In all the species mentioned, there is no cystidia resembling
those of C. sphaeropedunculata. It should be noted that cystidia in C. sphaeropedunculata are highly infrequent, so it is
likely that specimens of this species, previously deposited
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Fungal Diversity (2020) 104:1–266
Fig. 87 Clavulina sphaeropedunculata (FCME 27663, holotype). a
Simple basidiome, with acute tips. b Basidia and sphaeropedunculate
cystidia. c Sphaeropedunculate cystidia and spores. d. Sphaeropedun-
culate cystidia with smooth surface and thin-walled. Scale bars: a = 1
cm, b–d = 20 µm
in herbaria, may have them, but these have been overlooked
or collapsed with age.
hyaline, thin-walled, smooth spores. They include tropical
and temperate species (Jülich 1981).
Gomphales Jülich
Notes: The order Gomphales was proposed by Jülich in
1981. It is monophyletic and closely related to Geastrales,
Hysterangiales and Phallales (Villegas et al. 1999; Hosaka
et al. 2006; Giachini et al. 2010) and includes saprotrophic
and ectomycorrhizal representatives with wide morphological basidiocarp variation (e.g. resupinate, clavate, coralloid,
cantharelloid, hydnoid, and gasteroid). However, all species
in the Gomphales have a positive reaction to ferric salts in
the hymeniun, chiastic basidia and most taxa have cyanophilic ornamented spores (Donk 1964; Petersen 1971; Pine
et al. 1999; Villegas et al. 2005).
Lentaria Corner. Monograph of Clavaria and allied Genera,
(Annals of Botany Memoirs No. 1): 437, 696 (1950).
Notes: Initially, Lentaria included clavarioid species,
simple or branched, with smooth, hyaline spores, ellipsoid
to allantoid and phycophile or saprotrophic habitat. Phycophile species were later segregated in the genus Multiclavula
(Petersen 1967). Nowadays, Lentaria species are recognized
for developing coralloid basidiocarps and being saprotrophic, with humicolous or lignicolous habits. The basidiocarps emerge from a subiculum, superficial or immersed on
the substrate. The spores are hyaline, ellipsoid to sigmoid,
smooth and thin-walled and the hyphal system is monomitic.
However, it has been hypothesized that the genus is polyphyletic (Binder et al. 2010; Liu et al. 2017b). Twenty-three
species have been described, of which five are tropical.
Lentariaceae Jülich, Bibliotheca Mycologica 85: 375 (1982)
Notes: Lentariaceae includes the genera Delentaria and
Lentaria. Delentaria is monotypic, with simple to slightly
branched basidiocarps, which are sub-gelatinous and have
positive geotropism (Corner 1970). On the contrary, Lentaria develops basidiocarps with negative geotropism that
are widely branched and fleshy to leathery. These taxa have
a monomitic hyphal system, with thickened hyphal walls and
13
Lentaria gossypina R. Salas-Lizana, M. Villegas & E.
Pérez-Pazos, sp. nov.
MycoBank number: MB828900; Facesoffungi number:
FoF 05497; Figs. 89, 90
Fungal Diversity (2020) 104:1–266
125
Fig. 88 Phylogenetic tree of Clavulina obtained from the analyses
of ITS and LSU sequences showing the phylogenetic position of C.
sphaeropedunculata. Numbers above branches represent support val-
ues of maximum likelihood bootstrap and Bayesian inference posterior probabilities, respectively. Sequences generated in this work are
shown in bold. The tree was rooted with Hydnum repandum
Etymology: From Latin gossypium, referring to the
cottony mycelium that covers the stipe and basidiocarp
branches.
Holotype: FCME 27625.
Basidiocarps solitary, gregarious or caespitose, annual,
20–150 mm long. Subiculum superficial over the substrates,
extensive, white, plushy or cottony, with numerous whitish mycelial chords 1–2 mm thick. Stipe 10–30 × 2–10 mm,
cylindrical to flattened, greyish orange, brownish orange to
light brown (5B3, 5C4–6D4). Branches with four to five levels, predominantly dichotomous, flattened to sub-cylindrical,
with internode length decreasing gradually towards the apex,
parallel to semi-loose, greyish orange to light brown (5B4,
6D5) with light reddish brown tones (7/4 2.5Y), internodes
rounded to sub-acute; surface covered irregularly with a
thin layer of whitish mycelium, giving a cottony-tomentose
appearance. Apices sub-rounded to acute, white, yellowish white, orange-white or light brown (4A2, 5A2, 5D5).
Leathery, easily broken when dried; concolor context to the
surface. Smell slightly sweet, astringent flavor. Hyphal system monomitic, hyphae with thickened walls and conspicuous fibulae; occasionally ampulliform up to 12 μm wide.
Subiculum and mycelial chords with 1–4 μm wide hyphae,
covered with numerous crystals, irregular in shape and easily
detachable. Branch context with 3.5–10 μm wide hyphae and
some H connections. Hymenium thickened, amphigenous;
basidia sub-claviform, hyaline, tetrasporic, sterigmata 2–10
μm long, with fibulae at the base, homogeneous content,
28–48 × 6–8.5 μm. Spores sub-sigmoid, smooth, thin-walled,
(8–)9.3–12(13.5) × 2.5–3.5 μm, L = 11.5 μm, W = 2.3 μm,
Q = 3.9 μm (n = 30/1), which kept together in groups of two
to four once detached from the sterigma.
Habitat and distribution: Lignicolous, mostly growing on
highly rotten wood, producing white rot, sometimes developing on leaves or thorns of dead plants. Developing in subperennial and perennial forests, alt. 200 m.
Material examined: MEXICO, Campeche, Calakmul municipality, Escárcega-Chetumal highroad, Km 25
along the exit to the archaeological remains of Calakmul,
18o18′02" N, 89o50′47" W, 22 September 2003, VillegasRíos M. 2389 (FCME 27623); Chiapas, Tumbalá municipality, Cascadas de Bolo Naháu, 13 July 2010, Garibay-Orijel
2010–51 (FCME 27621); Tabasco, Macuspana municipality, Parque Estatal Agua Blanca 17°38′30" N, 92°27′18"
W, 3 September 2007, González Ávila 75 (FCME 27620);
Veracruz, San Andrés Tuxtla municipality, Estación de
Biología de Los Tuxtlas, Circuito 1 pathway, 18º35′0.174″
N, 95º4′25.8″ W, 13 September 2010, Villegas- Ríos M.
2749 (FCME 27624); ibid. 28 August 2011, Villegas-Ríos
M. 2796 (FCME 27625, holotype); ibid. 25 August 2012,
Villegas-Ríos M. 2854 (FCME 27627); ibid. Villegas-Ríos
M.2855 (FCME 27628); EL Vigia 1 pathway, 17 September
2011, Villegas-Ríos M. 2820 (FCME 27626); ibid. 28 July
2013, Ramírez-López I. 2013–14 (FCME 27622).
13
126
Fungal Diversity (2020) 104:1–266
Fig. 89 Majority-rule consensus tree of Lentaria gossypina, L. variabilis and reference sequences using a ITS and LSU concatenated
matrix. Bayesian posterior probabilities (PP) ≥ 0.84 and bootstrap
support values (ML) ≥ 70% are labeled. Type material for the new
species are in bold. Outgroups are Dyctiophora rubrovolvata and
Clathrus ruber
GenBank numbers: ITS: MK253203, MK253201,
MK253195, MK253198, MK253199, MK253202,
MK253197, MK253200, MK253196; LSU: MK253212,
MK253204, MK253218, MK253219, MK253221,
MK253220, MK253209).
Notes: The phylogenetic analyses of concatenated ITSLSU sequences place the new species in a well-supported
monophyletic group, independent from any other species
sequenced so far (Fig. 89). This species is different from all
the tropical Lentaria species described so far on account of
a very well-developed subiculum over the substrate, with the
mycelium extending over the basidiocarp surface, giving a
tomentose to cottony appearance. Additional observations
showed that developing basidiocarps of L. gossypina form
more mycelium or new basidiocarps when they touch the
surface of a substrate. Spore size range is similar to that of L.
bambusina P. Zhang & Zuo H. Chen (9.0–13 × 2.5–3.5 μm),
whose stipe is also covered with a plushy layer of mycelium. However, L. bambusina has a smaller basidiocarp size
(20–60 mm), different color (cinnamon brown) and the habitat is dead leaves of bamboo plants.
13
Lentaria variabilis M. Villegas, R. Garibay-Orijel & N.
Matías-Ferrer, sp. nov.
MycoBank number: MB828899; Facesoffungi number:
FoF 05496; Fig. 91
Etymology: From latin variabilis, variable, referring to its
variation in color and branch position.
Holotype: FCME 21524.
Basidiocarps gregarious, annual, 20–75 mm in length.
Subiculum and mycelial chords whitish, up to 2 mm thick,
generally immersed in the substrate, rarely superficial.
Stipe 10–35 × 1–7 mm, subcylindrical, pale yellow, greyish orange with brown tones to reddish brown (4A3, 5B3,
5C4, 7D3, 8/4 10YR). Branching up to five levels, predominantly dichotomous, cylindrical to slightly flattened, internode space reduces gradually towards the apex; parallel or
a bit loose, pale yellow, orange-grey with brown or light
brown tones (4A3, 6B2, 6C3, 7C–D4), rounded internodes
Fungal Diversity (2020) 104:1–266
127
Fig. 90 Basidiocarps of Lentaria gossypina (Pictures, Ramírez-López I.). a (FCME 27622). b (FCME 2796, holotype). Microscopic structures
(FCME 2796, holotype). c Basidiospores. d Subiculum hyphae. Scale bars: a–b = 10 mm, c–d = 1 μm
and smooth surface. Apices acute, white or pale yellow with
orange tones of different intensity (5A2–6C4). Sub-leathery consistency, easily broken when dry; context concolorous with surface. Smell not distinct. Taste slightly sour or
astringent flavor. Hyphal system monomitic, thick-walled
hyphae, conspicuously fibulate; occasionally ampulliform
fibulae up to 15 μm width. Subiculum and mycelial chords
with hyphae 1–5 μm width, covered with numerous crystals irregularly shaped, easily detachable. Branch context
with hyphae 3–8 μm thick and occasional H connections.
Hymenium thickened, amphigenous; basidia sub-claviform,
hyaline, tetrasporic, sterigmata 2–10 μm long, fibulate at the
base, homogeneous content or sometimes uni to multiguttulate, 20–74 × 6–11 µm. Spores subcylindrical to subsigmoid, smooth, thin-walled, (8–)10–15 × 2–5 µm, L = 11.7
µm, W = 2.7 μm Q = 3.13 µm (n = 30/1), remain together in
groups of four after basidium detachment.
Habitat and distribution: Lignicolous, growing in highly
decomposed wood in sub-perennial tropical forest, alt.
100–250 m.
Material examined: MEXICO, Campeche, Calakmul
municipality, Km 26 Escárcega-Chetumal highroad, Calakmul archaeological site deviation, 18°18′02" N, 89°50′47"
W, 25 September 2003, Villegas-Ríos M. 2429 (FCME
21524, holotype); ibid. 27 October 2002, García-Sandoval
2002–2034 (FCME 19864); ibid. 1 November 2001, GarcíaSandoval 2001–63 (FCME 19862). Archaeological site
Xpuhil, 19 September 2003, Villegas-Ríos M. 2382 (FCME
21518); ibid. 20 September 2003, Villegas-Ríos M. 2390
(FCME 21520); ibid. 28 October 2002, García-Sandoval
2002–42 (FCME 19861). Km 25.5 Escárcega-Chetumal
highroad, on the exit to Union 20 de Agosto, 22 September
2003, Villegas-Ríos M. 2408 (FCME 21522); ibid. 25 September 2003, Villegas-Ríos M. 2431 (FCME 21525); ibid.
Villegas-Ríos M. 2434 (FCME 21526); ibid. 25 October
2002, Villegas-Ríos M. 2274 (FCME 21516).
GenBank numbers: ITS: MK253189, MK253185,
MK253184, MK253192, MK253186, MK253191,
MK253193, MK253188, MK253194, MK253187;
LSU: MK253215, MK253206, MK253205, MK253211,
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Fig. 91 Basidiocarps of Lentaria variabilis (Pictures, J. Cifuentes).
a (FCME 21524, holotype). b (FCME 21520). c (FCME 21520).
Microscopic structures (FCME 21524, holotype) d Basidia, basidiole
and basidiospores. e Subiculum hyphae. f Ampulliform fibula. Scale
bars: a–c = 10 mm, d = 20 μm, e–f = 10 μm
LSU: MK253213, MK253207, MK253214, MK253216,
MK253217, MK253210.
Notes: As its epithet says, Lentaria variabilis is morphologically variable as is evident from the various forms
of basidiocarp branching, coloration and subiculum position. However, its microscopic characters are homogeneous
regarding spores, basidia and hyphal shape and size. Molecular data also showed that, despite this macro-morphological
variation, all sequenced individuals belong to a monophyletic group of an independent lineage (Fig. 89) of tropical
species. Some light-colored specimens of L. variabilis
resemble L. surculus Corner. However, spores of the latter
are smaller (8.6–12.6 × 2.9–4 μm, Petersen 2000).
13
Hymenochaetales Oberw.
Notes: The order Hymenochaetales now includes five
accepted families, viz. Hymenochaetaceae, Schizoporaceae,
Neoantrodiellaceae, Oxyporaceae and Nigrofomitaceae.
However, some genera in this order have uncertain positions at the family level and as well certain genera are not
monophyletic. Therefore, much more effort is needed to
construct a comprehensive phylogenetic framework for the
Hymenochaetales.
Fungal Diversity (2020) 104:1–266
Hymenochaetaceae Donk, Bull. Bot. Gdns Buitenz. 17(4):
474 (1948)
Notes: Hymenochaetaceae, typified by Hymenochaete
rubiginosa (Dicks.) Lév., belongs to the Hymenochaetales
and currently comprises 27 genera and about 490 known
species (Kirk et al. 2008). The family includes species with
ferruginous brown, coriaceous to woody, stipitate, pileate to
resupinate basidiocarps, poroid, hydnoid or smooth hymenophore, and xanthocroic microstructures (Dai 2010). The
hyphal system is monomitic or dimitic and clamp connections are absent. Dextrinoid basidiospores may be present in
some genera, also, sterile elements (setae) in the hymenium
or in the trama, a characteristic that, although not present
in all species, is very particular to the family (Ryvarden
2004; Dai 2010). Species produce white rots and several
are important phytopathogens (Zhou et al. 2016a). The genera in Hymenochaetaceae have been revised and the two
largest genera Phellinus Quél. and Inonotus P. Karst. have
been segregated into more natural groups (Wagner and Fischer 2001, 2002; Zhou et al. 2016b). Additionally, several
new genera and species have been proposed in the last years
(Rajchenberg et al. 2015; Drechsler-Santos et al. 2016; Wu
et al. 2016; Chen and Yuan 2017; Ji et al. 2017; SalvadorMontoya et al. 2015). Some important forest pathogens and
medicinal fungi are members of the Hymenochaetaceae (Dai
2012a, b; Wu et al. 2019).
Fuscoporia Murrill, N. Amer. Fl. (New York) 9(1): 3 (1907)
Notes: Fuscoporia is a well-established genus in the
Hymenochaetaceae and was originally described by Murrill (1907) with F. ferruginosa (Schrad.) Murrill as the type
species. The genus includes species with a wide geographic
distribution (Wagner and Fischer 2001, 2002; Baltazar et al.
2009; Vlasák et al. 2011; Raymundo et al. 2013; Chen and
Yuan 2017, Chen et al. 2019), characterized by resupinate
to pileate basidiocarps, dimitic hyphal system with hyaline,
narrow, thin-walled and simple-septate generative hyphae,
usually with encrustations and globose to ovoid, smooth and
hyaline basidiospores (Wagner and Fischer 2001, 2002; Baltazar et al. 2009; Chen and Yuan 2017).
Fuscoporia licnoides (Mont.) Oliveira-Filho & Gibertoni,
comb. nov.
MycoBank number: MB825474; Facesoffungi number:
FoF 05768; Figs. 92, 93
Basionym: Polyporus licnoides Mont., Annls Sci. Nat.,
Bot., Sér. 2 13: 204 (1840)
Material examined: BRAZIL, Distrito Federal, Brasília, PARNA de Brasília, October 2013, T.B. Gibertoni 26
(URM 87970 as F. callimorpha); Pará, Melgaço, FLONA de
Caxiuanã, Melgaço, 8 August 2013, A.S. Silva 495 (URM
87971 as F. callimorpha); Paraíba, Areia, RE Mata do PauFerro, 16 April 2012, C.R.S. Lira 634 (URM 83698 as F.
129
callimorpha); ibid. 16 July 2013, C.R.S. Lira 1307 (URM
85067 as P. gilvus); ibid. 21 July 2012, C.R.S. Lira 207
(URM 84107 as P. gilvus); Pernambuco, Recife, Campus
UFPE, 10 April 2017, J.R.C. Oliveira-Filho JRF140 (URM
91231 as P. gilvus); Rio de Janeiro, Angra dos Reis, PE
de Ilha Grande, December 2012, T.B. Gibertoni 05 (URM
87969 as F. callimorpha); Rondônia, Porto Velho, ESEC de
Cuniã, 11 February 2011, A.C. Gomes-Silva, E.A. Souza
& M.C. Flores 2080 (URM 83001 as P. gilvus); ibid. A.C.
Gomes-Silva, E.A. Souza & M.C. Flores 2209 (URM 83003
as P. gilvus); ibid. A.C. Gomes-Silva, E.A. Souza & M.C.
Flores 2173 (URM 83004 as P. gilvus).
GenBank numbers: ITS: MH392553, MH392554,
MH392555, MH392556, MH392557, MH392558,
MH392559, MH392560, MH392561; LSU: MH407352,
MH407353, MH407354, MH407355, MH407356,
MH407357, MH407358, MH407359, MH407360.
Notes: see discussion under F. semiarida.
Fuscoporia marquesiana Gibertoni & C.R.S. de Lira, sp.
nov.
MycoBank number: MB825476; Facesoffungi number:
FoF 05766; Fig. 94
Etymology: In honour of the late Marcos Marques, a
young and remarkable Brazilian mycologist.
Holotype: URM 83094.
Basidiomata annual to perennial, pileate, sessile, applanate, widely attached, rarely effused-reflexed, semicircular,
8–17 cm long, 4–10 cm wide, 0.1–1.2 cm thick, woody hard
in thick specimens and coriaceous in thin ones when dried.
Upper surface light to dark brown (12 fulvous to 16 cigar
brown), glabrous, sulcate in concentric and narrow zones,
rarely 2 zonate. Margin acute, rarely obtuse, entire, concolorous with upper surface. Context homogeneous, light
brown (12 fulvous), 0.1–0.2 cm at the middle of basidiomata. Pores tiny, almost invisible to the naked eye, circular,
8–9 per mm, dark brown (17 snuff brown), dissepiments
entire, thick, tubes light brown (12 fulvous), 0.1–1 cm.
Basidiospores broadly ellipsoid, thin-walled, hyaline to
pale golden yellow, 4–6 × 3–4 µm (L = 4.95 µm, W = 3.85
µm, Q = 1.29 µm), IKI–. Basidia clavate, hyaline, thinwalled, 9–10 × 4–5 µm. Hyphal system dimitic, generative
hyphae hyaline, narrow, thin-walled and simple-septate,
2–2.5 µm wide; skeletal hyphae dominating the context
and dissepiments, yellow to brownish, thick-walled, 2–3
µm wide. Hymenial setae lanceolate to ventricose, thickwalled, apex acuminate or hooked, brown, 20–30 × 5–11 µm
(L = 26.27 µm, W = 8 µm, setal index = 3.28 µm).
Material examined: BRAZIL, Bahia, Santa Teresinha,
Serra da Jibóia, 22 September 2010, T.B. Gibertoni 19
(URM 83094 as F. senex).
GenBank numbers: ITS: MH392544; LSU: MH407343.
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130
Fungal Diversity (2020) 104:1–266
Fig. 92 Phylogenetic tree of the pileate Fuscoporia obtained by
analyses of combined dataset ITS and nLSU sequences. Sequences
obtained in this study are in bold. Support values are maximum likelihood, maximum parsimony and Bayesian analyses. Bootstrap sup-
port values (ML and MP) ≥ 50% and Bayesian posterior probabilities ≥ 0.95 are given near nodes respectively. The tree was rooted with
Amyloporia carbonica and Fomes fomentarius
Notes: Fuscoporia marquesiana is characterized by
the robust, concentrically zoned basidiomata, and usually
hooked setae. It matches the description of F. wahlbergii
(Fr.) T. Wagner & M. Fisch. (Ryvarden and Johansen 1980),
a species described from South Africa (originally Natal, currently KwaZulu-Natal) (type not located: no answer from
UPS). However, the images of Pyropolyporus robinsoniae
Murrill (NY 743008), a neotropical synonym of F. wahlbergii, reveal macromorphological differences. Additionally, F.
marquesiana has only 97% identity with the only sequence
of F. wahlbergii available (AF311045, LSU, no answer
from REG). This sequence, furthermore, is from material
collected in Spain (Canary Islands).
Fuscoporia scruposa (Mont.) Gibertoni & Oliveira-Filho,
comb. nov.
MycoBank number: MB825475; Facesoffungi number:
FoF 05767; Fig. 95
Basionym: Polyporus scruposus Fr., Epicr. Syst. Mycol.
(Upsaliae): 473 (1838) [1836–1838]
Material examined: BRAZIL, Alagoas, Pilar, RPPN
Fazenda São Pedro, 12 November 2016, J.R.C. OliveiraFilho JRF90 (URM 91211 as P. gilvus); Maranhão, Bacurituba, Mata da Iracema, 8 January 2017, J.R.C. OliveiraFilho JRF119 (URM 91223 as P. gilvus); Humberto de
Campos, Sede, 19 January 2017, J.R.C. Oliveira-Filho
JRF130 (URM 91228 as P. gilvus); Minas Gerais, Brumadinho, Instituto Inhotim, 11 November 2016, TB
13
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131
Fig. 93 Fuscoporia licnoides (URM 83001; URM 83004). a Abhymenial surface. b Hymenial surface. c Setae. d Basidiospores. Photos: R.L.
Alvarenga. Scale bars: a = 1 cm, b = 1 cm, c = 10 μm, d = 5 μm
Fig. 94 Fuscoporia marquesiana (URM 83094, holotype). a Abhymenial surface. b Hymenial surface. c Setae. d Basidiospores. Photos: R.L.
Alvarenga. Scale bars: a = 1 cm, b = 1 cm, c = 10 μm, d = 5 μm
Gibertoni JRF99 (URM 91214 as P. gilvus); Pernambuco,
Igarassu, Refúgio Ecológico Charles Darwin, 12 May 2017,
J.R.C. Oliveira-Filho JRF148 (URM 91236 as P. gilvus);
Piauí, Caracol, Serra das Confusões, 29 March 2011, A.C.
Gomes-Silva 295 (URM 83185, as P. gilvus); ibid. 31 March
2011, A.C. Gomes-Silva 361 (URM 83187, as P. gilvus);
ibid. 15 March 2012, C.R.S. Lira 570 (URM 83957, as P.
gilvus); Rio de Janeiro, Rio de Janeiro, PARNA Tijuca,
November 2012, T.B. Gibertoni 33 (URM 87972, as F.
gilva).
Additional material examined: Fuscoporia formosana:
BRAZIL, Maranhão, São José de Ribamar, Praia do Araçagi,
13
132
26 February 2016, J.R.C. Oliveira-Filho 4B (URM 91197 as
P. gilvus); ibid. 26 February 2016, J.R.C. Oliveira-Filho 3B
(URM 91196 as P. gilvus); ibid. 26 February 2016, J.R.C.
Oliveira-Filho 6B (URM 91198 as P. gilvus); Pernambuco,
Recife, Jardim Botânico do Recife, 16 May 2017, J.R.C.
Oliveira-Filho JRF165 (URM 91243 as P. gilvus).
GenBank numbers: Fuscoporia scruposa ITS:
MH392545, MH392546, MH392547, MH392548,
MH392549, MH392550, LSU: MH407344, MH407345,
MH407346, MH407347, MH407348, MH407349. Fuscoporia formosana ITS: MH392551, MH392552; LSU:
MH407350, MH407351.
Notes: see discussion under F. semiarida.
Fuscoporia semiarida Lima-Júnior, C.R.S. de Lira & Gibertoni, sp. nov.
MycoBank number: MB825473; Facesoffungi number:
FoF 05769; Fig. 96
Etymology: (L.) semiaridus, referring to the dryer areas
of Brazil
Holotype: URM 83800.
Basidiomata annual to biannual, pileate, sessile, occasionally effused-reflexed, applanate, widely attached, semicircular, sometimes imbricate, 0.8–3.5 cm long, 0.7–2.8 cm
wide, 0.3–0.7 cm thick, corky in thick specimens and coriaceous to tough in thin ones when dried. Upper surface light,
greyish to dark brown (12 fulvous, 17 snuff brown, 28 milk
coffee), glabrous, sulcate in concentric and narrow zones.
Margin acute, entire or rarely weakly lobed, concolorous
with upper surface, occasionally golden yellow (H). Context
homogeneous, light brown (12 fulvous), 0.1–0.2 cm at the
middle of basidiomata. Pores tiny, almost invisible to the
naked eye, circular, 7–9/mm, dark brown (12 fulvous, 17
snuff brown), dissepiments entire, thick, tubes light to dark
brown (12 fulvous, 17 snuff brown), 0.1 cm.
Basidiospores ellipsoid, thin-walled, hyaline, 4–5 × 2–3.0
µm (L = 4.4 µm, W = 2.55 µm, Q = 1.73 µm), IKI–. Basidia
clavate, hyaline, thin-walled, 9–13 × 4–4.5 µm. Hyphal system dimitic, generative hyphae hyaline, narrow, thin-walled
and simple septate, 1.5–2.8 µm wide; skeletal hyphae
dominating in the context and dissepiments, yellow to
brownish, thick-walled, 2.5–3.9 µm wide. Hymenial setae
lanceolate to ventricose, thick-walled, apex acuminate,
brown, 18–26 × 5–7 µm (L = 22.93 µm, W = 5.47 µm, setal
index = 4.2 µm).
Material examined: BRAZIL, Ceará, Crato, Floresta
Nacional do Araripe-Apodi, C.R.S. Lira 872, May 15 2012,
(URM 83800, holotype; isotype O), on dead wood; Bahia,
Santa Terezinha, Serra da Jibóia, 24 September 2010, T.B.
Gibertoni 34 (URM 83436 as Phellinus gilvus); Ceará,
Crato, FLONA Araripe-Apodi, 17 May 2012, C.R.S. Lira
919 (URM 83798 as P. gilvus); Goiás, Alto Paraíso de
Goiás, Loquinhas, 17 March 2016, T.B. Gibertoni PH2
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(URM 91200 as P. gilvus); Pernambuco, Triunfo, Sítio Carro
Quebrado, 28 January 2010, C.R.S. Lira & C.A.S. Trajano
23 (URM 82508 as P. gilvus); ibid. 1 March 2010, C.R.S
Lira & N.C. Lima-Júnior 16 (URM 82510 as P. gilvus); ibid.
1 March 2010, C.R.S. Lira & N.C. Lima-Júnior 9 (URM
82511 as P. gilvus); Piauí, Caracol, PARNA Serra das Confusões, 14 March 2012, C.R.S. Lira 541 (URM 83926 as P.
gilvus).
Additional specimens examined: Fuscoporia callimorpha: CAMEROON, Campo Province, Akok lowland rain
forest reserve, 2 December 1991, M. Núñez & L. Ryvarden
31041 (O F-915116); ibid. 2 December 1991, M. Núñez
& L. Ryvarden 31066 (O F-915115); ZAMBIA, Victoria
Falls, 27 January 1988, L. Ryvarden 25297 (O F-915114);
ZAMBIA, Victoria Falls Riverine forest, 4 January 1990, L.
Ryvarden 27254 (O F-915112); ZIMBABWE, Manicaland
Province, J. Meikle Res. Station, 24 January 1994, M. Núñez
& L. Ryvarden 33811 (O F-915113); ZIMBABWE, Mvuma
district, Beacon Hill, 5 January 1987, L. Ryvarden 25028
(O F-915111).
GenBank numbers: ITS: MH392562, MH392563,
MH392564, MH392565; LSU: MH407361, MH407362,
MH407363, MH407364.
Notes: Fuscoporia semiarida is characterized by annual
to biannual basidiomata, sulcate in concentric narrow zones,
ellipsoid basidiospores and lanceolate to ventricose hymenial setae. It was previously identified as Phellinus gilvus
(Schwein.) Pat.[= F. gilva (Schwein.) T. Wagner & M.
Fisch.] and F. callimorpha (Lév.) Groposo, Log.-Leite &
Góes-Neto. P. gilvus has a blackish abhymenial surface and
thicker basidiomata, as seen in the type (PH 74281), and
was originally collected in the USA (Pennsylvania), while F.
callimorpha has perennial basidiomata, with zones that may
be absent, and narrower basidiospores; also the type was
originally collected in Madagascar (Ryvarden and Johansen
1980).
Fidalgo and Fidalgo (1968) considered F. gilva as a cosmopolitan, macromorphologically variable species, but recognized three major morphotypes in the “Phellinus gilvus
complex”: 1) P. gilvus sensu stricto, with thick and hard
basidiomata with blackish abhymenial surface, as seen in the
type (PH 74281); 2) P. licnoides (Mont.) Pat., with corky,
more flexible basidiomata with narrow concentric zones,
as seen in reference material (NY 3029273, 3029275, type
not located: no answer from PC); and 3) P. scruposus (Fr.)
Cunn., with corky, more flexible basidiomata without narrow concentric zones, as seen in reference material (NY
3029272, type not located: no answer from PC). These morphotypes were considered varieties of F. gilva [Polyporus
gilvus var. scruposus (Fr.) Henn., P. gilvus var. licnoides
(Mont.) Cleland & Cheel, and P. gilvus var. gilvus (Schwein.) Fr.], but they were usually treated as synonyms of F.
Fungal Diversity (2020) 104:1–266
133
gilva (www.indexfungorum.org/names/Names.asp, www.
mycobank.org/).
In our study, the Brazilian specimens identified as F. gilva
sensu lato are distributed in four clades (Fig. 92), representing the new species F. semiarida, the new combinations F.
licnoides (Mont.) Oliveira-Filho & Gibertoni and F. scruposa (Mont.) Gibertoni & Oliveira-Filho, and F. formosana,
a new occurrence in the Americas. Both F. semiarida and
F. licnoides would belong to the “P. licnoides group” in the
“P. gilvus complex”, while F. scruposa and F. formosana
would belong to the “P. scruposus group”. So far, no analyzed material or images match the type of F. gilva (PH
74281) and, thus, no F. gilva sensu stricto clade can be recognized. While morphologically three morphotypes can be
separated, microscopically the four species and the type of
F. gilva are almost identical (Table 3).
Fuscoporia semiarida is macro- and micromorphologically almost identical to F. licnoides (Figs. 93, 96; Table 3)
which was originally collected in French Guiana and to
which it is not related in the phylogenetic tree (Fig. 92).
Both species have strongly zonate, somewhat flexible basidiomata, and similar basidiospores and setae. They can be
roughly separated by geographical distribution: despite several collections in Southeast, North, Northeast and mid-West
Brazil, F. semiarida was found in vegetation surrounded by
dry forests (Caatinga), with one collection in Cerrado (Brazilian savanna), while F. licnoides was usually collected in
wetter forests in the Atlantic Rain Forest domain and in the
Amazonia, with also one collection in Cerrado, which seems
to be the overlapping region of occurrence of both species.
So far, F. semiarida seems restricted to Brazil, while F. licnoides has also been collected, besides from the type locality, in the USA (Everglades, Florida, JF692191, JF692190;
American Virgin Islands, JF692193, JF692192; no specific
location, AY558649, voucher not located), and unknown
localities (GU054105, GU054028, AF200240, vouchers not
located); all these sequences are only from the ITS region.
The specimens from Florida and the Virgin Islands were previously identified as F. callimorpha (https://mykoweb.prf.
Table 3 Basidiospore and seta
measures for species in the
“Phellinus gilvus complex”
Basidiospores
L
W
Q
Setae
L
W
Setal index
jcu.cz/polypores/); AY558649 as Phellinus torulosus (Pers.)
Bourdot & Galzin [= F. torulosa (Pers.) T. Wagner & M.
Fisch.]; AF200240 as P. tremulae (Bondartsev) Bondartsev & P.N. Borisov; and GU054105 and GU054028 were
treated as unidentified fungi. P. torulosus and P. tremulae
are morphologically different from both F. semiarida and
F. licnoides and were originally described from France
and Russia (Bryansk Region) respectively (Gilbertson and
Ryvarden 1987; Ryvarden and Gilbertson 1994). In addition,
P. tremulae belongs to Phellinus sensu stricto and AF200240
is a misidentification.
Fuscoporia formosana (T.T. Chang & W.N. Chou) T.
Wagner & M. Fisch. was originally described from Taiwan Island as Inonotus formosanus T.T. Chang & W.N.
Chou (Chang and Chou 1998) and was so far known only
from the type locality. Our specimens have 100% identity
with the only sequence available (AY059034, LSU), fortunately from the type. Basidiospores were not found in
our materials, but setae were abundant and similar to the
original description (ventricose, 18–30 × 5–7 μm) (Chang
and Chou 1998). F. formosana is morphologically similar
(Fig. 95, 97), but not related to F. scruposa (Fig. 92). The
F. scruposa clade is composed of Brazilian and American
specimens and MG966313 (ITS) from Mexico, KU139196
(ITS), KU139196 (ITS), KX065958 (ITS), KJ140621 (ITS)
and AY059025 (LSU; voucher not located, no answer from
REG) from the USA, and AF518636 (LSU) from unknown
locality (voucher not located) also seem to be F. scruposa.
Images of KP859305/KP859295, MG966313, KU139196,
KU139196, KX065958 and KJ140621 were analyzed and
share similar morphology.
Insumran et al. (2012) studied specimens of Fuscoporia gilva from Thailand and showed that Asian specimens
identified as such were distributed in three clades, but no
information about their morphology was provided. Recently,
F. atlantica Motato-Vásq., R.M. Pires & Gugliotta, a new
species macromorphologically similar to F. gilva sensu
lato was described from Brazil, but it differs by the hooked
setae (Pires et al. 2015). Our results are similar to those
F. gilva (PH 74281)
F. semiarida
(URM 83800)
F. licnoides
(URM 85067)
F. scruposa
(URM 83957)
F. formosana
(URM 91197)
N = 30*
4.49 µm
2.75 µm
1.63 µm
N = 15*
24 µm
5.9 µm
4.07 µm
N = 10
4.4 µm
2.55 µm
1.73 µm
N = 15
22.93 µm
5.47 µm
4.2 µm
N = 15
4 µm
2.27 µm
1.76 µm
N = 15
20.6 µm
4.6 µm
4.48 µm
N = 10
4.3 µm
2.1 µm
2.05 µm
N = 15
19.05 µm
4.95 µm
3.85 µm
–
–
–
–
N = 15
21.5 µm
5.25 µm
4.1 µm
*Measured by Erast Parmasto in 07/04/1983.
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Fungal Diversity (2020) 104:1–266
Fig. 95 Fuscoporia scruposa (URM 83187). a Abhymenial surface. b Hymenial surface. c Setae. d Basidiospores. Photos: R.L. Alvarenga.
Scale bars: a = 1 cm, b = 1 cm, c = 10 μm, d = 5 μm
Fig. 96 Fuscoporia semiarida
(URM 83800, holotype) a
Abhymenial surface b Hymenial
surface c Setae d Basidiospores.
Photos: R.L. Alvarenga. Scale
bars: a = 1 cm, b = 1 cm, c = 10
μm, d = 5 μm
of Insumran et al. (2012); specimens once identified as F.
gilva sensu lato from Brazil do not represent a single lineage, being distributed in at least four clades representing two
13
morphotypes of F. gilva sensu Fidalgo and Fidalgo
(1968) (Fig. 92). Our results also show that species in
Fungal Diversity (2020) 104:1–266
135
Fig. 97 Fuscoporia formosana
(URM 91243). a Abhymenial
surface. b Hymenial surface.
c Setae. Scale bars: a = 1 cm,
b = 1 cm, c–d = 20 μm
each morphotype are morphologically indistinguishable
(Figs. 93, 95, 96, 97) and so far can only be separated by
DNA analyses.
Rigidoporus Murrill, Bull. Torrey Bot. Club 32(9): 478
(1905)
Notes: Rigidoporus was first described by Murrill in
1905. The basionym of its type is Polyporus microporus
Fr. This genus is one of the most species-rich genera of
basidiomycete wood-decaying saprobes, causing a white
rot in deciduous and coniferous tree. Most of the species of
Rigidoporus have been described from the Northern Hemisphere (Murrill 1905; Cooke 1949; Corner 1987; Zeng 1992;
Ryvarden and Gilbertson 1994; Dai 1998, 2010, 2012a, b;
Hattori 2011; Zmitrovich and Malysheva 2014). Recently,
multi-gene molecular phylogenies, inferred from ribosomal
DNA sequence data, demonstrated the polyphyletic nature of
Rigidoporus, and the generic type was nested within a clade
in the Hymenochaetales. Moreover, the genus Oxyporus was
treated as a synonym of Rigidoporus and most of the Oxyporus species were transferred to Rigidoporus (Wu et al. 2017).
Rigidoporus juniperinus Gafforov, L.W. Zhou & E. Langer,
sp. nov.
MycoBank number: MB830651; Facesoffungi number:
FoF 06108; Figs. 98, 99
Etymology: The epithet reflects the name of the host
genus Juniperus.
Holotype: TASM 6139.
Basidiocarp annual or perennial, resupinate, without
odor or taste, soft when fresh and becoming hard when dry,
sometimes with moss cover and, up to 10 cm long, 4 cm
wide, and 3 mm thick at the center; margin sterile or fertile, whitish, soft, up to 2 mm wide. Pore surface whitish
when fresh, drying cream-colored to pale ochraceous; pores
circular to angular, 4–5 per mm; dissepiments thin, sometimes thick where close to margin of basidiocarp. Subiculum cream, up to 3 mm thick. Tubes concolorous with pore
surface, stratified, each tube layer up to 2 mm long. Hyphal
system monomitic; all hyphae hyaline with simple septate,
lacking clamp connections, negative in Melzer’s reagent,
cyanophilous in Cotton Blue; unchanged in KOH. Contextual skeletal hyphae straight and flexuous, thick-walled
with a large lumen, unbranched, usually regularly arranged,
interwoven, 4–5 μm diam. Tramal hyphae thin- to distinctly
thick-walled with a narrow to wide lumen, branched, frequently simple septate, strongly flexuous, interwoven, 1.8–3
μm diam. Hymenial cystidia present; clavate, or narrowly
clavate, sometimes curved, with apical encrustation, simpleseptate at base, hyaline, thin-walled, smooth, partly projecting into the hymenium, 11–17 × 4–5 µm; some hyphae at the
dissepiment edge bearing crystals and resembling hyphoid
cystidia; cystidioles absent. Basidia clavate, without basal
clamp connection, with a simple septum and four sterigmata,
15–20 × 4–5 µm; basidioles in shape similar to basidia, but
smaller, 10–18 × 3–5 µm. Basidiospores broadly ellipsoid to
globose or subglobose, hyaline, thin-walled, smooth, negative in Melzer’s reagent, acyanophilous in Cotton Blue, (3.9–
)4.2–4.5(–4.9) × 2.9–3(–3.5) µm, L = 4.1 µm, W = 3.02 µm,
Q = 1.29–1.50 (n = 30/1).
Material examined: UZBEKISTAN, Jizzakh Province,
Zaamin districts, Zaamin National Park, northern slope of
Turkistan range of Pamir-Alay mountain system, Juniper forest, on rotten stump and stem of Juniperus polycarpos var.
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Fungal Diversity (2020) 104:1–266
Fig. 98 Macroscopic structures of Rigidoporus juniperinus (TASM 6139, holotype). a Basidiocarps cover with moses. b Pores surface
seravschanica (Kom.) Kitam. (Cupressaceae), 10 September
2017, 1700 m alt., Y. Gafforov, YG575 (TASM 6139, holotype); ibid. Y. Gafforov, YG1070 (TASM 6140).
GenBank numbers: ITS: MK433640, MK433641; LSU:
MK433642, MK433643.
Notes: Rigidoporus juniperinus is characterized usually
by annual resupinate basidiocarps covered in moss, white to
cream-coloured pores, presence of hyphoid cystidia and lack
of cystidioles and globose to broadly ellipsoid basidiospores.
It grows on coniferous wood in juniper forests. From the
phylogenetic perspective inferred from the concatenated ITS
and LSU regions (Fig. 100), two specimens of R. juniperinus
formed a distinct clade with full support in Rigidoporus. The
closest phylogenetic species to R. juniperinus is R. cuneatus
(Murrill) F. Wu, Jia J. Chen & Y. C. Dai. However, there are
morphological differences between R. juniperinus and R.
cuneatus. R. juniperinus has broadly ellipsoid to subglobose,
basidiospores (3.9–)4.2–4.5(–4.9) × 2.9–3(–3.5) μm while
R. cuneatus has mostly globose and bigger basidiospores
(3–5 μm). Morphologically, R. juniperinus is very similar to
Oxyporus corticola (Fr.) Ryvarden. However, O. corticola
has larger and ovoid basidiospores ((4.5)5–6 × 3.5–4.5 μm,
Ryvarden and Gilberston 1994). R. juniperinus also resembles O. populinus (Schumach.) Donk by sharing annual,
resupinate basidiocarps covered with moss at the base,
cream-coloured pore surface, approximately the same shape
and size of basidiospores, and abundant encrusted cystidia
(Ryvarden and Gilberston 1994). However, the latter species
has small pores (6–8 per mm), capitate to cylindrical larger
cystidia, interwoven hyphae in the subiculum, and grows
on angiosperm wood, while R. juniperinus has bigger pores
13
(4–5 per mm), most of the skeletal hyphae are regularly
arranged in the subiculum, an absence of cystidioles and
distinctly thick-walled hyphoid cystidia, clavate and bigger
basidia and grow on coniferous wood. These fungal species
have many crystals and encrustations on the hyphae which
is a common ecological adaption for aphylloporoid species
that minimizes water loss in arid and semiarid regions of
Central Asia (Gafforov et al. 2017).
Polyporales Gäum.
Notes: The order Polyporales is strongly supported as a
clade of the Agaricomycetes. About 13 families, 216 genera and 1801 species had been recorded in the order before
2008 according to the 10th edition of Dictionary of the
Fungi (Kirk et al. 2008). In the last ten years, many studies
focused on the taxonomy and phylogeny of taxa belonging
to the Polyporales; one new family, Fragiliporiaceae (Zhao
et al. 2015), and many new genera have been proposed in
the order (Cui et al. 2011, 2019; Li and Cui 2013; Zhao
et al. 2013, 2014; Li et al. 2014; Han et al. 2016; Chen et al.
2017; Shen et al. 2019; Sun et al. 2020). Recently, Justo et al.
(2017) provided a revised family-level classification of the
Polyporales and proposed three new families. In addition,
some economically important species are the memebrs of
Polyporales (Dai and Yang 2008; Cao et al. 2012).
Fomitopsidaceae Jülich, Biblthca Mycol. 85: 367 (1981)
Notes: Fomitopsidaceae was established by Jülich (1981)
and typified by the genus Fomitopsis P. Karst. Around 24
genera and 197 species were accepted in the family before
Fungal Diversity (2020) 104:1–266
137
Fig. 99 Microscopic structures
of Rigidoporus juniperinus
(TASM 6139, holotype). a
Basidiospores. b Basidia. c
Basidioles. d Hymenial cystidia
e Cystidioid hyphal ends. f
Hyphae from trama. g Hyphae
from subiculum
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138
Fungal Diversity (2020) 104:1–266
Fig. 100 Phylogenetic position of Rigidoporus juniperinus inferred
from the ITS and LSU sequences. Bootstrap support values (ML and
MP) ≥ 50% and Bayesian posterior probabilities ≥ 0.95 are given near
nodes respectively. The tree is rooted with Phellinus ferrugineovelutinus. The new isolates are in bold
2008 according to the 10th edition of Dictionary of the
Fungi (Kirk et al. 2008). Recently, taxonomic and phylogentic studies were carried out on Fomitopsidaceae and
eleven new genera were proposed: Amaropostia B.K. Cui,
L.L. Shen & Y.C. Dai, Calcipostia B.K. Cui, L.L. Shen &
Y.C. Dai, Cystidiopostia B.K. Cui, L.L. Shen & Y.C. Dai,
Fuscopostia B.K. Cui, L.L. Shen & Y.C. Dai, Fragifomes
B.K. Cui, M.L. Han & Y.C. Dai, Niveoporofomes B.K. Cui,
M.L. Han & Y.C. Dai, Piptoporellus B.K. Cui, M.L. Han &
Y.C. Dai, Rhodofomitopsis B.K. Cui, M.L. Han & Y.C. Dai,
Rubellofomes B.K. Cui, M.L. Han & Y.C. Dai, Ungulidaedalea B.K. Cui, M.L. Han & Y.C. Dai and Pseudofibroporia
Yuan Y. Chen & B.K. Cui (Han et al. 2016; Chen et al. 2017;
Shen et al. 2019), and many new species were described
(Cui 2013; Cui and Dai 2013; Li et al. 2013; Han et al. 2014,
2015, 2016; Shen and Cui 2014; Shen et al. 2014, 2015,
2019; Song et al. 2014, 2018c; Chen et al. 2015a, b; Han
and Cui 2015; Chen and Cui 2016; Chen et al. 2017; Song
and Cui 2017). Fomitopsidaceae is the most important family of the brown-rot fungi. Species previously belonging to
Fomitopsidaceae were divided into three clades at family
level according to Justo et al. (2017), which needs to be
clarified in the future.
Notes: Rhodofomitopsis was recently segregated from
Fomitopsis based on morphological characters and phylogenetic evidence. It differs from Fomitopsis by its rose, violaceous to pinkish brown pore surface and context, round,
angular, or slightly labyrinthiform, subdaedaleoid to sinuous-daedaleoid pores and absence of cystidia or other sterile
hymenial elements (Han et al. 2016). In our phylogenetic
analysis based on the combined dataset of ITS + nLSU + nS
SU + mtSSU + RPB2 + TEF1 sequences, species of Rhodofomitopsis formed a separate lineage (Fig. 101), and were
distant from Fomitopsis and Rubellofomes B.K. Cui, M.L.
Han & Y.C. Dai.
Rhodofomitopsis B.K. Cui, M.L. Han & Y.C. Dai, Fungal
Diversity 80: 365 (2016)
13
Rhodofomitopsis pseudofeei B.K. Cui & Shun Liu, sp. nov.
MycoBank number: MB829661; Facesoffungi number:
FoF 06041; Figs. 102, 103
Etymology: Referring to the species being similar to Rhodofomitopsis feei (Fr.) B.K. Cui, M.L. Han & Y.C. Dai in
morphology.
Holotype: Cui 16794 (BJFC).
Basidiocarps annual, pileate, sessile, solitary or imbricate, easily separable from the substrate, without odor or
taste, corky when fresh, becoming hard corky and light in
weight when dry. Pilei applanate, semicircular to flabelliform, projecting up to 4.2 cm, 8.7 cm wide and 2.5 cm
thick at base. Pileal surface greyish brown to fuscous at base
and cream to honey yellow toward the margin, glabrous to
Fungal Diversity (2020) 104:1–266
139
Fig. 101 Phylogenetic analyses of Rhodofomitopsis inferred from the
ITS, nLSU, mtSSU, nSSU, RPB2 and TEF1 sequences. Bootstrap
support values (ML and MP) ≥ 50% and Bayesian posterior probabili-
ties ≥ 0.95 are given near nodes respectively. The tree is rooted with
Antrodia tanakae (Cui 9743 and Yuan 1106). The new species and
new combinations are in bold
slightly velutinate, azonate, slightly sulcate or not; margin
obtuse to acute. Pore surface clay buff to clay pink when
dry; sterile margin distinct, pale mouse grey to greyish
brown, up to 4 mm wide; pores round and small, but somewhat variable, mostly 5–7 per mm and almost invisible to
the naked eye; dissepiments thick, entire. Context buff to
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140
Fungal Diversity (2020) 104:1–266
3.2(–3.3) µm, L = 6.57 µm, W = 2.87 µm, Q = 2. 19-2.32
(n = 90/3).
Material examined: AUSTRALIA, Queensland, Cairns,
nearby Mount Whitfield Conservation Park, on fallen angiosperm trunk, 7 May 2018, Cui 16794 (BJFC, holotype); on
fallen angiosperm trunk, 7 May 2018, Cui 16762 (BJFC);
on fallen angiosperm trunk, 7 May 2018, Cui 16803 (BJFC);
on fallen angiosperm trunk, 7 May 2018, Cui 16807 (BJFC).
GenBank numbers: ITS: MK461952, MK461951,
MK461953, MK461954; LSU: MK461956, MK461955,
MK461957, MK461958; mtSSU: MK461960, MK461959,
MK461961, MK461962; nSSU: MK461964, MK461963,
MK461965, MK461966; RPB2: MK463984, MK463983,
TEF1: MK463986, MK463985, MK463987.
Notes: Rhodofomitopsis pseudofeei was found from tropical areas of Australia. Morphologically, it is similar to R.
feei by the pileate and pinkish basidiocarps (Gilbertson and
Ryvarden 1986). However, the latter species has smaller
basidiospores (5–6.5 × 2–3 µm), and lacks cystidioles (Han
and Cui 2015). Phylogenetically, R. pseudofeei is distinct
from R. feei (Fig. 101). Fomitopsis subfeei B.K. Cui & M.L.
Han was recently described from China (Han and Cui 2015)
and transferred to Rhodofomes Kotl. & Pouzar as a new combination: Rhodofomes subfeei (B.K. Cui & M.L. Han) B.K.
Cui, M.L. Han & Y.C. Dai (Han et al. 2016), which also
has pileate and pinkish basidiocarps and may be confused
with Rhodofomitopsis pseudofeei, but it is distinguished by
its bigger pores (4–6 per mm) and smaller basidiospores
(4–5 × 1.9–2.4 mm, Han and Cui 2015).
Fig. 102 Basidiocarps of Rhdofomitopsis pseudofeei (Cui 16794,
holotype). Scale bar: = 2 cm
reddish brown or cinnamon brown, corky, up to 15 mm
thick. Tubes concolorous with pore surface, hard corky, up
to 2 mm long. Hyphal system dimitic; generative hyphae
bearing clamp connections; skeletal hyphae IKI–, CB–; tissues becoming blackish brown in KOH. Context generative
hyphae hyaline, thin-walled, occasionally branched, 1.7–3.8
µm diam; skeletal hyphae dominant, hyaline to pale yellowish, thick-walled with a wide to narrow lumen, frequently
simple-septate, occasionally branched, interwoven, 2–5.5
µm diam. Trama generative hyphae hyaline, thin-walled,
occasionally branched, 1.8–2.6 µm diam; skeletal hyphae
dominant, hyaline to pale yellowish, thick-walled with a
wide to narrow lumen, frequently simple-septate, occasionally branched, interwoven, 1.5–4 µm diam. Cystidia absent,
but fusoid cystidioles occasionally present, hyaline, thinwalled, 12–23 × 3–5 µm. Basidia clavate, bearing four sterigmata and a basal clamp connection, 12–23 × 4–6.5 µm;
basidioles dominant, in shape similar to basidia, but smaller.
Basidiospores cylindrical, sometimes curved, hyaline, thinwalled, smooth, IKI–, CB–, (5.2–)5.9–7(–8.4) × (2–)2.5–
13
Rhodofomitopsis monomitica (Yuan Y. Chen) B.K. Cui,
Yuan Y. Chen & Shun Liu, comb. nov.
MycoBank number: MB829662; Facesoffungi number:
FoF 06042
≡ Antrodia monomitica Yuan Y. Chen, Mycosphere 8(7):
882 (2017)
Notes: Antrodia monomitica was recently described from
China (Chen and Wu 2017). According to the phylogenetic
analyses based on ITS + nLSU + nSSU + mtSSU + RPB2 +
TEF1 sequences, it fell into the Rhodofomitopsis group, thus
we transferred it to Rhodofomitopsis as a new combination.
Rhodofomitopsis oleracea (R.W. Davidson & Lombard)
B.K. Cui, Yuan Y. Chen & Shun Liu, comb. nov.
MycoBank number: MB829663; Facesoffungi number:
FoF 06043
≡ Poria oleracea R.W. Davidson & Lombard, Mycologia
39(3): 317 (1947)
≡ Antrodia oleracea (R.W. Davidson & Lombard)
Ryvarden, Prelim. Polyp. Fl. E. Afr. (Oslo): 252 (1980)
Notes: Antrodia oleracea was reported from North America and Africa previously (Gilbertson and Ryvarden 1986;
Ryvarden and Johansen 1980), and has been reported as a
Fungal Diversity (2020) 104:1–266
141
Fig. 103 Microscopic structures
of Rhdofomitopsis pseudofeei (Cui 16794, holotype). a
Basidiospores. b Basidia and
basidioles. c Basidioles. d
Hyphae form trama. e Hyphae
from context
new Chinese record based on morphological characters (Du
et al. 2009). According to the phylogenetic analyses based
on ITS + nLSU + nSSU + mtSSU + RPB2 + TEF1 sequences,
it grouped together with R. monomitica in the Rhodofomitopsis group, thus we transferred it to Rhodofomitopsis as a
new combination.
Polyporaceae Fr. ex Corda [as ’Polyporei’], Icon. Fung.
(Prague) 3: 49 (1839)
Notes: The family was typified by Polyporus P. Micheli ex
Adans. (Fries 1838), and its concept was originally described
as a family of the Aphyllophorales that included all fungi
with poroid hymenophores. Subsequently, the concept of
Polyporaceae has been revised by many mycologists, and
the classifications of Polyporaceae have been changing for
a long time (Karsten 1879, 1881, 1892; Patouilliard 1900;
Donk 1960, 1964; Jülich 1981; Ryvarden 1991; Zhao 1998;
Kirk et al. 2008). Recently, systematic studies on the taxonomy and phylogeny of some genera of Polyporaceae have
been reported, but comprehensive studies on the family level
in the Polyporaceae are still lacking. Although Justo et al.
(2017) presented a systematic revision of Polyporales at the
order level based on sequences from LSU, ITS and RPB1
genes, only limited taxa of Polyporaceae were included in
their analyses. Cui et al. (2019) performed a comprehensive study on the species diversity, taxonomy and phylogeny
of Polyporaceae in China. They recorded 217 species and
provided an outline of the phylogeny of the Polyporaceae
in China. The definition of Polyporaceae still needs to be
revised in the future.
Antrodiella Ryvarden & I. Johans., Prelim. Polyp. Fl. E. Afr.
(Oslo): 256 (1980)
13
142
Notes: Antrodiella was established in 1980 to embrace
the Polyporus semisupinus Berk. & M.A. Curtis complex
(Ryvarden and Johansen 1980). It was characterized by resupinate, effused-reflexed to pileate basidiocarps with elastic consistency when fresh and dense and hard when dry,
small pores, light-colored pore surface, a dimitic to trimitic
hyphal structure, small ellipsoid to cylindrical basdiospores
and producing a white rot (Núñez and Ryvarden 2001; Dai
2004). More than 50 species have been described worldwide (Yuan 2014). The circumscription of Antrodiella has
been expanded greatly, but the cyanophilic reaction of the
skeletal hyphae was always considered an important and
consistent character. Recently, DNA-based molecular phylogenetic studies have been revealed the non-monophyletic
background of Antrodiella, and showed very complicated
relationships among the genera of Antrodiella, Flaviporus,
Junghuhnia and Steccherinum (Vampola and Vlasák 2011;
Miettinen et al. 2012; Yuan 2013, 2014).
Antrodiella descendena (Corner) C.L. Zhao & Y.C. Dai,
comb. nov.
Index Fungorum number: IF556447; Facesoffungi number:
FoF 06040; Figs. 104, 105c, d
Basionym: Tyromyces descendens Corner, Beih. Nova Hedwigia 96: 167 (1989).
Basidiocarp basidiocarps annual, pileate. Pileus semicircular, projecting up to 1 cm, 1.5 cm wide, and 1 cm thick at
base. Pileal surface buff to pale brown on drying, smooth,
azonate; margin obtuse. Pore surface orange-brown to brown
upon drying; pores angular, 4–5 per mm; dissepiments thin.
Context cream to yellowish brown, corky, about 2 mm
thick. Tubes concolorous with pore surface, corky, up to 8
mm thick. Hyphal system dimitic; generative hyphae with
clamp connections; skeletal hyphae IKI–, CB–, present in
trama only; tissues unchanged in KOH. Subicular generative
hyphae hyaline, thin-walled, occasionally branched, 4.0–5.0
µm diam; skeletal hyphae yellowish brown, thick-walled,
with a wide lumen, unbranched, loosely interwoven, 3–4.5
µm diam. Tramal generative hyphae infrequent, hyaline,
thin-walled, usually unbranched, 3.0–4.5 µm diam; skeletal
hyphae dominant, hyaline, thick-walled with a wide to narrow lumen, unbranched, interwoven, 3.0–5.0 µm diam. Cystidia and cystidioles absent; basidia and basidioles absent.
Basidiospores cylindrical, hyaline, thin-walled, smooth,
IKI–, CB–, 3.0–3.4(–3.6) × 1.0–1.3(–1.5) µm, L = 3.16 µm,
W = 1.12 µm, Q = 2.82 (n = 30/1).
Material examined: MALAYSIA, Johore, Gunong Panti,
28 April 1941, leg. E.J.H. Corner, E00159582 (E).
Notes: The type specimen of Tyromyces descendens was
examined by Hattori (2002) but he did not find basidiospores
and treated it as Antrodiella sp. based on a dimitic hyphal
system. In the present study, we found the basidiospores
by re-examining the type specimen and confirmed that the
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specimen belongs to Antrodiella. Therefore, we propose a
new combination A. descendena although we did not get
DNA from the type material. A. americana Ryvarden &
Gilb. and A. descendena share basidiospores of similar size
(2.5–3.5 × 1.0–1.8 µm). However, the former differs by
resupinate basidiocarps and larger-sized pores (1–2 per mm,
Núñez and Ryvarden 2001).
Tyromyces P. Karst., Revue Mycol., Toulouse 3(no. 9): 17
(1881)
Notes: Tyromyces is a large, cosmopolitan genus that
occurs in almost all types of forests from boreal to the tropics (Gilbertson and Ryvarden 1987; Ryvarden and Melo
2014; Dai 2012a, b). The genus is characterized by annual,
pileate basidiocarps which are sappy when fresh becoming
rigid and fragile when dry; a white to cream pore surface,
a momomitic to dimitic hyphal system with clamp connections on the generative hyphae; allantoid to ovoid, thinwalled basidiospores without reaction in IKI and production
of a white rot-type (Núñez and Ryvarden 2001; Ryvarden
and Melo 2014).
Tyromyces minutulus Y.C. Dai & C.L. Zhao, sp. nov.
Index Fungorum number: IF556446; Facesoffungi number:
FoF 06044; Figs. 105a, b, 106
Etymology: Referring to the small pileus.
Holotype: BJFC017406.
Basidiocarp annual, pileate, occasionally attached by a lateral tapering base, soft and sappy when fresh, becoming
corky upon drying. Pilei more or less semicircular, projecting up to 1 cm, 1.5 cm wide, 1.5 mm thick at base. Pileal
surface concentrically zonate, white when juvenile, becoming light chocolate to pale brown in centre, more or less
glabrous to slightly rough. Pore surface white when fresh,
white to cream upon drying; pores round to angular, 7–9
per mm; dissepiments thin, entire. Sterile margin white, up
to 2 mm wide. Context white, corky, up to 0.5 mm thick.
Tubes concolorous with pore surface, corky, up to 1 mm
long. Hyphal system monomitic; generative hyphae with
clamp connections, IKI–, CB–; tissues unchanged in KOH.
Subicular generative hyphae hyaline, thick-walled with a
wide lumen, unbranched, interwoven, 2.5–4.5 µm diam.
Tramal generative hyphae hyaline, thin- to thick-walled,
frequently branched, interwoven, 2.0–3.0 µm diam. Cystidia absent, but fusoid cystidioles present, hyaline, thinwalled, 7.0–8.0 × 3.0–3.5 µm; basidia more or less barrelshaped, with four sterigmata and a basal clamp connection,
9.0–11.0 × 3.0–4.5 µm; basidioles dominant; basidioles in
shape similar to basidia, but slightly smaller. Basidiospores
cylindrical, hyaline, thin-walled, smooth, tapering at apiculus, usually bearing one or two guttules, IKI–, CB–, (3.5–)
3.7–4(–4.2) × 1.0–1.3(–1.5) µm, L = 3.89 µm, W = 1.18 µm,
Q = 3. 19-3.45 (n = 60/2).
Fungal Diversity (2020) 104:1–266
143
Fig. 104 Microscopic structures
of Antrodiella descendena
(E00159582, holotype). a
Basidiospores. b Hyphae from
trama. c Hyphae from context.
Scale bar: a = 5 μm, b–c = 10
μm
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Fungal Diversity (2020) 104:1–266
Fig. 105 Basidiocarps of Tyromyces minutulus (BJFC017406, holotype) and the combination taxon Antrodiella descendena (holotype). a, b
Tyromyces minutulus. c, d Antrodiella descendena. Scale bars: a = 0.5 cm, e–f = 2 mm
Material examined: CHINA, Hainan Province, Changjiang County, Bawangling Nature Reserve, on rotten angiosperm wood, 16 June 2014 Dai 13666 (holotype); on fallen
angiosperm trunk, 8 May 2009 Cui 6427 (BJFC).
GenBank numbers: ITS: KM598443; LSU: KM598445.
Notes: Phylogenetically, the new species, Tyromyces
minutulus, groups with T. chioneus, T. kmetii (Bres.) Bondartsev & Singer, Piloporia sajanensis and Skeletocutis amorpha (Fig. 107). However, morphologically T. chioneus differs from T. minutulus by its larger pores (3–4 per mm) and
basidiospores (4.0–5.0 × 1.5–2.0 µm, Núñez and Ryvarden
2001). T. kmetii is separated from T. minutulus by the orange
pore surface and larger basidiospores (4.0–4.5 2.5–3.0 µm,
Gilbertson and Ryvarden 1987). Skeletocutis amorpha
differs in its resupinate to effuso-reflexed basidiocarps, a
dimitic hyphal system and allantoid basidiospores (Núñez
and Ryvarden 2001; Ryvarden and Melo 2014). Piloporia
13
sajanensis is distinguished from T. minutulus by the distinctly duplex structure of the context and the brown hyphae
(Niemelä 1982). Morphologically the basidiospores of the
new species are reminiscent of several similar Tyromyces
species. T. polyporoides Ryvarden & Iturriaga differs in
its laterally stipitate basidiocarps (Ryvarden and Iturriaga
2003); T. americanus (D. Reid) Ryvarden & Iturriaga differs from T. minutulus by substipitate basidiocarps and pale
orange pore surface (Ryvarden and Iturriaga 2003); T. leucomallus (Berk. & M.A. Curtis) Murrill differs in having
larger basidiocarps and gloeoplerous hyphae (Núñez and
Ryvarden 2001); and T. subviridis Ryvarden & Guzmán is
distinguished by ochraceous basidiocarps and a grey to pale
green pore surface (Ryvarden and Guzmán 2001).
Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C. David
Fungal Diversity (2020) 104:1–266
145
Fig. 106 Microscopic structures of Tyromyces minutulus
(BJFC017406, holotype). a
Basidiospores. b Basidia and
basidioles. c Cystidioles. d
Hyphae from trama. e Hyphae
from context. Scale bars: a = 5
μm, b–e = 10 μm
Notes: The taxonomic placement of the family was very
controversial in earlier days. Pegler (1977) and Singer
(1986) agreed with Lotsy (1907) and proposed the family
name Russulaceae and placed it in the Agaricales (Smith
1973; Pegler 1977; Singer 1986). Based on Singer (1986),
the family possessed a wide range of characters like fleshy
(mostly), often vividly coloured basidiocarps with free to
decurrent lamellae, heteromerous lamellae trama without
clamp-connections, and amyloid basidiospores with distinct
ornamentation. These features easily help to distinguish the
family from all the other macrofungal families. Similarly,
other mycologists like Kreisel (1969) and Kühner (1980)
proposed to include the family under Russulales due to the
presence of heteromerous trama, and amyloid and ornamented basidiospores. Kirk et al. (2001) fixed the proper
placement of the family in Russulales. Recently, a molecular
phylogenetic approach confined the family to the “russuloid
clade” that consists of a morphologically diverse group of
fungi forming a distinct lineage (Hibbett and Thorn 2001;
Larsson and Larsson 2003). Finally, to our present knowledge, Russulaceae is classified under Russulales in the class
Agaricomycetes (Hibbett et al. 2007).
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146
Fungal Diversity (2020) 104:1–266
Fig. 107 Maximum Parsimony strict consensus tree illustrating the
phylogeny of Tyromyces minutulus with related species in Polyporales based on ITS and LSU sequences. Branches are labeled with
maximum likelihood bootstrap ≥ 70%, parsimony bootstrap propor-
tions ≥ 50% and Bayesian posterior probabilities ≥ 0.95 respectively.
Sequences of Stereum hirsutum and Heterobasidion annosum were
used as outgroup taxa. The new isolates are in bold
Russulaceae Lotsy, Vortr. Bot. Stammesgesch. 1: 706
(1907)
Notes: Russulaceae is one of the important ectomycorrhizal groups of fungi which belongs in order Russulales (Kirk
et al. 2001). Historically, the family was established by Roze
in 1876 as Russulariéés (nom. nud). Late in the year of 1907
it was validated as Russulaceae by Lotsy and placed under
Agaricales. To our recent knowledge, Russulaceae includes
seven genera viz. Russula Pers., Lactarius Pers., Lactifluus
(Pers.) Roussel, Cystangium Singer & A.H. Sm., Multifurca
Buyck & V. Hofst., Boidinia Stalpers & Hjortstam and Pseudoxenasma K.H. Larss. & Hjortstam. There are records of
more than 1248 species belonging to the family across the
world (Atri et al. 2016).
Notes: Persoon (1796a, b) was the pioneer mycologist
who established the genus Russula with morphological features like the presence of fleshy fruitbodies with a depressed
cap, and equal gills. About 900 taxa have been reported in
this genus worldwide and 158 taxa have been reported from
India (Paloi et al. 2018).
Russula Pers., Observ. Mycol. (Lipsiae) 1: 100 (1796)
13
Russula benghalensis S. Paloi & K. Acharya, sp. nov.
MycoBank number: MB 831253; Facesoffungi Number: FoF
06141; Figs. 108, 109
Etymology: ‘benghalensis’ refers to the type locality West
Bengal.
Holotype: CUH AM654.
Basidiocarp small to medium. Pileus 14‒35 mm diam,
convex to broadly convex at early stage, becoming broadly
convex to applanate, or sometime upturned with a central
Fungal Diversity (2020) 104:1–266
Fig. 108 Russula benghalensis (CUH AM654, holotype). a Fresh
basidiocarps. b SEM photograph of basidiospores. Scale bars: a = 10
mm, b = 2 µm
depression, surface smooth when young, becoming cracked
or areolate, moist to semi-moist, gelatinized, brownish orange (7C5‒6), light brown (7D6) to reddish brown
(8D5) at center, towards margin greyish orange (5B6; 6B3),
brownish orange (6C4‒5) or greyish red (7B4), no change
after bruising, turns orange-red with KOH, orange-pink
with SV, light pink to pink with phenol and guaiacol; context very thin (< 1 mm), white (1A1) to cream, no change
when exposed to air. Lamellae ca. 2‒3 mm broad, adnate,
regular, white (1A1), turns brown-orange to brick red with
guaiacol and phenol, negative with SV; margins even, concolorous; lamellulae 1‒2 tiers. Stipe central, 20‒40 × 3‒6
mm, slightly curved, more or less equal, semi moist, ruinose;
greyish orange (5B3) to pale orange (6A3) or light orange
(6A4), no change on bruising, turning yellow with KOH,
negative reaction to phenol and guaiacol; context chambered, white (1A1), no change when exposed to air. Odour
unknown. Taste mild. Spore print white (1A1) to cream.
Basidiospores (5.5‒)6.3‒7.5(‒8.0) × (4.6‒)5.0‒6.5(‒7.5)
µm, L = 6.7, W = 5.8, Q = 1.07‒1.25, subglobose to broadly
ellipsoid, ornamentation amyloid, composed of short
(0.3‒0.4 µm) to high (0.6‒0.8 µm) isolated individual warts,
obtuse to sometime lobed apex, never reticulated. Basidium
35.5‒61.0 × 5.5‒11.5 µm, clavate to subclavate, hyaline,
thin-walled. Lamellar trama pseudoparanchymatus. Lamellae edge sterile. Hymenial cystidia 30.5‒68.5 × 6.0‒12.5
µm, sub-clavate to sub-cylindrical with moniliform to appendiculate apex, oil granules present in KOH near gill edge;
147
pleurocystidia: two types of cystidia present, one same as
gill edge, another 53‒70 × 9.0‒14.0 µm, clavate to subclavate with round or obtuse apex, thin-walled, hyaline, oil
granule present when viewed with KOH. Pileipellis a trichoderm, orthocromatic in Cresyl Blue, distinctly three layers,
underlining cortex composed of spherocytes; subpileus
50‒85 µm deep, composed of tightly arranged hyphal cells,
3.5‒5.5 µm broad, thin-walled, hyaline, end obtuse, oil granules present when mounted in KOH; suprapellis 30‒65 µm
deep, non-gelatinous, composed of erect to suberect hyphae,
3.0‒5.0 µm broad, thin-walled, hyaline, end obtuse or round;
oleiferous hyphae 3.5‒7.0 µm broad, abundant towards cap
centre, thin-walled; pileocystidia 27.0‒40.0 × 5.0‒7.0 µm,
scattered at margin, cylindrical to subcylindrical with less
pointed apex or sometimes rounded, thin-walled, hyaline,
oil granules present when mounted in KOH. Stipitipellis
composed of erect to suberect hyphae, 4.0‒6.0 µm broad,
non-gelatinous, thin-walled, hyaline, end obtuse; caulocystidia 43.0‒69.0 × 5.0‒7.0 µm, cylindrical with moniliform to
pointed apex, thin-walled, hyaline, oil granules present when
viewed with KOH. Clamp-connections absent in all tissues.
Habit and habitat: solitary, scattered, grows under Shorea
robusta tree.
Material examined: INDIA, West Bengal, Nadia district,
near Bethuradhari forest, 23°35′52.2492′′ N, 88°23′34.0944′′
E (elev. 25 m), 20 August 2016, S. Paloi and A. K. Dutta,
CUH AM654 (holotype); Nadia district, near Bethuradhari
forest, 23°35′52.1088″ N, 88°23′33.8424″ E (elev. 27 m), 5
August 2017, S. Paloi and K. Acharya, CUH AM655.
GenBank numbers: ITS: MK414583, MK414584; LSU:
MK496844.
Notes: Russula benghalensis is easily recognized by the
presence of a small-sized pileus, which turned orange-red
with KOH, adnate lamellae with one to two lamellulae, a
mild taste, basidiospores with ornamentation composed of
isolated warts, a trichodermal type of pileipellis, the presence of oleiferous hyphae and pileocystidia.
Phylogenetically related species are Russula pila (Pat.)
Trappe & T.F. Elliott and R. mistiformis (Mattir.) Trappe
& T.F. Elliott (Fig. 110); both species were reported from
Spain. However, R. pila differs from the new species due to
the presence of a subglobose fruitbody, larger basidiospores
(9‒13 µm, Q = 1) with warty ornamentation and absence of
cystidia (Vidal et al. 2019); R. mistiformis is associated with
Quercus, Pinus and Castanea; has a tuber-like brown fruitbody and larger basidiospores (8.5‒12.5 × 8‒10.5 µm) and
is easily separated from R. benghalensis (Vidal et al. 2019).
When considering species with similar basidiospore
size and ornamentation in the subgenus Ingratula, Russula
benghalensis appears closely related to R. lilacipes Shear,
R. granulata (Peck) Peck and R. amoenolens Romagn. However, R. lilacipes has a comparatively larger pileus (70‒150
mm diam), short, cylindrical cheilocystidia with ventricose
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Fungal Diversity (2020) 104:1–266
Fig. 109 Morphological features of Russula benghalensis (CUH AM654, holotype). a Basidia. b Cystidia near gill edge. c Cystidia near gill
side. d Pileipellis. e Oleiferous hyphae. f Pileocystidia. g Basidiospores. Scales bars: a = 10 mm, b–g = 10 µm, h = 5 µm
to fusiform apex and pileipellis covered by a gelatinous
layer (Shaffer 1972). R. granulata differs by a larger pileus
(40‒105 mm diam), a stipe surface turning reddish with
KOH, and pileocystidia with a fusiform, lanceolate or short
appendiculate apex (Shaffer 1972). R. amoenolens has a
strong acrid taste, a pileus surface that remains unchanged or
turns very faintly orange with KOH, a cutis type of pileipellis and an absence of pileocystidia (Romagnesi 1952).
Russula laurocerasi Melzer and R. senecis S. Imai
(reported from India) morphologically appear close to
R. benghalensis. R. laurocerasi differs from R. benghalensis by the presence of a dull yellow to yellow–brown
pileus, an acrid taste, comparatively larger basidiospores
(7.5‒11.0 × 7.5‒9 µm) with partial or nearly reticulate
ornamentation and a cutis type of pileipellis (Shaffer 1972).
R. senecis differs from the present taxon by the presence
of a discolored lamellae edge, a pileus surface which turns
rust to rusttawny with KOH, adnexed lamellae attachment,
13
a very acrid taste, strong fishy odour, larger-sized basidiospores with very high ornamentation (up to 3.2 µm tall)
(Khatua et al. 2015).
Thelephorales Corner ex Oberw.
Notes: The order was erected by Oberwinkler (1976),
based on Thelephoraceae ss. Stalpers. The Thelephorales is
composed of two families: Thelephoraceae and Bankeraceae
ss. Stalpers (Stalpers 1993; most species in this order are
considered to be ectomycorrhizal (Agerer 1987–2008, 1995;
Taylor and Bruns 1997; Erland and Taylor 1999; Kõljalg
et al. 2000). Currently around 18 genera and 269 species are
accepted in the order (Kirk et al. 2008).
Thelephoraceae Chevall.[as ’Thelephoreae’], Fl. Gén. Env.
Paris (Paris) 1: 84 (1826)
Notes: Thelephoraceae was introduced by Chevall (1826),
with the type genus Thelephora Ehrh. ex Willd. It is the
Fungal Diversity (2020) 104:1–266
149
Fig. 110 Phylogram generated from Bayesian analysis based on ITS
sequence dataset for 45 Russula and two outgroup sequences (Amylostereum laevigatum and Stereum hirsutum). Maximum likelihood
bootstrap support values ≥ 50% and Bayesian posterior probabilities ≥ 0.50 are indicated above the nodes. The newly described taxon
is indicated in bold
most important family in the Thelephorales, as many species
in the family play an important role forming ectomycorrhizal associations in forests from temperate to tropical zones
(Jakucs and Erős-Honti 2008; Peay et al. 2010; Tedersoo
et al. 2010; Gao et al. 2014). Currently around 12 genera and
171 species are accomodated in the family (Kirk et al. 2008).
Tomentella Pers. ex Pat., Hyménomyc. Eur. (Paris): 154
(1887)
Notes: The name “Tomentella” was first used as the subgenus of Corticium Pers. (Persoon 1799) and was validated
as the generic name by Patouillard (1887). Tomentella is
delimited from the other genera mostly by the lack of a special combinations of features, which are present in the other
genera (Agerer et al. 2001; Kõljalg 1996). It is characterized
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150
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Fig. 111 Phylogenetic position of Tomentella species inferred from the ITS sequences. Bootstrap support values for ML ≥ 50% and Bayesian
posterior probabilities ≥ 0.95 are given near nodes respectively. The tree is rooted with Odontia fibrosa (TU115714). The new species are in bold
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151
Fig. 111 (continued)
13
152
Fig. 112 A basidiocarp of Tomentella asiae-orientalis (IFP 019275,
holotype)
Fig. 113 SEM of basidiospores of Tomentella asiae-orientalis (IFP
019275, holotype)
by strictly resupinate basidiocarps separable from or adhered
to the substrates, a smooth or granulose hymenophore, the
presence or absence of rhizomorphs, subicular hyphae with
clamp connections or simple septa and ornamented basidiospores (Wakefield 1960; Larsen 1966, 1969, 1970; Kõljalg
1996). Fungi in this genus are cosmopolitan, and the species
diversity based on fruitbodies have been extensively taxonomically studied from temperate Eurasia (Kõljalg 1996),
North America (Larsen 1974), Iberian Peninsula (Melo et al.
2002), West Africa (Yorou and Agerer 2008; Yorou et al.
2007, 2008, 2011), Australia (Agerer and Bougher 2001),
South America (Kuhar et al. 2016) and Asia (Thind and
Rattan 1971). In addition, using high throughput sequencing
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techniques a large number of DNA sequences from environmental samples or plant ectomycorrhizal root tips from
different regions of the world have been identified as Tomentella spp. with most of them not matching any known species (Gao et al. 2013; Nouhra et al. 2013). These sequences
provide distribution information despite lacking a visible
fruitbody. About 400 names have been recorded and around
150 species accepted in the database of Index Fungorum;
80 species were recorded in the Dictionary of the Fungi
(10th ed.) (Kirk et al. 2008). However, taxonomic studies
based on fruitbodies of Tomentella in China were very few.
Based on morphological characteristics and phylogenetic
analyses of rDNA ITS sequences (Fig. 111), new species of
Tomentella are described and illustrated in this paper. The
types (type A–F) of rhizomorphic hyphal organization are
important charateristics; we follow the definition of Agerer
(1987–2008).
Tomentella asiae-orientalis H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555723; Facesoffungi number:
FoF 05601; Figs. 112, 113, 114
Etymology: Refers to the distribution in East Asia.
Holotype: IFP 019275.
Basidiocarps annual, resupinate, adherent to the substrate,
arachnoid, without odour or taste when fresh, 0.5–0.8 mm
thick, continuous. Hymenophoral surface smooth, pale
brown to dark brown (6D8–6F7) and concolorous with subiculum when dry. Sterile margin often indeterminate, byssoid, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped,
thick-walled, frequently branched, 5–7 μm diam, without
encrustation, pale brown in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped, thin-walled, occasionally
branched, 7–9 μm diam; hyphal cells more or less uniform,
pale brown in KOH, acyanophilous, inamyloid. Cystidia
absent. Basidia 15–45 μm long and 5–13 μm diam at apex,
4–6 μm at base, with a clamp connection at base, clavate,
stalked, sinuous, without transverse septa, pale brown in
KOH and in distilled water, 4-sterigmate; sterigmata 6–8
μm long and 1–1.5 μm diam at base. Basidiospores thickwalled, (7–)7.5–8.5(–9) × (6–)6.5–7.5(–8) μm, L = 8.12 μm,
W = 7.08 μm, Q = 1.08–1.2 (n = 60/2), globose to subglobose
in frontal and lateral views, echinulate, pale brown in KOH
and distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 2.5 μm
long.
Material examined: CHINA, Heilongjiang Province,
Fenglin National Nature Reserve, on fallen angiosperm twig,
16 October 2016, Yuan 12022 (IFP 019275, holotype); on
fallen branch of Pinus koraiensis, 16 October 2016, Yuan
11918 (IFP 019276).
Fungal Diversity (2020) 104:1–266
153
Fig. 114 Microscopic structures
of Tomentella asiae-orientalis
(IFP 019275, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
GenBank numbers: ITS: MK211711, MK211710; LSU:
MK446334, MK446333.
Notes: Tomentella bryophila (Pers.) M.J. Larsen is similar to T. asiae-orientalis by its basidiocarps adherent to the
substrate, smooth hymenophoral surface, clamped hyphae,
globose to subglobose basidiospores and the absence of rhizomorphs and cystidia. However, T. bryophila differs from
T. asiae-orientalis by having short and inflated subhymenial hyphal cells, utriform basidia and encrusted subicular hyphae (Kõljalg 1996). T. changbaiensis resembles T.
asiae-orientalis by light brown to dark brown basidiocarps
adherent to the substrate, smooth hymenophoral surface,
clamped hyphae, globose to subglobose basidiospores
and the absence of rhizomorphs and cystidia. However, T.
changbaiensis differs from T. asiae-orientalis by having
short and inflated subhymenial hyphal cells and utriform
basidia.
Tomentella atrobadia H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555627; Facesoffungi number: FoF 05602; Figs. 115, 116, 117
Etymology: Refers to the dark brown basidiocarps.
Holotype: IFP 019243.
Basidiocarps annual, resupinate, adherent to the substrate, crustose, without odour or taste when fresh, 0.5–0.8
mm thick, continuous. Hymenophoral surface smooth, dark
brown (6F5–7F5) and paler than subiculum when dry. Sterile margin often indeterminate, byssoid, concolorous with
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154
Fig. 115 A basidiocarp of Tomentella atrobadia (IFP 019243, holotype)
Fig. 116 SEM of basidiospores of Tomentella atrobadia (IFP
019243, holotype)
hymenophore. Subiculum mostly dark brown. Rhizomorphs
absent. Subicular hyphae monomitic, generative hyphae
clamped, thick-walled, rarely branched, 6–7 μm diam, without encrustation, yellow in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped, thick-walled, frequently
branched, 6–8 μm diam; hyphal cells short and inflated,
yellow in KOH, cyanophilous, inamyloid. Cystidia absent.
Basidia 40–60 μm long and 8–15 μm diam at apex, 5–8 μm
at base, with a clamp connection at base, utriform, stalked,
not sinuous, rarely with transverse septa, yellow in KOH,
yellow in distilled water, 4-sterigmate; sterigmata 5–7 μm
long and 3–4 μm diam at base. Basidiospores thick-walled,
(9–)9.4–10.5(–11) × (7.5–)8–9.9(–10.5) μm in lateral and
frontal views, L = 9.87 μm, W = 8.8 μm, Q = 1.07–1.15
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(n = 60/2), globose, subglobose, triangular or lobed in frontal view and subglobose to ellipsoid in lateral view, echinulate to aculeolate, yellow in KOH, yellow in distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 1.5 μm long.
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on rotten angiosperm branch, 2 August 2016, Yuan 11099 (IFP 019243,
holotype); on rotten angiosperm wood debris, 2 August
2016, Yuan 11114 (IFP 019244).
GenBank numbers: ITS: KY686248, KY686249; LSU:
MK446335, MK446336.
Notes: Tomentella stuposa (Link) Stalpers is similar to T.
atrobadia by having continuous and brown-coloured basidiocarps adherent to the substrate, a smooth hymenophoral
surface, thick-walled hyphae, globose basidiospores of
approximately the same size and the absence of rhizomorphs. However, T. stuposa is differentiated by having
simple septate subicular hyphae, thin-walled subhymenial
hyphae and triangular or lobed basidiospores (Kõljalg 1996).
T. alpina Peintner & Dämmrich is similar to T. atrobadia by
having continuous basidiocarps adherent to the substrate,
smooth hymenophore, the absence of rhizomorphs, clamped
and thick-walled subicular hyphae and globose or subglobose basidiospores. But, the former species has distinctly
smaller basidospores (6.5–8.5 μm diam, Peintner and Dämmrich 2012).
Tomentella atrocastanea H.S. Yuan, X. Lu & Y.C. Dai, sp.
nov.
Index Fungorum number: IF555628; Facesoffungi number: FoF 05603; Figs. 118, 119, 120
Etymology: Refers to the dark chestnut basidiocarps.
Holotype: IFP 019245.
Basidiocarps annual, resupinate, separable from the
substrate, mucedinoid, without odour or taste when fresh,
0.2–0.6 mm thick, continuous. Hymenophoral surface
smooth, dark chestnut (6F7) and concolorous with subiculum when dry. Sterile margin often determinate, farinaceous,
concolorous with hymenophore. Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped and
rarely simple septate, thick-walled, occasionally branched,
5–7 μm diam, without encrustation, pale brown in KOH,
acyanophilous, inamyloid. Subhymenial hyphae clamped and
rarely simple septate, thin-walled, occasionally branched,
7–9 μm diam; hyphal cells more or less uniform, pale brown
in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
10–40 μm long and 5–10 μm diam at apex, 5–10 μm at base,
with a clamp connection at base, utriform, not stalked, sinuous, rarely with transverse septa, pale brown in KOH and in
distilled water, 4-sterigmate; sterigmata 4–6 μm long and
2–2.5 μm diam at base. Basidiospores thick-walled, (7–)7
.5–8.5(–9) × (6–)6.5–8(–8.5) μm, L = 7.96 μm, W = 7.28 μm,
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155
Fig. 117 Microscopic structures
of Tomentella atrobadia (IFP
019243, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
Q = 1.07–1.13 (n = 60/2), globose to subglobose in frontal
and lateral views, echinulate, pale brown in KOH and distilled water, cyanophilous, inamyloid; echinuli usually isolated, up to 3 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 31 July 2017, Yuan 12179 (IFP
019245, holotype), Yuan 12170 (IFP 019246).
GenBank numbers: ITS: MK211743, MK211742; LSU:
MK446338, MK446337.
Notes: Tomentella tenuirhizomorpha X. Lu et al. is similar to T. atrocastanea by having mucedinoid and continuous
basidiocarps separable from the substrate, a smooth hymenophoral surface, a farinaceous sterile margin, clamped and
simple septate hyphae, more or less uniform subhymenial
hyphal cells, the absence of cystidia, and globose to subglobose basidiospores. However, T. tenuirhizomorpha differs from T. atrocastanea by having thin-walled subicular
hyphae, clavate basidia and slender rhizomorphs (Lu et al.
2018a). T. agereri Yorou also resembles T. atrocastanea by
having continuous basidiocarps separable from the substrate,
more or less uniform subhymenial hyphal cells, the absence
of cystidia, and globose to subglobose basidiospores. But, it
differs by fulvous or sienna, arachnoid basidiocarps with a
13
156
Fig. 118 A basidiocarp of Tomentella atrocastanea (IFP 019245,
holotype)
Fig. 119 SEM of basidiospores of Tomentella atrocastanea (IFP
019245, holotype)
byssoid sterile margin, clamped hyphae, clavate basidia and
the presence of rhizomorphs (Yorou et al. 2011).
Tomentella aureomarginata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555629; Facesoffungi number: FoF 05604; Figs. 121, 122, 123
Etymology: Refers to the golden yellow sterile margin of
the basidiocarps.
Holotype: IFP 019247.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh, 1–1.5
mm thick, continuous. Hymenophoral surface smooth,
golden brown to yellowish brown (5D7–5F5) and turning
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darker than subiculum when dry. Sterile margin often determinate, byssoid, ligher than hymenophore, golden brown
(5D7). Rhizomorphs present in subiculum and margins,
15–35 μm diam; rhizomorphic surface more or less smooth;
hyphae in rhizomorphs monomitic, undifferentiated, of type
B, compactly arranged and uniform; single hyphae with
both clamps and simple septa, thick-walled, unbranched,
3.5–4 μm, pale brown in KOH, cyanophilous, inamyloid.
Subicular hyphae monomitic; generative hyphae clamped
and rarely simple septate, slightly thick- to thick-walled,
frequently branched, 2–4 μm diam, without encrustation,
pale brown in KOH, cyanophilous, inamyloid. Subhymenial
hyphae clamped and simple septate, thin-walled, frequently
branched, 3.5–4 μm diam, without encrustation; hyphal cells
short and not inflated, pale brown in KOH, acyanophilous,
inamyloid. Cystidia absent. Basidia 20–45 μm long and 4–8
μm diam at apex, 3–4 μm at base, with a clamp connection at base, utriform, not stalked, sinuous, with transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 2–7 μm long and 1.5–2 μm diam at base.
Basidiospores slightly thick-walled, (6–)6.5–7(–7.5) × (5.5–
)6–6.5(–7) μm, L = 6.72 μm, W = 6.26 μm, Q = 1.06–1.22
(n = 60/2), subglobose to bi- or tri-lobed in frontal and lateral views, echinulate, pale brown in KOH and distilled
water, cyanophilous, inamyloid; echinuli usually isolated,
sometimes grouped in 2 or more, up to 1 μm long.
Material examined: CHINA, Liaoning Province, Huanren County, Laotudingzi National Nature Reserve, on rotten
angiosperm wood debris, 21 October 2015, Yuan 10671 (IFP
019247, holotype), Yuan 10683 (IFP 019248).
GenBank numbers: ITS: MK211744, MK211745; LSU:
MK446339, MK878395.
Notes: Tomentella botryoides (Schwein.) Bourdot & Galzin has characteristics similar to T. aureomarginata, such as
pelliculose and continuous basidiocarps, a byssoid sterile
margin, the absence of cystidia, the presence of rhizomorphs
and subglobose to bi- or tri-lobed basidiospores. However, T.
botryoides differs from T. aureomarginata by having fuscous
black, mouse grey or greyish sepia basidiocarps, more or
less uniform subhymenial hyphal cells and clavate basidia
(Kõljalg 1996; Daemmrich 2006). T. separata resembles
T. aureomarginata by having continuous basidiocarps, a
smooth hymenophoral surface, a byssoid sterile margin,
clamped and simple septate hyphae, short and inflated subhymenial hyphal cells, the presence of rhizomorphs, the
absence of cystidia, and slightly thick-walled basidiospores.
But T. separata differs by the honey yellow to raw umber
basidiocarps, clavate basidia and subglobose to bi-, tri- or
quadra-lobed basidiospores.
Tomentella brevis H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555630; Facesoffungi number: FoF 05605; Figs. 124,125, 126
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157
Fig. 120 Microscopic structures
of Tomentella atrocastanea
(IFP 019245, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
Fig. 121 A basidiocarp of Tomentella aureomarginata (IFP 019247,
holotype)
Etymology: Refers to the short sterigmata of the basidia.
Holotype: IFP 019249.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh, 0.5–1
mm thick, discontinuous. Hymenophoral surface granulose,
brownish beige to greyish brown (6E3–6F3) and concolorous
with subiculum when dry. Sterile margin often determinate,
farinaceous, concolorous with hymenophore. Rhizomorphs
present in subiculum and margins, 25–40 μm diam; rhizomorphic surface more or less smooth; hyphae in rhizomorph
monomitic, differentiated, of type C; central hyphae with
both clamps and simple septa, somewhat enlarged, thickwalled, unbranched, 3–4 μm diam; hyphae outer part of rhizomorphs with both clamps and simple septa, slightly thickwalled, unbranched, 2–2.5 μm diam; pale-brown in KOH,
acyanophilous, inamyloid. Subicular hyphae monomitic;
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having continuous basidiocarps adherent to the substrate,
clamped hyphae, clavate basidia and the absence of cystidia.
But, the former species is differentiated by brick or umber
basidiocarps with a granulose hymenophoral surface, subglobose to bi-lobed basidiospores and the absence of rhizomorphs (Kõljalg 1996; Daemmrich 2006; Melo et al. 2006).
Fig. 122 SEM of basidiospores of Tomentella aureomarginata (IFP
019247, holotype)
generative hyphae clamped, thick-walled, occasionally
branched, 3–5 μm diam, without encrustation, pale brown
in KOH, cyanophilous, inamyloid. Subhymenial hyphae
clamped, thin- to slightly thick-walled, rarely branched,
3–6 μm diam; hyphal cells short and inflated, pale brown in
KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
10–30 μm long and 3–5.5 μm diam at apex, 2.5–3 μm at
base, with clamp connection at base, clavate, not stalked,
sinuous, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 2–3.5 μm and 1–1.5 μm diam at base.
Basidiospores slightly thick-walled, (4.5–)5–6.5(–7) × (4–)
4.5–5.5(–6) μm, L = 5.93 μm, W = 4.83 μm, Q = 1.17–1.23
(n = 60/2), subglobose to bi-, tri- or quadra-lobed in frontal and lateral views, echinulate, pale brown in KOH and
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 1 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 7 August
2016, Yuan 11328 (IFP 019249, holotype); on rotten angiosperm stump, 7 August 2016, Yuan 11332 (IFP 019250).
GenBank numbers: ITS: MK211746, MK211747; LSU:
MK446340, MK878396.
Notes: The fuscous black, mouse grey or greyish sepia,
continuous basidiocarps of Tomentella botryoides are similar to T. brevis. In addition, they also share some other characteristics: clamped hyphae, clavate basidia, the presence of
rhizomorphs and the absence of cystidia. However, T. botryoides differs from T. brevis by having pelliculose basidiocarps separable from the substrate, a byssoid sterile margin
and subglobose to bi-, tri-lobed basidiospores (Kõljalg 1996;
Daemmrich 2006). T. lateritia Pat. resembles T. brevis by
13
Tomentella brunneoflava H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555631; Facesoffungi number: FoF 05606; Figs. 127, 128, 129
Etymology: Refers to the brownish yellow hymenophoral
surface.
Holotype: IFP 019251.
Basidiocarps annual, resupinate, adherent to the substrate, arachnoid, without odour or taste when fresh, 0.8–1.5
mm thick, continuous. Hymenophoral surface smooth,
brownish yellow (5C8) and concolorous with subiculum
when dry. Sterile margin often indeterminate, byssoid, concolorous with hymenophore. Rhizomorphs present in subiculum and margins, 5–35 μm diam; rhizomorphic surface more
or less smooth; hyphae in rhizomorph monomitic, undifferentiated, of type B, compactly arranged and of uniform; single hyphae with both clamps and simple septa, thick-walled,
unbranched, 2–4 μm, pale brown in KOH and in distilled
water, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae clamped and rarely simple septate,
thin- to slightly thick-walled, occasionally branched, 2–5
μm diam, without encrustation, pale brown in KOH, acyanophilous, inamyloid. Subhymenial hyphae clamped and
rarely simple septate, thin-walled, occasionally branched,
3–5 μm diam; hyphal cells short and not inflated, pale brown
in KOH, inamyloid. Cystidia absent. Basidia 10–30 μm
long and 3–5 μm diam at apex, 2–3.5 μm at base, with a
clamp connection at base, clavate, not stalked, sinuous, with
transverse septa, pale brown in KOH and in distilled water,
4-sterigmate; sterigmata 3–5 μm long and 1–1.5 μm diam
at base. Basidiospores slightly thick-walled, (6.5–)7–8(–8.
5) × (5–)5.5–7(–7.5) μm, L = 7.78 μm, W = 6.29 μm, Q = 1.
19-1.24 (n = 60/2), subglobose to bi- or tri-lobed in frontal
and lateral views, echinulate, pale brown in KOH and distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 1.5 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 31 July 2017, Yuan 12162 (IFP
019251, holotype); on fallen branch of Larix sp., 31 July
2017, Yuan 12161 (IFP 019252).
GenBank numbers: ITS: MK211749, MK211748; LSU:
MK850194, MK446341.
Notes: Tomentella botryoides has characteristics similar to T. brunneoflava, such as continuous basidiocarps,
a byssoid sterile margin, clavate basidia and the presence
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159
Fig. 123 Microscopic structures
of Tomentella aureomarginata
(IFP 019247, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
of rhizomorphs. However, T. botryoides differs from T.
brunneoflava by having fuscous black, mouse grey or
greyish sepia and pelliculose basidiocarps, more or less
uniform subhymenial hyphal cells and subglobose to
bi-, tri-lobed basidiospores (Kõljalg 1996; Daemmrich
2006). T. aureomarginata resembles T. brunneoflava by
having continuous basidiocarps adherent to the substrate,
a smooth hymenophoral surface, a byssoid sterile margin, the presence of rhizomorphs, the absence of cystidia,
clamped and rarely simple septate hyphae and short subhymenial hyphal cells. T. aureomarginata differs by the
honey yellow to raw umber colour of the basidiocarps and
utriform basidia.
Tomentella brunneogrisea H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Fig. 124 A basidiocarp of Tomentella brevis (IFP 019249, holotype)
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160
Fig. 125 SEM of basidiospores of Tomentella brevis (IFP 019249,
holotype)
Fig. 126 Microscopic structures of Tomentella brevis (IFP
019249, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Index Fungorum number: IF555632; Facesoffungi number: FoF 05607; Figs. 130, 131 132
Etymology: Refers to the brownish grey basidiocarps.
Holotype: IFP 019253.
Basidiocarps annual, resupinate, adherent to the substrate,
mucedinoid, without odour or taste when fresh, 0.8–1.5 mm
thick, continuous. Hymenophoral surface granulose, brownish grey to dark brown (7E2–7F4) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped
and rarely simple septate, thick-walled, frequently branched,
4–9 μm diam, without encrustation, pale brown in KOH,
cyanophilous, inamyloid. Subhymenial hyphae clamped and
rarely simple septate, thin-walled, occasionally branched,
4–7 μm diam; hyphal cells short and not inflated, pale brown
in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
10–50 μm long and 6.5–10 μm diam at apex, 4–6 μm at base,
with a clamp connection at base, utriform, not stalked, sinuous, without transverse septa, pale brown in KOH and in
distilled water, 4-sterigmate; sterigmata 8–10 μm long and
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161
adherent to the substrate, short subhymenial hyphal cells,
utriform basidia and the absence of rhizomorphs and cystidia. However, T. atrobadia differs from T. brunneogrisea
by having crustose basidiocarps with a byssoid sterile margin, and thick-walled subhymenial hyphae. T. pertenuis and
T. brunneogrisea share the following similar morphological and anatomical characteristics: mucedinoid basidiocarps
adherent to the substrate, farinaceous and indeterminate sterile margin, utriform basidia, thick-walled basidiospores and
the absence of rhizomorphs and cystidia. But, T. pertenuis
has yellowish brown basidiocarps and slightly thick-walled
subhymenial hyphae.
Fig. 127 Basidiocarps of Tomentella brunneoflava (IFP 019251,
holotype)
Fig. 128 SEM of basidiospores of Tomentella brunneoflava (IFP
019251, holotype)
2–3 μm diam at base. Basidiospores thick-walled, (7–)8–
9(–9.5) × (6.5–)7.5–8.5(–9) μm, L = 8.47 μm, W = 7.97 μm,
Q = 1.04–1.08 (n = 60/2), globose to subglobose in frontal
and lateral views, echinulate, pale brown in KOH and in
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 2 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 30 July 2017, Yuan 12147 (IFP
019253, holotype), Yuan 12146 (IFP 019254).
GenBank numbers: ITS: MK211751, MK211750; LSU:
MK446343, MK446342.
Notes: Tomentella atrobadia is similar to T. brunneogrisea by the brownish grey to dark brown basidiocarps
Tomentella capitatocystidiata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555724; Facesoffungi number: FoF 05608; Figs. 133, 134, 135
Etymology: Refers to the capitate cystidia.
Holotype: IFP 019255.
Basidiocarps annual, resupinate, separable from the substrate, arachnoid, without odour or taste when fresh, 0.8–1.2
mm thick, continuous. Hymenophoral surface smooth, dark
blonde to yellowish brown (5D4–5E8) and turning lighter
than the subiculum when dry. Sterile margin often indeterminate, byssoid, ligher than hymenophore, dark blonde
(5D4). Rhizomorphs present in subiculum and margins, 5–30
μm diam; rhizomorphic surface more or less smooth; hyphae
in rhizomorphs monomitic, undifferentiated, of type B, compactly arranged and of uniform; single hyphae with both
clamps and simple septa, thick-walled, unbranched, 2–4 μm
diam, pale yellow in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae clamped and
rarely simple septate, thick-walled, occasionally branched,
3–6 μm diam, without encrustation, pale brown in KOH,
cyanophilous, inamyloid. Subhymenial hyphae clamped and
rarely simple septate, thin-walled, occasionally branched,
5–8 μm diam; hyphal cells short and inflated, pale brown
in KOH, acyanophilous, inamyloid. Cystidia present, capitate, arising from subhymenial hyphae, 55–100 μm long,
3–5 μm diam at base and 12–16 μm diam at apex, projecting outside of the hymenium, clavate, without simple septa
and encrustation, pale yellow in KOH. Basidia 15–40 μm
long and 5–10 μm diam at apex, 3–5.5 μm at base, with
a clamp connection at base, clavate, not stalked, sinuous,
without transverse septa, pale yellow in KOH and in distilled
water, 4-sterigmate; sterigmata 3–3.5 μm long and 1.5–2 μm
diam at base. Basidiospores slightly thick-walled, (6.5–)7–
7.5(–8) × (6–)6.5–7(–7.5) μm, L = 7.38 μm, W = 6.56 μm,
Q = 1.09–1.19 (n = 60/2), subglobose to bi-, tri- or quadralobed in frontal and lateral views, echinulate, pale yellow in
KOH and distilled water, cyanophilous, inamyloid; echinuli
usually isolated, sometimes grouped in 2 or more, up to 1.5
μm long.
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Fig. 129 Microscopic structures
of Tomentella brunneoflava
(IFP 019251, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
Fig. 130 A basidiocarp of Tomentella brunneogrisea (IFP 019253,
holotype)
13
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 10 August
2016, Yuan 11459 (IFP 019255, holotype), Yuan 11494
(IFP 019256).
GenBank numbers: ITS: MK211700, MK211701; LSU:
MK446344, MK446345.
Notes: Tomentella pilosa (Burt) Bourdot & Galzin and
T. capitatocystidiata formed a clade in our phylogeny
(Fig. 111) and both species share these characteristics:
arachnoid basidiocarps separable from the substrate with
a byssoid sterile margin, the presence of capitate cystidia
and rhizomorphs. However, T. pilosa differs from T. capitatocystidiata by having utriform basidia, rhizomorphs with
a dimitic hyphal system and subglobose to bi-, tri-lobed
basidiospores (Wakefield 1960; Kõljalg 1996). T. capitata
Yorou & Agerer and T. capitatocystidiata also share arachnoid basidiocarps separable from the substrate, a smooth
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163
hymenophoral surface, a byssoid sterile margin, clamped
and simple septate hyphae, clavate basidia and the presence
of rhizomorphs and capitate cystidia. But, the former species
is differentiated by brown to dark brown basidiocarps, more
or less uniform subhymenial hyphal cells and subglobose to
bi-, tri-lobed basidiospores (Yorou et al. 2007).
at base. Basidiospores thick-walled, (7.5–)8–9(–9.5) × (7–)
7.5–8.5(–9) μm, L = 8.45 μm, W = 7.83 μm, Q = 1.06–1.12
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 1.5 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 10 August
2016, Yuan 11496 (IFP 019257, holotype), Yuan 11477
(IFP 019258).
GenBank numbers: ITS: MK211739, MK211738; LSU:
MK446347, MK446346.
Notes: Tomentella bryophila and T. changbaiensis formed
a clade in the phylogeny (Fig. 111). The former is similar
to T. changbaiensis by having basidiocarps adherent to the
substrate, the absence of rhizomorphs and cystidia, short and
inflated subhymenial hyphal cells, utriform basidia and globose to subglobose basidiospores (Kõljalg 1996). However,
T. bryophila differs from T. changbaiensis by having dark
blonde to raw umber basidiocarps and encrusted subicular
hyphae. T. pallidobrunnea resembles T. changbaiensis by
pale brown to dark brown, mucedinoid or arachnoid and
continuous basidiocarps adherent to the substrate, a byssoid
sterile margin, short and inflated subhymenial hyphal cells,
the absence of rhizomorphs and cystidia, and globose to
subglobose basidiospores. However, T. pallidobrunnea can
be distinguished by the pale brown sterile margin and clavate
basidia.
Tomentella changbaiensis H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555725; Facesoffungi number: FoF 05609; Figs. 136, 137, 138
Etymology: Refers to the location of the collection of the
type specimen.
Holotype: IFP 019257.
Basidiocarps annual, resupinate, duplex, easily separable
from the substrate, arachnoid, without odour or taste when
fresh, 0.5–1 mm thick, continuous. Hymenophoral surface
smooth, pale brown to dark brown (6D8–6F7) and turning
lighter than subiculum when dry. Sterile margin often indeterminate, byssoid, lighter than hymenophore, pale brown
(6D8). Rhizomorphs absent. Subicular hyphae monomitic;
generative hyphae clamped, thick-walled, occasionally
branched, 3–6 μm diam, without encrustation, pale brown
in KOH, cyanophilous, inamyloid. Subhymenial hyphae
clamped, thin-walled, occasionally branched, 4–6 μm diam;
hyphal cells short and inflated, pale brown in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia 20–55 μm long
and 5–11 μm diam at apex, 3–5.5 μm at base, with a clamp
connection at base, utriform, not stalked, sinuous, without
transverse septa, pale brown in KOH and in distilled water,
4-sterigmate; sterigmata 4–5.5 μm long and 2.5–3 μm diam
Tomentella citrinocystidiata H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555692; Facesoffungi number: FoF 05610; Figs. 139, 140, 141
Etymology: Refers to the pale orange cystidia in KOH.
Holotype: IFP 019331.
Basidiocarps annual, resupinate, separable from the
substrate, pelliculose, without odour or taste when fresh,
0.1–0.2 mm thick, continuous. Hymenophoral surface
smooth, dark brown (6F5–7) when dry. Sterile margin often
indeterminate, byssoid, concolorous with subiculum, more
or less yellowish brown. Subiculum mostly yellowish brown,
paler than hymenophore. Rhizomorphs present in subiculum
and margins, 15–30 μm diam; rhizomorphic surface more
or less smooth; hyphae in rhizomorph monomitic, differentiated, of type C, compactly arranged; hyphae in center of
rhizomorph usually clamped, rarely simple septate, thickwalled, unbranched, 3–3.5 μm diam; hyphae at outer part of
rhizomorph clamped, thick-walled, unbranched, 2–2.5 μm
diam; all the hyphae pale orange in KOH, cyanophilous,
inamyloid. Subicular hyphae monomitic; generative hyphae
mostly clamped and rarely simple septate, thick-walled,
rarely branched, 3–5 μm diam, occasionally collapsed and
encrusted, pale orange in KOH, cyanophilous, inamyloid.
Subhymenial hyphae thin- to slightly thick-walled, clamped,
Fig. 131 SEM of basidiospores of Tomentella brunneogrisea (IFP
019253, holotype)
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164
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Fig. 132 Microscopic structures
of Tomentella brunneogrisea
(IFP 019253, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
frequently branched, 3–5 μm diam; hyphal cells short and
inflated, pale orange in KOH, cyanophilous, inamyloid. Cystidia arising from subhymenial or from subicular hyphae,
60–80 μm long, 5–7.5 μm diam at base and 10–15 μm diam
at apex, capitate, embedded, without encrustation, pale
orange in KOH. Basidia 30–70 μm long and 5–10 μm diam
at apex, 5–7 μm at base, with a clamp connection at base,
clavate, stalked, sinuous, without transverse septa, pale
orange in KOH, yellowish brown in distilled water, 4-sterigmate; sterigmata 4–7 μm long and 1–2 μm diam at base.
Basidiospores thick-walled, (7–)7.4–8.5(–9) × (6.5–)7.1–
7.9(–8.5) μm, L = 7.92 μm, W = 7.41 μm, Q = 1.07–1.12
(n = 60/2), irregular globose or lobed in frontal view and
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ellipsoid in lateral view, echinulate to aculeate, pale brown
in KOH, pale brown in distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2
or more, up to 1.5 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
wood debris, 21 October 2015, Yuan 10680 (IFP 019331,
holotype); Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 22 October
2015, Yuan 10743 (IFP 019332).
GenBank numbers: ITS: KY686246, KY686247; LSU:
MK446348, MK446349.
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hyphae, no encrustated subicular hyphae, and basidiospores
bearing long echinuli.
Fig. 133 A basidiocarp of Tomentella capitatocystidiata (IFP
019255, holotype)
Fig. 134 SEM of basidiospores of Tomentella capitatocystidiata (IFP
019255, holotype)
Notes: Tomentella capitata, T. cystidiata and T. citrinocystidiata share similar morphological and anatomical characteristics: brown to dark brown, continuous basidiocarps
separable from the substrate, a smooth hymenophore, a byssoid sterile margin, the presence of rhizomorphs and capitate cystidia, thin- to thick-walled subhymenial hyphae and
basidiospores of approximately the same shape and size.
However, T. capitata is differentiated from T. citrinocystidiata by having a granulose or farinose hymenophore and the
presence of cystidia from the rhizomorphic surface (Yorou
et al. 2007); T. cystidiata differs from T. citrinocystidiata
by having arachnoid basidiocarps, a uniform subhymenial
Tomentella coffeae H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555693; Facesoffungi number: FoF 05611; Figs. 142, 143 144
Etymology: Refers to the coffee coloured basidiocarps.
Holotype: IFP 019259.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.1–0.3 mm thick, continuous. Hymenophoral surface
smooth, dark brown (7F5–7) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
with both clamps and simple septa, thick-walled, occasionally branched, 4–7 μm diam, without encrustation, deep yellow in KOH, slightly cyanophilous, inamyloid. Subhymenial
hyphae clamped, thick-walled, frequently branched; hyphal
cells short and inflated, 5–9 μm diam, deep yellow in KOH,
cyanophilous, inamyloid. Cystidia absent. Basidia 30–50
μm long and 5–10 μm diam at apex, 3–5 μm at base, with
a clamp connection at the base, clavate, stalked, sinuous,
occasionally with transverse septa, deep yellow in KOH,
deep yellow in distilled water, 4-sterigmate; sterigmata 3–5
μm long and 1–2 μm diam at base. Basidiospores thin- to
slightly thick-walled, (6.5–)7–9(–9.5) × (6.5–)6.8–8.3(–8.5)
μm, L = 8.37 μm, W = 7.62 μm, Q = 1.09–1.26 (n = 60/2),
globose in frontal view and subglobose in lateral view, aculeate, yellowish orange in KOH, pale yellow in distilled
water, cyanophilous, inamyloid; echinuli usually isolated,
sometimes grouped in 2, up to 2 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
wood debris, 20 October 2015, Yuan 10629 (IFP 019259,
holotype); Xifeng County, Binglashan National Forest Park,
on fallen angiosperm branch, 2 August 2016, Yuan 11100
(IFP 019260); Qingyuan County, Experimental Station of
Forest Ecology, on fallen angiosperm branch, 23 October
2016, Yuan 10762.
GenBank numbers: ITS: KY686255, KY686254; LSU:
MK446350, MK446351.
Notes: Tomentella intsiae Suvi & Kõljalg resembles T.
coffeae by having continuous basidiocarps adherent to the
substrate, a smooth hymonophoral surface, the absence of
rhizomorphs, thick-walled subicular hyphae and globose,
echinulate basidiospores with long echinuli. However, the
former species is differentiated by the absence of simple septate subicular hyphae and bigger basidiospores (9.5–11 μm,
Suvi et al 2010). T. lapida (Pers.) Stalpers is similar to T.
coffeae by having continuous and mucedinoid basidiocarps
adherent to the substrate, indeterminate sterile margins, the
absence of rhizomorphs, and slightly globose basidiospores
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Fig. 135 Microscopic structures
of Tomentella capitatocystidiata (IFP 019255, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
of approximately the same size. But T. lapida differs from T.
coffeae by its smoke grey basidiocarps, clamped subicular
hyphae and thin-walled subhymenial hyphae (Kõljalg 1996).
Tomentella conclusa H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
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Index Fungorum number: IF555735; Facesoffungi number: FoF 05612; Figs. 145, 146, 147
Etymology: Refers to the spores having crowded echinuli.
Holotype: IFP 019263.
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Fig. 136 A basidiocarp of Tomentella changbaiensis (IFP 019257,
holotype)
Fig. 137 SEM of basidiospores of Tomentella changbaiensis (IFP
019257, holotype)
Basidiocarps annual, resupinate, duplex, adherent to the
substrate, mucedinoid, without odour or taste when fresh,
0.3–0.6 mm thick, continuous. Hymenophoral surface
smooth, pale brown to brown (6D4–6E5) and concolorous
with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent. Subicular hyphae monomitic; generative
hyphae clamped, thick-walled, occasionally branched, 5–7
μm diam, without encrustation, pale brown in KOH, acyanophilous, inamyloid. Subhymenial hyphae clamped, thinwalled, occasionally branched, 6–9 μm diam; hyphal cells
short and inflated, pale brown in KOH, cyanophilous, inamyloid. Cystidia absent. Basidia 15–55 μm long and 7–12
167
μm diam at apex, 5–8 μm at base, with a clamp connection at
base, utriform, not stalked, sinuous, without transverse septa,
pale brown in KOH, 4-sterigmate; sterigmata 10–15 μm long
and 2–3 μm diam at base. Basidiospores thick-walled, (8.5–
)9–10(–10.5) × (7.5–)8–9(–10) μm, L = 9.46 μm, W = 8.57
μm, Q = 1.06–1.14 (n = 60/2), globose to subglobose in frontal and lateral views, echinulate, pale brown in KOH and
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, up to 2 μm long.
Material examined: CHINA, Heilongjiang Province,
Fenglin National Nature Reserve, on fallen angiosperm
branch, 17 October 2016, Yuan 12086 (IFP 019263, holotype); on fallen trunk of Pinus koraiensis, 16 October 2016,
Yuan 11986 (IFP 019264).
GenBank numbers: ITS: MK211703, MK211702; LSU:
MK446352, MK850195.
Notes: Tomentella longiechinuli and T. conclusa formed a
clade in the phylogeny (Fig. 111). They share similar characteristics: continuous basidiocarps adherent to the substrate, a
farinaceous sterile margin, short subhymenial hyphal cells,
utriform basidia, and the absence of rhizomorphs and cystidia. However, T. longiechinuli differs from T. conclusa by
having pelliculose basidiocarps, non-inflated subhymenial
hyphal cells and subglobose to bi-lobed basidiospores with
longer echinuli (echinuli up to 3 μm long). T. longisterigmata X. Lu, K. Steffen & H.S. Yuan resembles T. conclusa
by having mucedinoid basidiocarps adherent to the substrate,
utriform basidia, basidiospores of approximately the same
shape and size, and the absence of rhizomorphs and cystidia.
However, the former species is differentiated by dark brown
to chestnut basidiocarps, and slightly thick-walled subhymenial hyphae (Lu et al. 2018b).
Tomentella cystidiata H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555694; Facesoffungi number: FoF 05613; Figs. 148, 149, 150
Etymology: Refers to the species having cystidia.
Holotype: IFP 019262.
Basidiocarps annual, resupinate, separable from the substrate, arachnoid, without odour or taste when fresh, 0.4–0.6
mm thick, continuous. Hymenophoral surface smooth,
brown to dark brown (7E5–7F6) and darker than subiculum when dry. Sterile margin often indeterminate, byssoid,
paler than hymenophore. Subiculum mostly brown. Rhizomorphs present in subiculum, 10–30 μm diam; rhizomorphic
surface more or less smooth; hyphae in rhizomorph monomitic, differentiated, of type C, compactly arranged; hyphae
in central rhizomorph clamped, thick-walled, unbranched,
3–4 μm diam; hyphae at outer part of rhizomorph clamped,
thick-walled, unbranched, 1.5–2.5 μm diam; all the hyphae
greyish orange in KOH, cyanophilous, inamyloid. Subicular
hyphae monomitic; generative hyphae with both of clamps
and simple septa, slightly thick- to distinctly thick-walled,
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Fig. 138 Microscopic structures
of Tomentella changbaiensis
(IFP 019257, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
Fig. 139 A basidiocarp of Tomentella citrinocystidiata (IFP 019331,
holotype)
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rarely branched, 4–6 μm diam, occasionally collapsed, not
encrusted, orange in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, thin- to slightly thick-walled,
rarely branched, 4–6 μm diam; hyphal cells more or less
uniform, orange in KOH, cyanophilous, inamyloid. Cystidia
arising from subhymenial or subicular hyphae, thin-walled,
50–65 μm long, 5–6 μm diam at base and 9–10 μm diam at
apex, capitate, embedded, without encrustation, pale brown
in KOH. Basidia 25–65 μm long and 6–8 μm diam at apex,
5–7 μm at base, with a clamp connection at base, clavate,
stalked, sinuous, rarely with transverse septa, orange in
KOH, yellowish brown in distilled water, 4-sterigmate; sterigmata 2.5–4.5 μm long and 1–2.5 μm diam at base. Basidiospores thick-walled, (7.5–)7.9–9(–9.5) × (5.5–)6.1–8(–8.5)
μm, L = 8.53 μm, W = 7.47 μm, Q = 1.12–1.16 (n = 60/2),
irregularly subglobose or lobed in frontal and lateral views,
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Fig. 140 SEM of basidiospores of Tomentella citrinocystidiata (IFP
019331, holotype)
aculeate, reddish orange in KOH, reddish orange in distilled
water, cyanophilous, inamyloid; echinuli usually isolated,
sometimes grouped in 2 or more, up to 2.5 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
wood debris, 21 October 2015, Yuan 10693 (IFP 019262,
holotype), Yuan 10704; Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood
debris, 10 October 2015, Yuan 10620 (IFP 019261).
GenBank numbers: ITS: KY686219, KY686218; LSU:
MK446354, MK446353.
Notes: Tomentella capitata is similar to T. cystidiata by
having brown to dark brown, continous basidiocarps separable from the substrate, the presence of rhizomorphs and
capitate cystidia, thin- to thick-walled subhymenial hyphae
and basidiospores of approximately the same shape and size.
However, it differs in its granulose or farinose hymenophore,
the presence of cystidia from the rhizomorphic surface and
the absence of simple septa in the subicular hyphae (Yorou
et al. 2007). T. pilosa resembles T. cystidiata by having
arachnoid basidiocarps separable from the substrate, a byssoid sterile margin, the presence of rhizomorphs and capitate
cystidia, and basidiospores of approximately the same shape
and size. But, it differs by the pale coloured hymenophore
surface, the rhizomorphs with a monomitic to dimitic hyphal
structure and the absence of simple septa in the subicular
hyphae (Kõljalg 1996).
Tomentella dimidiata H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
169
Index Fungorum number: IF555738; Facesoffungi number: FoF 05614; Figs. 151, 152, 153
Etymology: Refers to the dimitic hyphal structure of the
rhizomorphs.
Holotype: IFP 019265.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.2–0.4 mm thick, continuous. Hymenophoral surface
smooth, golden brown to brown (5D7–6F5) and turning
lighter than the subiculum when dry. Sterile margin often
determinate, byssoid, concolorous with hymenophore. Rhizomorphs present in subiculum and margins, 12–50 μm
diam; rhizomorphic surface more or less smooth; hyphae
in rhizomorph dimitic, highly differentiated, of type G (this
a new type, see in the notes); central hyphae usually simple septate, thick-walled, unbranched, 2–4 μm diam, pale
brown in KOH, cyanophilous, inamyloid; skeletal hyphae
from outer part of rhizomorphs thick-walled, unbranched,
1.5–2.5 μm diam, pale brown in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae
clamped, thick-walled, 4–6 μm diam, without encrustation,
pale brown in KOH, acyanophilous, inamyloid. Subhymenial hyphae clamped, thin-walled, 4–6 μm diam; hyphal
cells short and not inflated, pale brown in KOH, cyanophilous, inamyloid. Cystidia absent. Basidia 10–40 μm long
and 4.5–12.5 μm diam at apex, 3.5–5.5 μm at base, with
a clamp connection at base, utriform, not stalked, sinuous,
without transverse septa, pale brown in KOH and in distilled
water, 4-sterigmate; sterigmata 5.5–11.5 μm long and 2–3
μm diam at base. Basidiospores slightly thick-walled, (6.5–
)7–7.5(–8) × (5–)5.5–7(–7.5) μm, L = 7.07 μm, W = 5.99 μm,
Q = 1.15–1.26 (n = 60/2), subglobose to bi-, tri- or quadralobed in frontal and lateral views, echinulate, pale brown in
KOH and distilled water, cyanophilous, inamyloid; echinuli
usually grouped in 2 or more, up to 2 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 3 August 2016, Yuan 11205 (IFP
019265, holotype); Jilin Province, Changbaishan Nature
Reserve, on fallen angiosperm branch, 7 August 2016, Yuan
11267 (IFP 019266).
GenBank numbers: ITS: MK211704, MK211705; LSU:
MK446355, MK446356.
Notes: Some species of Tomentella have been reported
to have dimitic rhizomorphs, for instance, T. botryoides, T.
brunneorufa M.J. Larsen, T. fungicola (Litsch.) M.J. Larsen,
T. pilosa, T. punicea (Alb. & Schwein.) J. Schröt. and T.
umbrinospora M.J. Larsen (Kõljalg 1996; Daemmrich 2006;
Peintner and Dämmrich 2012). In addition, some species
of Pseudotomentella Svrček and Odontia Pers. also have
dimitic rhizomorphs (Agerer et al. 2001; Tedersoo et al.
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Fig. 141 Microscopic structures
of Tomentella citrinocystidiata
(IFP 019331, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
2014; Yuan et al. 2018). Agerer recognized six different
types (type A–F) of the rhizomorphic hyphal organization,
but all these types are monomitic hyphal systems (Agerer
1987–2008). Yorou mentioned the presence of dimitic rhizomorphs in T. punicea and T. umbrinospora, but he treated
this kind of structure as type C. However, the dimitic hypal
13
system of rhizomorphs is significantly different from the
monomitic hyphal structure. We name this kind of structure
in rhizomorphs as type G.
Tomentella globospora and T. dimidiata formed a clade in
the phylogenetic tree (Fig. 111), and they share the similar
morphological and anatomical characteristics: basidiocarps
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Fig. 142 A basidiocarp of Tomentella coffeae (IFP 019259, holotype)
Fig. 143 SEM of basidiospores of Tomentella coffeae (IFP 019259,
holotype)
adherent to the substrate, thick-walled subicular hyphae,
utriform basidia and the absence of cystidia. However, T.
globospora differs from T. dimidiata by having crustose
basidiocarps, globose to subglobose basidiospores and the
absence of rhizomorphs. The key feature of T. dimidiata is
the presence of dimitic rhizomorphs in subiculum. T. incrustata and T. dimidiata both have a byssoid sterile margin,
short subhymenial hyphal cells, and subglobose to bi-, trior quadra-lobed basidiospores; but T. incrustata has clavate
basidia, encrusted subicular hyphae and generative hyphae
occasionally inflated.
171
Tomentella duplexa H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555462; Facesoffungi number: FoF 05615; Figs. 154, 155, 156
Etymology: Refers to the duplex basidiocarps.
Holotype: IFP 019267.
Basidiocarps annual, resupinate, duplex, adherent to the
substrate, pelliculose, without odour or taste when fresh,
0.6–1 mm thick, continuous. Hymenophoral surface smooth,
greyish brown to brown (7D3–7E4) and slightly lighter
than subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent. Subicular hyphae monomitic; generative
hyphae clamped and rarely simple septate, thick-walled,
occasionally branched, 3–6 μm diam, without encrustation,
pale brown in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped and rarely simple septate, thin-walled,
occasionally branched, 5–8 μm diam; hyphal cells short
and inflated, pale brown in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia 20–45 μm long and 5–9 μm
diam at apex, 3–5 μm at base, with a clamp connection at
base, clavate, not stalked, sinuous, rarely with transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 4–7 μm long and 1.5–2 μm diam at base.
Basidiospores slightly thick-walled, (7–)7.5–8.5(–9.5) × (6
.5–)7–8.5(–9) μm, L = 8.07 μm, W = 7.55 μm, Q = 1.03–1.08
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and in distilled water,
cyanophilous, inamyloid; echinuli usually isolated, up to 2.5
μm long.
Material examined: CHINA, Liaoning Province, Dalian,
Zhuanghe County, Xianrendong, Bingyugou National Geopark, on rotten angiosperm wood debris, 5 August 2017, Yuan
12207 (IFP 019267, holotype), Yuan 12202 (IFP 019268).
GenBank numbers: ITS: MK211707, MK211706; LSU:
MK446358, MK446357.
Notes: Tomentella bresadolae (Brinkmann) Bourdot &
Galzin and T. duplexa are closely related in the phylogeny
(Fig. 111) and they share greyish brown to brown basidiocarps adherent to the substrate, globose to subglobose basidiospores, the absence of rhizomorphs and cystidia. However,
T. bresadolae differs from T. duplexa by the stalked, utriform
basidia and more or less uniform subhymenial hyphal cells
(Wakefield 1960; Larsen 1969; Kõljalg 1996; Daemmrich
2006). T. griseofusca resembles T. duplexa by greyish brown
to brown basidiocarps adherent to the substrate, short and
inflated subhymenial hyphal cells, globose to subglobose
basidiospores, and the absence of rhizomorphs and cystidia.
But, T. griseofusca is differentiated by crustose basidiocarps
with a byssoid sterile margin and utriform basidia.
Tomentella efibulata H.S. Yuan & Y.C. Dai, sp. nov.
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Fig. 144 Microscopic structures
of Tomentella coffeae (IFP
019259, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
Fig. 145 A basidiocarp of Tomentella conclusa (IFP 019263, holotype)
13
Index Fungorum number: IF555695; Facesoffungi number: FoF 05616; Figs. 157, 158, 159
Etymology: Refers to the species having simple septate
hyphae.
Holotype: IFP 019269.
Basidiocarps annual, resupinate, easily separated from
the substrate, pelliculose, without odour or taste when
fresh, 0.2–0.3 mm thick, continuous. Hymenophoral surface smooth to indistinctly granulose, dark brown (6F5–7),
darker than or concolorous with subiculum when dry. Sterile margin often indeterminate, byssoid or fimbriate, paler
than hymenophore. Subiculum mostly yellowish brown.
Rhizomorphs present in subiculum and margins, 10–40 μm
diam; rhizomorphic suraface more or less smooth; hyphae
in rhizomorph monomitic, undifferentiated, of type B,
compactly arranged and of uniform; single hyphae simple
septate, thick-walled, unbranched, 1.5–2.5 μm diam, pale
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173
from the substrate, a dark brown and granulose hymenophoral surface, frequently simple septate hyphae, the presence of
rhizomorphs in subiculum and margin, and basidiospores of
approximately the same size and shape. However, T. africana
is differentiated by its subicular and subhymenial hyphae
with clamp connections and differentiated rhizomorphs of
type C (Yorou and Agerer 2008). T. badia (Link) Stalpers
resembles T. efibulata by having continuous basidiocarps
with a smooth to granulose hymenophoral surface of similar
colour, simple septate and thick-walled hyphae, the absence
of cystidia, and irregular globose to lobed, echinulate basidiospores. However, T. badia differs by having basidiocarps
adherent to the substrate, the absence of rhizomorphs, and
bigger basidiospores (8–11 μm, Kõljalg 1996).
Fig. 146 SEM of basidiospores of Tomentella conclusa (IFP 019263,
holotype)
yellow in KOH, cyanophilous, inamyloid. Subicular hyphae
monomitic; hyphae simple septate, thick-walled, occasionally branched, 3–5 μm diam, occasionally collapsed and
encrusted, pale yellow in KOH, slightly cyanophilous, inamyloid. Subhymenial hyphae simple septate, thin- to slightly
thick-walled, frequently branched, 2.5–5.5 μm diam; hyphal
cells more or less uniform, pale yellow in KOH, slightly
cyanophilous, inamyloid. Cystidia absent. Basidia 40–70 μm
long and 5–10 μm diam at apex, 3–5 μm at base, simple septate at base, clavate, stalked, sinuous, rarely with transverse
septa, pale yellow in KOH, pale yellow in distilled water,
4-sterigmate; sterigmata 3–7 μm long and 1–1.5 μm diam at
base. Basidiospores thick-walled, (5.5–)6–8.2(–8.5) × (5–)5
.2–7.3(–7.5) μm, L = 6.94 μm, W = 6.63 μm, Q = 1.04–1.12
(n = 60/2), subglobose to bi-, tri- or quadra-lobed in frontal
view and ellipsoid in lateral view, echinulate, greyish yellow
in KOH, reddish yellow in distilled water, cyanophilous, inamyloid; echinuli usually grouped in 2 or more, up to 0.3 μm.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
wood debris, 21 October 2015, Yuan 10699 (IFP 019269,
holotype); Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 3 August
2016, Yuan 11167 (IFP 019270).
GenBank numbers: ITS: KY686228, KY686229; LSU:
MK446359, MK446360.
Notes: Tomentella africana Yorou & Agerer is similar
to T. efibulata by having continuous basidiocarps separable
Tomentella efibulis H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555740; Facesoffungi number: FoF 05617; Figs. 160, 161, 162
Etymology: Refers to the simple septate hyphae of the
rhizomorphs.
Holotype: IFP 019271.
Basidiocarps annual, resupinate, adherent to the substrate, arachnoid, without odour or taste when fresh, 0.2–0.5
mm thick, continuous. Hymenophoral surface granulose,
brownish grey to brown (5C2–5F4) and concolorous with
subiculum when dry. Sterile margin often determinate, byssoid, concolorous with hymenophore. Rhizomorphs present
in subiculum only, 35–55 μm diam; rhizomorphic surface
more or less smooth; hyphae in rhizomorph monomitic,
undifferentiated, of type B, compactly arranged and of uniform; single hyphae simple septate, thick-walled, rarely
branched, 4–5 μm diam, pale brown in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative
hyphae clamped and rarely simple septate, thick-walled,
occasionally branched, 3–5 μm diam, without encrustation,
pale brown in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped and rarely simple septate, thin-walled,
rarely branched, 3–5 μm diam; hyphal cells more or less
uniform, pale brown in KOH, acyanophilous, inamyloid.
Cystidia absent. Basidia 15–35 μm long and 3–7 μm diam
at apex, 2–4 μm at base, with a clamp connection at base,
clavate, not stalked, sinuous, without transverse septa, pale
brown in KOH and in distilled water, 4-sterigmate; sterigmata 3–5 μm long and 1–1.5 μm diam at base. Basidiospores
slightly thick-walled, (4.5–)5–6(–6.5) × (4–)4.5–5(–5.5) μm,
L = 5.36 μm, W = 4.54 μm, Q = 1.15–1.21 (n = 60/2), subglobose to bi-, tri- or quadra-lobed in frontal and lateral views,
echinulate, pale brown in KOH and distilled water, cyanophilous, inamyloid; echinuli usually grouped in 2 or more,
up to 1 μm long.
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174
Fig. 147 Microscopic structures
of Tomentella conclusa (IFP
019263, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 148 A basidiocarp of Tomentella cystidiata (IFP 019262, holotype)
Fig. 149 SEM of basidiospores of Tomentella cystidiata (IFP
019262, holotype)
175
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 3 August 2016, Yuan 11241 (IFP
019271, holotype); Jilin Province. Changbaishan Nature
Reserve, on fallen angiosperm branch, 7 August 2016, Yuan
11329 (IFP 019272).
GenBank numbers: ITS: MK211708, MK211709; LSU:
MK446361, MK446362.
Notes: Tomentella cinerascens (P. Karst.) Höhn. &
Litsch. and T. efibulis formed a clade with high support in
our phylogeny (Fig. 111). Both species have greyish brown
and arachnoid basidiocarps, the presence of rhizomorphs
and the absence of cystidia. However, T. cinerascens differs
from T. efibulis by the thin-walled and clamped rhizomorphic hyphae, and globose to subglobose basidiospores (Kõljalg 1996; Daemmrich 2006; Melo et al. 2006). T. asperula
(P. Karst.) Höhn. & Litsch. resembles T. efibulis by having
arachnoid basidiocarps with a granulose hymenophoral surface, a byssoid sterile margin, the presence of rhizomorphs
and the absence of cystidia. However, the former species
is differentiated by thin-walled and clamped rhizomorphic
hyphae, and globose to subglobose basidiospores (Larsen
1970; Daemmrich 2006).
Tomentella farinosa H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555696; Facesoffungi number: FoF 05618; Figs. 163, 164, 165
Etymology: Refers to the farinose hymenophore.
Holotype: IFP 019273.
Basidiocarps annual, resupinate, separable from the substrate, arachnoid, without odour or taste when fresh, 0.3–0.5
mm thick, continuous. Hymenophoral surface smooth to
farinose, pale brown to brown (6D7–6E8) and darker than
subiculum when dry. Sterile margin often indeterminate,
byssoid, paler than hymenophore, olive brown. Subiculum
mostly olive brown (4E6–8). Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped,
thick-walled, frequently branched, 5–10 μm diam, occasionally collapsed and encrusted, reddish yellow in KOH,
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176
Fig. 150 Microscopic structures
of Tomentella cystidiata (IFP
019262, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 151 A basidiocarp of Tomentella dimidiata (IFP 019265, holotype)
Fig. 152 SEM of basidiospores of Tomentella dimidiata (IFP
019265, holotype)
177
cyanophilous, inamyloid. Subhymenial hyphae clamped,
slightly thick- to distinctly thick-walled, frequently branched,
4–7 μm diam; hyphal cells more or less uniform, reddish
yellow in KOH, cyanophilous, inamyloid. Cystidia absent.
Basidia 30–70 μm long and 5–10 μm diam at apex, 3–5
μm at base, with a clamp connection at base, clavate, not
stalked, sinuous, rarely with transverse septa, pale yellow
in KOH, yellowish brown in distilled water, 4-sterigmate;
sterigmata 3–6 μm long and 1–1.5 μm diam at base. Basidiospores thick-walled, (4.5–)5–6.9(–7.5) × (4.5–)4.7–6.3(–6
.5) μm, L = 5.84 μm, W = 5.5 μm, Q = 1.04–1.08 (n = 60/2),
irregularly subglobose or lobed in frontal and lateral views,
aculeate to echinulate, pale yellow in KOH, reddish yellow
in distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 1.2 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
branch, 21 October 2015, Yuan 10666 (IFP 019273, holotype); on rotten angiosperm wood debris, 21 October 2015,
Yuan 10656 (IFP 019274); on rotten angiosperm trunk, 20
October 2015, Yuan 10655 & 10660.
GenBank numbers: ITS: KY686250, KY 686,251.
Notes: Tomentella brunneorufa M.J. Larsen is similar to
T. farinosa by having continuous, arachnoid basidiocarps
separable from the substrate, a byssoid sterile margin and
basidiospores of approximately the same size. However, it
is differentiated by the fulvous basidiocarps, the presence of
rhizomorphs and globose basidiospores (Larsen 1974; Kõljalg 1996). T. capitata has characteristics similar to T. farinosa, such as brown, continous basidiocarps separable from
the substrate, a farinose hymenophore surface, the presence
of rhizomorphs, clamped, thick-walled subicular hyphae and
approximately the same shape basidiospores. But it differs
by having capitate cystidia and bigger basidiospores (7–9
μm, Yorou et al. 2007).
Tomentella flavidobadia H.S. Yuan & Y.C. Dai, sp. nov.
13
178
Fig. 153 Microscopic structures
of Tomentella dimidiata (IFP
019265, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 154 A basidiocarp of Tomentella duplexa (IFP 019267, holotype)
Fig. 155 SEM of basidiospores of Tomentella duplexa (IFP 019267,
holotype)
179
Index Fungorum number: IF555697; Facesoffungi number: FoF 05619; Figs. 166, 167, 168
Etymology: Refers to the yellowish brown basidiocarps.
Holotype: IFP 019277.
Basidiocarps annual, resupinate, separable from the
substrate, mucedinoid, without odour or taste when fresh,
0.2–0.25 mm thick, continuous. Hymenophoral surface
smooth, yellowish brown (5D8) and concolorous with subiculum when dry. Sterile margin often indeterminate, byssoid,
paler than hymenophore, pale brown (6D7). Rhizomorphs
present in subiculum, 10–20 μm diam; rhizomorphic surface
more or less smooth; hyphae in rhizomorphs monomitic,
undifferentiated, of type B, compactly arranged and of uniform; single hyphae clamped, thin- to slightly thick-walled,
unbranched, 1–2 μm diam, golden yellow in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative
hyphae clamped, thin- to slightly thick-walled, occasionally
branched, 3–4 μm diam, occasionally collapsed, without
encrustation, yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, thin-walled, frequently branched,
3–4.5 μm diam; hyphal cells more or less uniform, golden
yellow in KOH, acyanophilous, inamyloid. Cystidia absent.
Basidia 30–50 μm long and 6–9 μm diam at apex, 3–5 μm at
base, with a clamp connection at base, clavate, stalked, not
sinuous, rarely with transverse septa, golden yellow in KOH,
deep yellow in distilled water, 4-sterigmate; sterigmata 3–5
μm long and 1.5–2.5 μm diam at base. Basidiospores thickwalled, (6.5–)6.7–9(–9.5) × (4.5–)5–7.9(–8.5) μm, L = 7.72
μm, W = 5.68 μm, Q = 1.22–1.41 (n = 60/2), subglobose to
bi-, tri- or quadra-lobed in frontal view and ellipsoid in lateral view, echinulate, oxide yellow in KOH, deep yellow in
distilled water, cyanophilous, inamyloid; echinuli usually
grouped in 2 or more, up to 1 μm long.
Material examined: CHINA, Liaoning Province, Kuandian County, Baishilazi Nature Reserve, on fallen angiosperm branch, 1 August 2016, Yuan 11044 (IFP 019277,
holotype), Yuan 11061 (IFP 019278).
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Fig. 156 Microscopic structures
of Tomentella duplexa (IFP
019267, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
GenBank numbers: ITS: KY686231, KY686230; LSU:
MK446364, MK446365.
Notes: Tomentella kentuckiensis M.J. Larsen is similar to
T. flavidobadia by having mucedinoid, continuous basidiocarps, a smooth hymenophore, the presence of rhizomorphs,
clamped and thin-walled subhymenial hyphae and irregular
globose or lobed basidiospores. However, the former species
is differentiated by paler coloured basidiocarps adherent to
the substrate and bigger basidiospores (8–11 μm, Larsen
1974). T. longiaculeifera resembles T. flavidobadia by having mucedinoid, yellowish brown basidiocarps, a smooth
hymenophoral surface and approximately the same-sized
basidiospores. However, T. longiaculeifera lacks rhizomorphs and has thick-walled subhymenial hyphae.
Fig. 157 A basidiocarp of Tomentella efibulata (IFP 019269, holotype)
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Fig. 158 SEM of basidiospores of Tomentella efibulata (IFP 019269,
holotype)
Tomentella fuscocrustosa H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555741; Facesoffungi number: FoF 05620; Figs. 169, 170, 171
Etymology: Refers to the brown and crustose basidiocarps.
Holotype: IFP 019279.
Basidiocarps annual, resupinate, adherent to the substrate,
crustose, without odour or taste when fresh, 0.6–1 mm thick,
continuous. Hymenophoral surface smooth, pale brown to
brown (6D4–6E4) and concolorous with subiculum when
dry. Sterile margin often determinate, byssoid, concolorous
with hymenophore. Rhizomorphs present in subiculum and
margins, 25–40 μm diam; rhizomorphic surface more or less
smooth; hyphae in rhizomorphs monomitic, undifferentiated,
of type A, loosely interwoven and of uniform; single hyphae
clamped, thick-walled, occasionally branched, 3.5–4.5 μm
diam, pale brown in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae clamped and
rarely simple septate, thick-walled, occasionally branched,
3.5–6.5 μm diam, without encrustation, pale brown in KOH,
cyanophilous, inamyloid. Subhymenial hyphae clamped and
rarely simple septate, thin- to slightly thick-walled, rarely
branched, 4–7 μm diam; hyphal cells short and inflated, pale
brown in KOH, cyanophilous, inamyloid. Cystidia absent.
Basidia 20–60 μm long and 5–9 μm diam at apex, 1.5–4 μm
181
at base, with a clamp connection at base, clavate, stalked,
sinuous, without transverse septa, pale brown in KOH and
in distilled water, 4-sterigmate; sterigmata 3–6 μm long
and 1.5–2 μm diam at base. Basidiospores slightly thickwalled, (6–)6.5–8.5(–9) × (5–)5.5–6.5(–7) μm, L = 7.35 μm,
W = 6.12 μm, Q = 1.17–1.35 (n = 60/2), subglobose to bi- or
tri-lobed in frontal and lateral views, echinulate, pale brown
in KOH and distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up
to 1.2 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 9 August
2016, Yuan 11420 (IFP 019279, holotype), Yuan 11399
(IFP 019280).
GenBank numbers: ITS: MK211713, MK211712; LSU:
MK446367, MK446366.
Notes: Tomentella ellisii (Sacc.) Jülich & Stalpers is
similar to T. fuscocrustosa by having continuous basidiocarps adherent to the substrate with a smooth hymenophoral surface, short and inflated subhymenial hyphae cells,
the presence of rhizomorphs, the absence of cystidia, and
subglobose to bi- or tri-lobed basidiospores. However, T.
ellisii differs in its utriform basidia and thin-walled subicular and subhymenial hyphae (Stalpers 1993; Kõljalg 1996).
T. pallidomarginata resembles T. fuscocrustosa by having
continuous basidiocarps adherent to the substrate, a smooth
hymenophoral surface, short and inflated subhymenial
hyphal cells, the presence of rhizomorphs, the absence of
cystidia, and subglobose to bi- or tri-lobed basidiospores.
However, the former species is differentiated by dark blonde
to brown basidiocarps, a pale yellow to champagne sterile
margin and utriform basidia.
Tomentella fuscofarinosa H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555742; Facesoffungi number: FoF 05621; Figs. 172, 173, 174
Etymology: Refers to the farinaceous sterile margin of
the basidiocarps.
Holotype: IFP 019281.
Basidiocarps annual, resupinate, adherent to the substrate, arachnoid, without odour or taste when fresh, 0.3–0.6
mm thick, continuous. Hymenophoral surface granulose,
brown to dark brown (6E5–6F5) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped,
13
182
Fig. 159 Microscopic structures
of Tomentella efibulata (IFP
019269, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fig. 160 A basidiocarp of Tomentella efibulis (IFP 019271, holotype)
Fig. 161 SEM of basidiospores of Tomentella efibulis (IFP 019271,
holotype)
thick-walled, frequently branched, 5–10 μm diam, without
encrustation, pale brown in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped, thin-walled, occasionally
branched, 5–11 μm diam; hyphal cells more or less uniform,
183
pale brown in KOH, acyanophilous, inamyloid. Cystidia
absent. Basidia 10–50 μm long and 6–10 μm diam at apex,
4–7 μm at base, with a clamp connection at base, utriform,
stalked, sinuous, without transverse septa, pale brown in
KOH and in distilled water, 4-sterigmate; sterigmata 3–6
μm long and 2–2.5 μm diam at base. Basidiospores thickwalled, (6.5–)7–8 (–8.5) × (5–)5.5–6.5(–7.5) μm, L = 7.35
μm, W = 6.25 μm, Q = 1.15–1.24 (n = 60/2), subglobose to
bi-, tri- or quadra-lobed in frontal and lateral views, echinulate, pale brown in KOH and distilled water, cyanophilous,
inamyloid; echinuli usually grouped in 2 or more, up to 1.5
μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 30 July 2017, Yuan 12142 (IFP
019281, holotype), Yuan 12125 (IFP 019282).
GenBank numbers: ITS: MK211715, MK211714; LSU:
MK446369, MK446368.
Notes: Tomentella atrobadia and T. fuscofarinosa are
closely related in our phylogeny (Fig. 111), and they share
similar morphological and anatomical characteristics: continuous basidiocarps adherent to the substrate, clamped
hyphae, utriform and stalked basidia, and the absence of
rhizomorphs and cystidia. However, T. atrobadia differs
from T. fuscofarinosa by having crustose basidiocarps, a
byssoid sterile margin and thick-walled subhymenial hyphae.
T. brunneogrisea resembles T. fuscofarinosa by brown to
dark brown basidiocarps, utriform basidia and the absence
of rhizomorphs and cystidia. But, T. brunneogrisea is differentiated from T. fuscofarinosa by having short subhymenial
hyphal cells and globose to subglobose basidiospores with
longer echinuli (echinuli up to 2 μm long).
Tomentella fuscogranulosa H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555698; Facesoffungi number: FoF 05622; Figs. 175, 176, 177
Etymology: Refers to the brown and granulose hymenophore of the basidiocarps.
Holotype: IFP 019283.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh,
0.4–0.7 mm thick, continuous. Hymenophoral surface
smooth or granulose, brown (7E6–8) when dry. Sterile
margin often indeterminate, byssoid, concorlorous with
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Fig. 162 Microscopic structures
of Tomentella efibulis (IFP
019271, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in lateral view. d
Basidiospores in frontal view
subiculum. Subiculum paler than hymenophore, mostly
pale brown. Rhizomorphs present in subiculum and margins, 20–50 μm diam; rhizomorphic surface more or less
smooth; hyphae in rhizomorphs monomitic, undifferentiated, of type B, compactly arranged and of uniform; single
hyphae clamped, thick-walled, unbranched, 1–1.5 μm diam,
greyish yellow in KOH, cyanophilous, inamyloid. Subicular
hyphae monomitic; generative hyphae with both clamps and
simple septa, thin- to slightly thick-walled, rarely branched,
2–3.5 μm diam, occasionally collapsed and encrusted, greyish yellow in KOH, acyanophilous, inamyloid. Subhymenial
hyphae clamped, thin-walled, frequently branched, 2.5–3.5
μm diam; hyphal cells more or less uniform, greyish yellow
13
in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
30–60 μm long and 5–10 μm diam at apex, 4–6 μm at base,
with a clamp connection at base, clavate, stalked, sinuous,
without transverse septa, pale orange in KOH, yellowish
brown in distilled water, 4-sterigmate; sterigmata 1.5–3
μm long and 1–2 μm diam at base. Basidiospores thickwalled, (5–)5.5–8(–8.5) × (4.5–)5–6.8(–7) μm, L = 6.55 μm,
W = 6.02 μm, Q = 1.08–1.13 (n = 60/2), irregularly globose
or lobed in frontal view and ellipsoid in lateral view, echinulate, reddish brown in KOH, reddish golden in distilled
water, cyanophilous, inamyloid; echinuli usually grouped in
2 or more, up to 1 μm long.
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Fig. 163 A basidiocarp of Tomentella farinosa (IFP 019273, holotype)
Fig. 164 SEM of Tomentella farinosa (IFP 019273, holotype) (a
Subicular hyphae. b–d Basidiospores)
Material examined: CHINA, Liaoning Province, Kuandian County, Baishilazi Nature Reserve, on rotten angiosperm branch, 22 October 2015, Yuan 10725 (IFP 019283,
holotype), Yuan 10733 (IFP 019284).
185
GenBank numbers: ITS: KY686233, KY686232; LSU:
MK446371, MK446370.
Notes: Tomentella subvinosa (Burt) Bourdot & Galzin is
similar to T. fuscogranulosa by having basidiocarps adherent to the substrate with a smooth or granulose hymenophore, indeterminate sterile margins, the presence of rhizomorphs, thin-walled subhymenial hyphae and irregularly
globose or lobed basidiospores. However, the former species is differentiated by vinaceous brown, usually discontinuous basidiocarps, the absence of simple septa in generative hyphae, subicular hyphae without encrustation and
smaller basidiospores (6–8 μm, Bourdot and Galzin 1924).
T. griseoviolacea Litsch. is similar to T. fuscogranulosa by
having continuous basidiocarps with a byssoid sterile margin, a smooth or granulose hymenophore, the presence of
rhizomorphs, encrusted subicular hyphae, and irregularly
globose or lobed, echinulate spores. However, T. griseoviolacea has greyish brown to blackish brown and arachnoid
basidiocarps, generative hyphae lacking of simple septa and
smaller basidiospores (5.5–8 μm, Larsen 1974).
Tomentella fuscopelliculosa H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555743; Facesoffungi number: FoF 05623; Figs. 178, 179, 180
Etymology: Refers to the brown and pelliculose
basidiocarps.
Holotype: IFP 019285.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh,
0.8–1.5 mm thick, continuous. Hymenophoral surface
smooth, brown to dark brown (6E5–6F5) and lighter than
subiculum when dry. Sterile margin often indeterminate,
farinaceous, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
clamped and rarely simple septate, thick-walled, occasionally branched, 3.5–5 μm diam, without encrustation, pale
brown in KOH and in distilled water, cyanophilous, inamyloid. Subhymenial hyphae clamped and rarely simple
septate, thin-walled, rarely branched, 4–6 μm diam; hyphal
cells short and inflated, pale brown in KOH, cyanophilous,
inamyloid. Cystidia absent. Basidia 10–55 μm long and 4–9
μm diam at apex, 3–6 μm at base, with a clamp connection
at base, utriform, not stalked, sinuous, rarely with transverse
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Fig. 165 Microscopic structures
of Tomentella farinosa (IFP
019273, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 6–8.5 μm long and 1.5–2.5 μm diam at
base. Basidiospores thick-walled, (8–)8.5–9.5(–11) × (7.5–
)8–9(–10.5) μm, L = 8.75 μm, W = 8.23 μm, Q = 1.05–1.09
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and in distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 2 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 7 August
2016, Yuan 11305 (IFP 019285, holotype), Yuan 11316
(IFP 019286).
GenBank numbers: ITS: MK211716, MK211717; LSU:
MK446372, MK446373.
13
Notes: Tomentella stipitata is similar to T. fuscopelliculosa by basidiocarps adherent to the substrate, a farinaceous
sterile margin, short and inflated subhymenial hyphal cells,
utriform basidia, thick-walled basidiospores and the absence
of rhizomorphs and cystidia. However, T. stipitata differs
from T. fuscopelliculosa by having mucedinoid basidiocarps and subglobose to bi- or tri-lobed basidiospores.
T. pertenuis resembles T. fuscopelliculosa by basidiocarps
adherent to the substrate with a farinaceous sterile margin,
utriform basidia, the absence of rhizomorphs and cystidia,
and thick-walled, globose to subglobose basidiospores. But,
it is differentiated by yellowish brown basidiocarps, slightly
thick-walled subhymenial hyphae, and more or less uniform
subhymenial hyphal cells.
Fungal Diversity (2020) 104:1–266
Fig. 166 A basidiocarp of Tomentella flavidobadia (IFP 019277,
holotype)
Fig. 167 SEM of basidiospores of Tomentella flavidobadia (IFP
019277, holotype)
Tomentella globospora H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555699; Facesoffungi number: FoF 05624; Figs. 181, 182, 183
Etymology: Refers to the globose basidiospores.
Holotype: IFP 019287.
187
Basidiocarps annual, resupinate, adherent to the substrate, crustose, without odour or taste when fresh, 0.4–0.6
mm thick, continuous. Hymenophoral surface smooth,
brown to darkbrown (6E5–6F7) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped,
thick-walled, occasionally branched, 5–6 μm diam, encrustated, greyish yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, slightly thick- to thick-walled,
frequently branched, 4–6 μm diam; hyphal cells more or less
uniform, greyish yellow in KOH, cyanophilous, inamyloid.
Cystidia absent. Basidia 20–50 μm long and 4–9 μm diam
at apex, 6–8 μm at base, with a clamp connection at the
base, clavate, stalked, sinuous, rarely with transverse septa,
greyish yellow in KOH, yellowish brown in distilled water,
4-sterigmate; sterigmata 3–5 μm long and 0.5–1.5 μm diam
at base. Basidiospores thin- to slightly thick-walled, (8.5–
)9.1–10.2(–10.5) × (8.5–)8.7–9.7(–10) μm, L = 9.66 μm,
W = 9.33 μm, Q = 1.03–1.08 (n = 60/2), globose to subglobose in frontal and lateral views, aculeate, golden yellow
in KOH, brownish yellow in distilled water, cyanophilous,
inamyloid; echinuli usually isolated, sometimes grouped in
2, up to 2 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
wood debris, 21 October 2015, Yuan 10668 (IFP 019287,
holotype), 22 October 2015, Yuan 10748 (IFP 019288).
GenBank numbers: ITS: KY686242, KY686243; LSU:
MK446374, MK446375.
Notes: Tomentella terrestris (Berk. & Broome) M.J.
Larsen is similar to T. globospora by having crustose basidiocarps adherent to the substrate, a smooth hymenophore,
indeterminate sterile margins, the absence of rhizomorphs
and cystidia, and clamped hyphae. However, the former species is differentiated by thin-walled subhymenial hyphae and
smaller, triangular to ellipsoid basidiospores (7.5–9.5 μm,
Kõljalg 1996). T. alpina resembles T. globospora by having continuous basidiocarps adherent to the substrates, the
absence of rhizomorphs and cystidia, clamped and thickwalled subicular hyphae, and globose to subglobose basidiospores. However, it differs from T. globospora by having
thick basidiocarps (up to 1 mm thick) and smaller basidiospores (6.5–8.5 μm, Peintner and Dämmrich 2012).
Tomentella gloeocystidiata H.S. Yuan & Y.C. Dai, sp. nov.
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188
Fig. 168 Microscopic structures
of Tomentella flavidobadia
(IFP 019277, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 169 A basidiocarp of Tomentella fuscocrustosa (IFP 019279,
holotype)
Fig. 170 SEM of basidiospores of Tomentella fuscocrustosa (IFP
019279, holotype)
Index Fungorum number: IF555700; Facesoffungi number: FoF 05625; Figs. 184, 185, 186
189
Etymology: Refers to the species having gloeocystidia.
Holotype: IFP 019289.
Basidiocarps annual, resupinate, separable from the substrate, arachnoid, without odour or taste when fresh, 0.1–0.2
mm thick, continuous. Hymenophoral surface smooth, olive
yellow to olive (3D6–3E8) and concolorous with subiculum when dry. Sterile margin often indeterminate, byssoid,
concolorous with hymenophore. Rhizomorphs present in
subiculum, 20–35 μm diam; rhizomorphic surface more or
less smooth; hyphae in rhizomorphs monomitic, undifferentiated, of type B, compactly arranged and of uniform; single hyphae clamped, slightly thick-walled, unbranched, 2–3
μm diam, yolk yellow in KOH, cyanophilous, inamyloid.
Subicular hyphae monomitic; generative hyphae clamped,
slightly thick-walled, occasionally branched, 3–5 μm diam,
without encrustation, yolk yellow in KOH, cyanophilous,
inamyloid. Subhymenial hyphae clamped, thin- to slightly
thick-walled, occasionally branched, 4–5.5 μm diam; hyphal
cells more or less uniform, yolk yellow in KOH, acyanophilous, inamyloid. Gloeocystidia present, arising from subhymenial or subicular hyphae, 60–80 μm long, 4–5.5 μm
diam at base and 7–10 μm diam at apex, capitate, embedded, aseptate, occasionally encrusted on stalk, yolk yellow
in KOH. Basidia 40–60 μm long and 5–8 μm diam at apex,
5–8 μm at base, with a clamp connection at base, clavate,
not stalked, not sinuous, rarely with transverse septa, Chinese yellow in KOH, yellow in distilled water, 4-sterigmate;
sterigmata 2–4 μm long and 1–2 μm diam at base. Basidiospores thick-walled, (6.5–)7–8(–8.5) × (6–)6.3–7.6(–8) μm,
L = 7.59 μm, W = 6.96 μm, Q = 1.07–1.13 (n = 60/2), irregularly globose or lobed in frontal and lateral views, aculeate,
Chinese yellow in KOH, Chinese yellow in distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 2.5 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 3 August 2016, Yuan 11171
(IFP 019289, holotype), Yuan 11200 (IFP 019290).
GenBank numbers: ITS: KY686220, KY686221; LSU:
MK446376, MK446377.
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190
Fig. 171 Microscopic structures
of Tomentella fuscocrustosa
(IFP 019279, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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191
Notes: Tomentella capitata is similar to T. gloeocystidiata
by having continuous basidiocarps separable from the substrate, the presence of rhizomorphs, thick-walled subicular
hyphae, capitate cystidia and basidiospores of approximately
the same shape and size. However, it is differentiated by the
presence of cystidia both in the hymenium and the rhizomorphs and the absence of gloeocystidia (Yorou et al. 2007).
T. cystidiata and T. citrinocystidiata have capitate cystidia
and rhizomorphs, but the cystidia in the two species are
without oily appearance.
Fig. 172 A basidiocarp of Tomentella fuscofarinosa (IFP 019281,
holotype)
Fig. 173 SEM of basidiospores of Tomentella fuscofarinosa (IFP
019281, holotype)
Tomentella griseocastanea H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555744; Facesoffungi number: FoF 05626; Figs. 187, 188, 189
Etymology: Refers to the greyish brown hymenophoral
surface of the basidiocarps.
Holotype: IFP 019291.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.5–0.8 mm thick, continuous. Hymenophoral surface
granulose, greyish brown to dark blond (5D3–5D4) and
concolorous with subiculum when dry. Sterile margin often
indeterminate, farinaceous, concolorous with hymenophore.
Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped and simple septate, thick-walled,
rarely branched, 3–5 μm diam, without encrustation, some
subicular hyphal cells short, pale brown in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped and simple septate, thin- to slightly thick-walled, rarely branched,
4–6 μm diam; hyphal cells short and inflated, pale brown
in KOH, cyanophilous, inamyloid. Cystidia absent. Basidia
15–35 μm long and 4–7 μm diam at apex, 3–5 μm at base,
with a clamp connection at base, clavate, not stalked, sinuous, rarely with transverse septa, pale brown in KOH and in
distilled water, 4-sterigmate; sterigmata 3–5 μm long and
1.5–2 μm diam at base. Basidiospores slighty thick-walled,
(5–)5.5–6.5(–7) × (4–)5–5.5(–6) μm, L = 5.92 μm, W = 5.06
μm, Q = 1.12–1.21 (n = 60/2), subglobose to bi-, tri- or
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192
Fig. 174 Microscopic structures
of Tomentella fuscofarinosa
(IFP 019281, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 175 A basidiocarp of Tomentella fuscogranulosa (IFP 019283,
holotype)
Fig. 176 SEM of basidiospores of Tomentella fuscogranulosa (IFP
019283, holotype)
193
quadra-lobed in frontal and lateral views, echinulate, pale
brown in KOH and distilled water, cyanophilous, inamyloid;
echinuli usually isolated, sometimes grouped in 2 or more,
up to 1 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 9 August
2016, Yuan 11409 (IFP 019291, holotype), Yuan 11401
(IFP 019292).
GenBank numbers: ITS: MK211719, MK211718; LSU:
MK446379, MK446378.
Notes: Tomentella coerulea Höhn. & Litsch. resembles
T. griseocastanea by having greyish brown and continuous basidiocarps adherent to the substrate with a granulose
hymenophoral surface, a farinaceous sterile margin, the
absence of cystidia, and subglobose to bi-, tri- or quadralobed basidiospores. However, T. coerulea is differentiated
by a whitish sterile margin, thin-walled subicular hyphae
and the presence of rhizomorphs (Kõljalg 1996; Daemmrich
2006; Melo et al. 2006). T. subclavigera Litsch., T. clavigera Litsch. and T. griseocastanea also share continuous
basidiocarps adherent to the substrate, short subhymenial
hyphal cells and the absence of rhizomorphs. However, T.
subclavigera and T. clavigera differ from T. griseocastanea
by having thin-walled subicular hyphae and clavate cystidia
(Kõljalg 1996; Melo et al. 1998; Daemmrich 2006).
Tomentella griseofusca H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555701; Facesoffungi number: FoF 05627; Figs. 190, 191, 192
Etymology: Refers to the greyish brown colour of the
basidiocarps.
Holotype: IFP 019293.
Basidiocarps annual, resupinate, adherent to the substrate, crustose, without odour or taste when fresh, 0.5–0.8
mm thick, continuous. Hymenophoral surface smooth, greyish brown to dark brown (7E4–7F4) and concolorous with
subiculum when dry. Sterile margin often indeterminate,
byssoid, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
clamped, thick-walled, rarely branched, 5–6.5 μm diam,
without encrustation, golden brown in KOH, cyanophilous,
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194
Fig. 177 Microscopic structures
of Tomentella fuscogranulosa
(IFP 019283, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 178 A basidiocarp of Tomentella fuscopelliculosa (IFP 019285,
holotype)
Fig. 179 SEM of basidiospores of Tomentella fuscopelliculosa (IFP
019285, holotype)
195
inamyloid. Subhymenial hyphae clamped, thick-walled,
occasionally branched, 6–9 μm diam; hyphal cells short and
occasionally inflated, golden brown in KOH, cyanophilous,
inamyloid. Cystidia absent. Basidia 35–55 μm long and
6–12 μm diam at apex, 5–9 μm at base, with a clamp connection at base, utriform, not stalked, not sinuous, without transverse septa, golden brown in KOH, golden brown in distilled
water, 4-sterigmate; sterigmata 4–7 μm long and 1.5–2.5 μm
diam at base. Basidiospores slightly thick-walled, (8–)8.2–
9.1(–9.5) × (7–)7.4–8.8(–9) μm, L = 8.60 μm, W = 8.21 μm,
Q = 1.03–1.18 (n = 60/2), globose to subglobose in frontal
and lateral views, aculeate, brownish yellow in KOH, brownish yellow in distilled water, cyanophilous, inamyloid; echinuli isolated, up to 2.5 μm.
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on rotten angiosperm branch, 2 August 2016, Yuan 11094 (IFP 019293,
holotype); on rotten angiosperm wood debris, 2 August
2016, Yuan 11104 (IFP 019294) & 11105.
GenBank numbers: ITS: KY686252, KY686253; LSU:
MK446380, MK446381.
Notes: Tomentella griseoumbrina Litsch. resembles T.
griseofusca by having continuous basidiocarps adherent
to the substrate with a smooth hymenophore, the absence
of rhizomorphs, clamped hyphae without encrustations
and globose to subglobose basidiospores. However, it differs from T. griseofusca by having pale brown basidiocarps
and smaller basidiospores with short echinuli (4.5–5.5 μm,
Kõljalg 1996). T. beaverae Suvi & Kõljalg is similar to T.
griseofusca by having continuous, greyish brown basidiocarps adherent to the substrate with a byssoid sterile margin,
clamped, thick-walled subicular hphae, the absence of cystidia, and short, inflated subhymenial hyphae. But, it differs
in the presence of rhizomorphs, thick-walled subhymenial
hyphae and smaller, ellipsoid or slightly triangular basidiospores (6.8–8.6 μm, Suvi et al. 2010).
Tomentella griseomarginata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
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196
Fig. 180 Microscopic structures
of Tomentella fuscopelliculosa
(IFP 019285, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 181 A basidiocarp of Tomentella globospora (IFP 019287, holotype)
Fig. 182 SEM of basidiospores of Tomentella globospora (IFP
019287, holotype)
197
Index Fungorum number: IF555748; Facesoffungi number: FoF 05628; Figs. 193, 194, 195
Etymology: Refers to the grey sterile margin of the
basidiocarps.
Holotype: IFP 019295.
Basidiocarps annual, resupinate, adherent to the substrate,
mucedinoid, without odour or taste when fresh, 0.3–0.6 mm
thick, continuous. Hymenophoral surface smooth, greyish
brown to dark brown (6D3–6F5) and turning lighter than
subiculum when dry. Sterile margin often indeterminate,
farinaceous, paler than hymenophore, grey to orange-grey
(5B1–5B2). Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped and rarely simple septate,
thick-walled, occasionally branched, 4–6 μm diam, without
encrustation; subicular hyphal cells short and not inflated,
brown in KOH, cyanophilous, inamyloid. Subhymenial
hyphae clamped and rarely simple septate, thin-walled,
rarely branched, 4–6 μm diam; hyphal cells short and not
inflated, pale brown in KOH, acyanophilous, inamyloid.
Cystidia absent. Basidia 15–40 μm long and 5–9 μm diam
at apex, 3–5.5 μm at base, with a clamp connection at base,
clavate, not stalked, sinuous, without transverse septa, pale
brown in KOH and in distilled water, 4-sterigmate; sterigmata 2–5 μm long and 1.5–2 μm diam at base. Basidiospores
slightly thick-walled, (5.5–)6.5–7(–7.5) × (5.5–)6–6.5(–7)
μm, L = 6.86 μm, W = 6.28 μm, Q = 1.05–1.15 (n = 60/2),
globose to subglobose in frontal and lateral views, echinulate, pale brown in KOH and in distilled water, cyanophilous,
inamyloid; echinuli usually isolated, sometimes grouped in
2 or more, up to 1 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 10 August
2016, Yuan 11468 (IFP 019295, holotype), Yuan 11458
(IFP 019296).
GenBank numbers: ITS: MK211721, MK211720; LSU:
MK446383, MK446382.
Notes: Tomentella clavigera, T. subclavigera and T. griseomarginata are closely related in the phylogeny (Fig. 111).
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198
Fig. 183 Microscopic structures
of Tomentella globospora (IFP
019287, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
199
They share similar characteristics: an indeterminate and
farinaceous sterile margin, short subhymenial hyphal cells,
clavate cystidia and the absence of rhizomorphs. However, T.
clavigera and T. subclavigera differ from T. griseomarginata
by having thin-walled subicular hyphae and subglobose to
bi- and tri-lobed basidiospores (Kõljalg 1996; Melo et al.
1998; Daemmrich 2006). T. griseocastanea also resembles
T. griseomarginata by having mucedinoid and continuous
basidiocarps adherent to the substrate, a farinaceous sterile
margin, clavate basidia and the absence of rhizomorphs and
cystidia. However, the former species is differentiated by a
greyish brown sterile margin and subglobose to bi-, tri- or
quadra-lobed basidiospores.
Fig. 184 A basidiocarp of Tomentella gloeocystidiata (IFP 019289,
holotype)
Fig. 185 SEM of basidiospores of Tomentella gloeocystidiata (IFP
019289, holotype)
Tomentella inconspicua H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555702; Facesoffungi number: FoF 05629; Figs. 196, 197, 198
Etymology: Refers to the inconspicuous basidiocarps.
Holotype: IFP 019297.
Basidiocarps annual, resupinate, adherent to the substrate,
mucedinoid, without odour or taste when fresh, 0.1–0.2 mm
thick, continuous. Hymenophoral surface smooth to granulose, dark brown (6F4–6) and concolorous with subiculum
when dry. Sterile margin often indeterminate, byssoid, concolorous with hymenophore. Rhizomorphs present in subiculum, 10–20 μm diam; rhizomorphic surface more or less
smooth; hyphae in rhizomorph monomitic, undifferentiated,
of type B, compactly arranged and of uniform; single hyphae
clamped, thick-walled, unbranched, 2–3 μm diam, brownish
yellow in KOH, cyanophilous, inamyloid. Subicular hyphae
monomitic; generative hyphae clamped, thick-walled, 4–6
μm diam, rarely branched, without encrustation, pastel yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae
clamped, thin- to slightly thick-walled, frequently branched,
5–7 μm diam; hyphal cells more or less uniform, pastel yellow in KOH, cyanophilous, inamyloid. Cystidia absent.
Basidia 30–50 μm long and 7–10 μm diam at apex, 5–8
μm at base, with a clamp connection at base, clavate, not
stalked, not sinuous, without transverse septa, pastel yellow
in KOH, yellow in distilled water, 4-sterigmate; sterigmata
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200
Fig. 186 Microscopic structures
of Tomentella gloeocystidiata
(IFP 019289, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 187 A basidiocarp of Tomentella griseocastanea (IFP 019291,
holotype)
Fig. 188 SEM of basidiospores of Tomentella griseocastanea (IFP
019291, holotype)
201
3–5 μm long and 1–2 μm diam at base. Basidiospores thickwalled, (6–)6.4–7.3(–7.5) × (5–)5.3–6.8(–7) μm, L = 7.01
μm, W = 6.54 μm, Q = 1.05–1.13 (n = 60/2), irregularly
globose or lobed in frontal and lateral views, echinulate,
mustard yellow in KOH, mustard yellow in distilled water,
cyanophilous, inamyloid; echinuli isolated or grouped in 2
or more, up to 1.5 μm long.
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on rotten angiosperm branch, 2 August 2016, Yuan 11107 (IFP 019297,
holotype); on rotten angiosperm wood debris, 1 August
2016, Yuan 11060 (IFP 019298); Huanren County, Laotudingzi Nature Reserve, on rotten angiosperm wood debris, 21
October 2015, Yuan 10684.
GenBank numbers: ITS: KY686234, KY686235; LSU:
MK446385, MK446384.
Notes: Tomentella badia resembles T. inconspicua by
having brown, mucedinoid basidiocarps adherent to the
substrate, a smooth to granulose hymenophore, indeterminate sterile margins, the absence of cystidia, and echinulate
basidiospores with long echinuli. However, it differs from T.
inconspicua by having simple septate hyphae, the absence
of rhizomorphs, and bigger basidiospores (8–11 μm, Kõljalg 1996). T. parmastoana Suvi & Kõljalg is similar to T.
inconspicua by having brown, continuous and mucedinoid
basidiocarps adherent to the substrate, an indeterminate sterile margin, the presence of rhizomorphs, clamped, thickwalled subicular hyphae and echinulate basidiospores of
approximately the same shape and size. But, it differs by
a smooth to granulose hymenophoral surface, thick rhizomorphs (49–120 mm diam) and short, inflated subhymenial
hyphae (Suvi et al. 2010).
Tomentella incrustata H.S. Yuan, X. Lu & Y.C. Dai, sp.
nov.
Index Fungorum number: IF555749; Facesoffungi number: FoF 05630; Figs. 199, 200, 201
Etymology: Refers to the encrusted generative hyphae.
Holotype: IFP 019299.
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202
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Fig. 189 Microscopic structures
of Tomentella griseocastanea
(IFP 019291, holotype). a
Section through a basidiocarp.
b Basidiospores in frontal and
lateral views
Basidiocarps annual, resupinate, separable from the substrate, pelliculose, without odour or taste when fresh, 1–1.5
mm thick, continuous. Hymenophoral surface smooth, pale
brown to brown (6D8–6E8) and concolorous with subiculum when dry. Sterile margin often determinate, byssoid,
concolorous with hymenophore. Rhizomorphs present in
subiculum and margins, 6.5–25 μm diam; rhizomorphic surface more or less smooth; hyphae in rhizomorphs dimitic,
highly differentiated, of type G; central hyphae clamped,
thick-walled, rarely branched, 3–4 μm diam, with external
13
encrustation, pale yellow in KOH, cyanophilous, inamyloid;
skeletal hyphae at outer part of rhizomorphs thick-walled,
unbranched, 1–1.5 μm diam, pale yellow in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative
hyphae clamped and rarely simple septate, thin- to slightly
thick-walled, frequently branched; dominant, mostly 2 μm
diam and occasionally inflated to 4 μm diam, with external
encrustation, pale brown in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped and rarely simple septate,
thin-walled, frequently branched, 3–6 μm diam; hyphal cells
Fungal Diversity (2020) 104:1–266
Fig. 190 A basidiocarp of Tomentella griseofusca (IFP 019293, holotype)
Fig. 191 SEM of basidiospores of Tomentella griseofusca (IFP
019293, holotype)
203
short and not inflated, pale brown in KOH, acyanophilous,
inamyloid. Cystidia absent. Basidia 10–30 μm long and 3–6
μm diam at apex, 3–5 μm at base, with a clamp connection
at base, clavate, not stalked, sinuous, rarely with transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate;
sterigmata 6–7 μm and 1.5–2 μm diam at base. Basidiospores
slightly thick-walled, (5–)5.5–7(–7.5) × (4–)5–6.5(–7) μm,
L = 6.55 μm, W = 5.83 μm, Q = 1.08–1.15 (n = 60/2), subglobose to bi-, or tri-lobed in frontal and lateral views, echinulate, pale brown in KOH and distilled water, cyanophilous,
inamyloid; echinuli usually isolated, sometimes grouped in 2
or more, up to 1 μm long.
Material examined: CHINA, Liaoning Province, Anshan,
Qianshan Park, on fallen angiosperm branch, 1 August 2017,
Yuan 12189 (IFP 019299, holotype); Qingyuan County,
Experimental Station of Forest Ecology, on fallen angiosperm branch, 3 August 2016, Yuan 11158 (IFP 019300).
GenBank numbers: ITS: MK211723, MK211722; LSU:
MK446387, MK446386.
Notes: Tomentella umbrinospora and T. incrustata formed
a clade in the phylogeny (Fig. 111), and they share similar
morphological and anatomical characteristics: continuous
basidiocarps separable from the substrate, a byssoid sterile margin, clavate basidia, a dimitic structure of the rhizomorphs and the absence of cystidia. However, T. umbrinospora differs from T. incrustata by having dark brick, umber
or fulvous and arachnoid basidiocarps, thin-walled subicular
hyphae and subglobose to bi-lobed basidiospores (Kõljalg
1996; Peintner and Dämmrich 2012). T. fuscogranulosa is
similar to T. incrustata by the brown, pelliculose and continuous basidiocarps, the byssoid sterile margin, clavate
basidia, the presence of rhizomorphs and subglobose to bi-,
or tri-lobed basidiospores. However, T. fuscogranulosa can
be distinguished by the occasionally collapsed subicular
hyphae and the more or less uniform subhymenial hyphal
cells.
Tomentella interrupta H.S. Yuan & Y.C. Dai, sp. nov.
13
204
Fig. 192 Microscopic structures
of Tomentella griseofusca (IFP
019293, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
Fig. 193 A basidiocarp of Tomentella griseomarginata (IFP 019295,
holotype)
205
Index Fungorum number: IF555624; Facesoffungi number: FoF 05631; Figs. 202, 203, 204
Etymology: Refers to the discontinous basidiocarps.
Holotype: IFP 019241.
Basidiocarps annual, resupinate, separable from the
substrate, pelliculose, without odour or taste when fresh,
0.1–0.2 mm thick, discontinuous. Hymenophoral surface
smooth, pale brown to brown (6D6–6D8) and concolorous
with subiculum when dry. Sterile margin often indeterminate, byssoid, paler than hymenophore. Rhizomorphs present
in subiculum, 10–20 μm diam; rhizomorphic surface more or
less smooth; hyphae in rhizomorphs monomitic, undifferentiated, of type B, compactly arranged and of uniform; single
hyphae clamped, slightly thick-walled, unbranched, 1–2 μm
diam, pale yellow in KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae mostly clamped
and rarely simple septate, thick-walled, rarely branched, 3–4
μm diam, occasionally collapsed, not encrusted, greyish yellow in KOH, slightly cyanophilous, inamyloid. Subhymenial
hyphae clamped, thin-walled, frequently branched, 4–6 μm
diam; hyphal cells more or less uniform, greyish yellow in
KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
15–40 μm long and 5–9 μm diam at apex, 4–6 μm at base,
with a clamp at base, clavate, stalked, sinuous, rarely with
transverse septa, greyish yellow in KOH, greyish yellow in
distilled water, 4-sterigmate; sterigmata 3–5 μm long and
1–2 μm diam at base. Basidiospores thick-walled, (7.5–)7
.9–9.2(–9.5) × (7–)7.4–8.7(–9) μm, L = 8.66 μm, W = 8.23
μm, Q = 1.03–1.08 (n = 60/2), irregularly globose or lobed
in frontal view and ellipsoid in lateral view, echinulate to
aculeate, reddish yellow in KOH, orange in distilled water,
slightly cyanophilous, inamyloid; echinuli usually isolated,
sometimes grouped in 2 or more, up to 2 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 23 October 2015, Yuan 10775
(IFP 019241, holotype); on rotten angiosperm branch, 3
August 2016, Yuan 11203 (IFP 019242), Yuan 11183, 11197
Fig. 194 SEM of basidiospores of Tomentella griseomarginata (IFP
019295, holotype)
13
206
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Fig. 195 Microscopic structures
of Tomentella griseomarginata
(IFP 019295, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
& 11243; Huanren County, Laotudingzi Nature Reserve, on
rotten angiosperm branch, 20 October 2015, Yuan 10661.
GenBank numbers: ITS: KY686236, KY686237; LSU:
MK446388, MK446389.
Notes: Tomentella galzinii Bourdot and T. viridula (Bourdot & Galzin) Svrček resemble T. interrupta in having discontinuous, very thin basidiocarps, thin-walled subhymenial hyphae, and the same size and triangular to irregularly
globose basidiospores. However, the former species is differentiated by an olivaceous hymenophoral surface, acuminate, encrusted cystidia and the absence of rhizomorphs. T.
viridula differs from T. interrupta by its capitate cystidia
and the absence of rhizomorphs (Kõljalg 1996). Tomentella
13
kentuckiensis is similar to T. interrupta by having a smooth
hymenophore, the presence of rhizomorphs, thick-walled,
clamped and simple septate subicular hyphae and irregular or lobed basidiospores of approximately the same size.
However, it differs by having mucedinoid, continuous basidiocarps and slightly bigger basidiospores (8–11 μm diam,
Larsen 1974).
Tomentella liaoningensis H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555703; Facesoffungi number: FoF 05632; Figs. 205, 206, 207
Etymology: Refers to the species having a distribution in
Liaoning Province of China.
Fungal Diversity (2020) 104:1–266
Fig. 196 A basidiocarp of Tomentella inconspicua (IFP 019297,
holotype)
Fig. 197 SEM of basidiospores of Tomentella inconspicua (IFP
019297, holotype)
Holotype: IFP 019301.
207
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.2–0.4 mm thick, continuous. Hymenophoral surface
smooth, brown (6D7–6E8) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped
and simple septate, thick-walled, occasionally branched,
3–5 μm diam, occasionally collapsed, without encrustation,
pale orange in KOH, cyanophilous, inamyloid. Subhymenial
hyphae clamped, thin-walled, occasionally branched, 3–5
μm diam; hyphal cells more or less uniform, pale orange in
KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
20–50 μm long and 5–9 μm diam at apex, 3–5 μm at base,
with a clamp connection at base, utriform, stalked, sinuous, without transverse septa, pale orange in KOH, yellowish brown in distilled water, 4-sterigmate; sterigmata 2–5
μm long and 0.5–1 μm diam at base. Basidiospores thickwalled, (6–)6.3–7.8(–8.5) × (5.5–)6.1–7.5(–8) μm, L = 7.07
μm, W = 6.89 μm, Q = 1.02–1.08 (n = 60/2), irregularly globose or lobed in frontal and lateral views, aculeate, orange
in KOH, golden yellow in distilled water, cyanophilous,
inamyloid; echinuli usually isolated, sometimes grouped in
2 or more, up to 1.5 μm long.
Material examined: CHINA, Liaoning Province, Huanren
County, Laotudingzi Nature Reserve, on rotten angiosperm
branch, 21 October 2015, Yuan 10707 (IFP 019301, holotype); on rotten angiosperm wood debris, 21 October 2015,
Yuan 10681 (IFP 019302); on fallen angiosperm branch, 4
August 2018, Yuan 12982.
GenBank numbers: ITS: KY686257, MK250814.
Notes: Tomentella stuposa resembles T. liaoningensis by
having brown, mucedinoid, continuous basidiocarps adherent to the substrate, a smooth hymenophore, the absence of
rhizomorphs, subicular hyphae mostly clamped and rarely
simple septate, thin-walled subhymenial hyphae, and echinulate basidiospores. However, T. stuposa differs by slightly
globose and bigger basidiospores (8.5–11 μm, Kõljalg
1996). T. clavigera is similar to T. liaoningensis by having
brown, mucedinoid, continuous basidiocarps adherent to the
substrate, a smooth hymenophore, indeterminate and byssoid
sterile margins, the absence of rhizomorphs, and irregularly
globose or lobed basidiospores. But, the former species is
differentiated by clavate cystidia, the absence of simple septa
13
208
Fig. 198 Microscopic structures
of Tomentella inconspicua (IFP
019297, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
209
on generative hyphae, and bigger basidiospores (7.5–8.5 μm,
Kõljalg 1996).
Fig. 199 A basidiocarp of Tomentella incrustata (IFP 019299, holotype)
Fig. 200 SEM of basidiospores of Tomentella incrustata (IFP
019299, holotype)
Tomentella longiaculeifera H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555704; Facesoffungi number: FoF 05633; Figs. 208, 209, 210
Etymology: Refers to the long aculeate ornamentation on
the basidiospores.
Holotype: IFP 019303.
Basidiocarps annual, resupinate, separable from the
substrate, mucedinoid, without odour or taste when fresh,
0.4–0.6 mm thick, continuous. Hymenophoral surface
smooth, brownish yellow to brownish orange (5C5–7) and
concolorous with subiculum when dry. Sterile margin often
indeterminate, farinaceous, concolorous with hymenophore.
Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae mostly clamped and rarely simple septate,
thick-walled, occasionally branched, 4–6 μm diam, occasionally collapsed, with fine crystal encrustation, greyish
yellow in KOH, cyanophilous, inamyloid. Subhymenial
hyphae clamped, thick-walled, occasionally branched, 4–6
μm diam; hyphal cells slightly short and inflated, greyish
yellow in KOH, cyanophilous, inamyloid. Cystidia absent.
Basidia 20–60 μm long and 5–11 μm diam at apex, 5–8
μm at base, with a clamp at base, clavate, stalked, sinuous,
without transverse septa, greyish yellow in KOH, yellowish brown in distilled water, 4-sterigmate; sterigmata 5–11
μm long and 1–3 μm diam at base. Basidiospores slightly
thick-walled, (6.5–)6.9–10.1(–10.5) × (6–)6.5–9.7(–10) μm,
L = 8.66 μm, W = 8.24 μm, Q = 1.03–1.08 (n = 60/2), globose in frontal view and subglobose in lateral view, echinulate to aculeate, golden yellow in KOH, pale yellow in
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, up to 2.5 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
bark of fallen angiosperm trunk, 22 October 2015, Yuan
10744 (IFP 019303, holotype); Xifeng County, Binglashan
National Forest Park, on rotten angiosperm branch, 2 August
2016, Yuan 11119 (IFP 019304), Yuan 11127 & 11133; on
13
210
Fig. 201 Microscopic structures
of Tomentella incrustata (IFP
019299, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
Fig. 202 A basidiocarp of Tomentella interrupta (IFP 019241, holotype)
Fig. 203 SEM of basidiospores of Tomentella interrupta (IFP
019241, holotype)
211
rotten angiosperm wood debris, 3 August 2016, Yuan 11149,
11168, 11175, 11191 & 11198.
GenBank numbers: ITS: KY686238, KY686239; LSU:
MK446391, MK446392.
Notes: Tomentella stuposa is similar to T. longiaculeifera by having continuous and mucedinoid basidiocarps, a
smooth hymenophore, indeterminate sterile margins, the
absence of rhizomorphs, clamped and simple septate hyphae
and subglobose basidiospores of approximately the same
size. However, T. stuposa is differentiated by hazel-coloured
basidiocarps adherent to the substrate, and thin-walled subhymenial hyphae (Kõljalg 1996). T. globospora resembles T.
longiaculeifera by having brown, continuous basidiocarps,
the absence of rhizomorphs and cystidia, globose to subglobose basidiospores of approximately the same size and long
echinuli. But, the basidiocarps of the former are crustose
and adherent to the substrate, and the subhymenial hyphal
cells are uniform.
Tomentella longiechinuli H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555750; Facesoffungi number: FoF 05634; Figs. 211, 212, 213
Etymology: Refers to the long echinuli on the
basidiospores.
Holotype: IFP 019305.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh, 1–1.5
mm thick, continuous. Hymenophoral surface smooth, pale
brown to brown (6D4–6E5) and slightly lighter than subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent.
Subicular hyphae monomitic; generative hyphae clamped,
thick-walled, occasionally branched, 4–9 μm diam, without
encrustation, pale brown in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped, thin-walled, occasionally
branched, 4–7 μm diam; hyphal cells short and not inflated,
pale brown in KOH, acyanophilous, inamyloid. Cystidia
absent. Basidia 10–50 μm long and 6–13 μm diam at apex,
5–9 μm at base, with a clamp connection at base, utriform,
not stalked, sinuous, without transverse septa, pale brown
13
212
Fig. 204 Microscopic structures
of Tomentella interrupta (IFP
019241, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
Fig. 205 A basidiocarp of Tomentella liaoningensis (IFP 019301,
holotype)
Fig. 206 SEM of basidiospores of Tomentella liaoningensis (IFP
019301, holotype)
213
in KOH and in distilled water, 4-sterigmate; sterigmata 3–8
μm long and 2–3 μm diam at base. Basidiospores thickwalled, (8–)8.5–10.5(–12) × (7–)7.5–9(–10) μm, L = 9.37
μm, W = 8.47 μm, Q = 1.08–1.25 (n = 60/2), subglobose to
bi-lobed in frontal and lateral views, echinulate, pale brown
in KOH and in distilled water, cyanophilous, inamyloid;
echinuli usually isolated, up to 3 μm long.
Material examined: CHINA, Heilongjiang Province,
Fenglin National Nature Reserve, on fallen angiosperm
branch, 16 October 2016, Yuan 11979 (IFP 019305, holotype), Yuan 12083 (IFP 019306).
GenBank numbers: ITS: MK211726, MK211727; LSU:
MK446393, MK446394.
Notes: Tomentella griseofusca is similar to T. longiechinuli by the brownish basidiocarps, clamped hyphae, short
subhymenial hyphal cells, utriform basidia and the absence
of rhizomorphs and cystidia. However, the former species
is differentiated by crustose basidiocarps, thick-walled subhymenial hyphae and globose to subglobose basidiospores.
T. conclusa resembles T. longiechinuli by the basidiocarps
adherent to the substrate, short subhymenial hyphal cells,
utriform basidia and the absence of rhizomorphs and cystidia. However, T. conclusa differs from T. longiechinuli by
the inflated subhymenial hyphal cells and globose to subglobose basidiospores with shorter echinuli (echinuli up to
2.5 μm long).
Tomentella megaspora H.S. Yuan, X. Lu & Y.C. Dai, sp.
nov.
Index Fungorum number: IF555751; Facesoffungi number: FoF 05635; Figs. 214, 215, 216
Etymology: Refers to the large basidiospores.
Holotype: IFP 019307.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh, 0.5–0.8
mm thick, continuous. Hymenophoral surface smooth,
brown to dark brown (6E8–6F8) and concolorous with
subiculum when dry. Sterile margin often indeterminate,
13
214
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Fig. 207 Microscopic structures
of Tomentella liaoningensis
(IFP 019301, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
byssoid, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
clamped, thick-walled, frequently branched, 5–9 μm diam,
without encrustation, pale brown in KOH, cyanophilous,
inamyloid. Subhymenial hyphae clamped, thin-walled,
occasionally branched, 6–9 μm diam, without encrustation;
hyphal cells short and inflated, pale brown in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia 10–50 μm long
and 3–12 μm diam at apex, 5–7 μm at base, with a clamp
connection at base, utriform, not stalked, sinuous, without
transverse septa, pale brown in KOH and in distilled water,
4-sterigmate; sterigmata 3–5 μm long and 1–1.5 μm diam
at base. Basidiospores thick-walled, (8–)9–11(–11.5) × (7.5–
13
)8–10(–11) μm, L = 9.88 μm, W = 9.08 μm, Q = 1.08–1.12
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 2.5 μm long.
Material examined: CHINA, Jilin Province. Changbaishan Nature Reserve, on fallen angiosperm branch, 7 August
2016, Yuan 11326 (IFP 019307, holotype); 10 August 2016,
Yuan 11472 (IFP 019308).
GenBank numbers: ITS: MK211724, MK211725; LSU:
MK446395, MK446396.
Notes: Tomentella longiaculeifera is similar to T. megaspora by the byssoid sterile margin, clamped subicular and
Fungal Diversity (2020) 104:1–266
215
subhymenial hyphae, short and inflated subhymenial hyphal
cells, globose to subglobose basidiospores and the absence
of rhizomorphs and cystidia. However, T. longiaculeifera differs from T. megaspora by having mucedinoid basidiocarps,
clavate basidia, and encrusted subicular hyphae which are
occasionally collapsed. T. changbaiensis resembles T. megaspora by the byssoid sterile margin, clamped hyphae, short
and inflated subhymenial hyphal cells, utriform basidia, globose to subglobose basidiospores and the absence of rhizomorphs and cystidia. However, T. changbaiensis differs from
T. megaspora by having arachnoid basidiocarps with pale
brown sterile margin and duplex subiculum.
Fig. 208 A basidiocarp of Tomentella longiaculeifera (IFP 019303,
holotype)
Fig. 209 SEM of basidiospores of Tomentella longiaculeifera (IFP
019303, holotype)
Tomentella olivacea H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555705; Facesoffungi number: FoF 05636; Figs. 217, 218, 219
Etymology: Refers to the olive brown hymenophoral
surface.
Holotype: IFP 019309.
Basidiocarps annual, resupinate, separable from the
substrate, pelliculose, without odour or taste when fresh,
0.25–0.5 mm thick, continuous. Hymenophoral surface
smooth to indistinctly granulose, olive brown (4F5–7) and
turning darker than subiculum when dry. Sterile margin
often determinate, byssoid, paler than hymenophore, almost
pale brown. Subiculum mostly pale brown (5D6–8), Rhizomorphs present in subiculum, 10–20 μm diam; rhizomorphic
surface more or less smooth; hyphae in rhizomorph monomitic, undifferentiated, of type B, compactly arranged and of
uniform; single hyphae clamped, thick-walled, unbranched,
1–2 μm diam, yolk yellow in KOH, cyanophilous, inamyloid.
Subicular hyphae monomitic; generative hyphae clamped,
thin- to slightly thick-walled, rarely branched 3–4 μm diam,
without encrustation, yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, thin-walled, frequently
branched, 6–8 μm diam; hyphal cells more or less uniform,
yellow in KOH, acyanophilous, inamyloid. Cystidia absent.
Basidia 20–70 μm long and 5–7 μm diam at apex, 5–8 μm at
base, with a clamp connection at base, clavate, stalked, not
sinuous, without transverse septa, yellow in KOH, yellow
in distilled water, 4-sterigmate; sterigmata 4–5 μm long and
13
216
Fig. 210 Microscopic structures
of Tomentella longiaculeifera
(IFP 019303, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
Fig. 211 A basidiocarp of Tomentella longiechinuli (IFP 019305,
holotype)
Fig. 212 SEM of basidiospores of Tomentella longiechinuli (IFP
019305, holotype)
217
1.5–2 μm diam at base. Basidiospores thick-walled, (6–)6.5–
7.9(–8.5) × (5–)5.5–6.8(–7) μm, L = 7.0 μm, W = 6.27 μm,
Q = 1.08–1.17 (n = 60/2), irregularly globose or lobed in lateral and frontal views, echinulate, yolk yellow in KOH, yolk
yellow in distilled water, cyanophilous, inamyloid; echinuli
usually grouped in 2 or more, up to 0.5 μm long.
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on rotten branch
of Quercus, 2 August 2016, Yuan 11139 (IFP 019309, holotype); on rotten angiosperm branch, 1 August 2016, Yuan
11043 (IFP 019310), 2 August 2016, Yuan 11082, 11095,
11118 & 11141; on rotten angiosperm wood debris, Yuan
11103, 11108, 11113, 11120, 11121 & 11143.
GenBank numbers: ITS: KY686225, KY686224; LSU:
MK446398, MK446397.
Notes: Tomentella olivascens (Berk. & M.A. Curtis)
Bourdot & Galzin resembles T. olivacea in having olivaceous grey basidiocarps, thin- to slightly thick-walled subicular hyphae, thin-walled subhymenial hyphae and echinulate basidiospores of approximately the same size and shape.
But, it differs by having basidiocarps adherent to the substrate and the absence of rhizomorphs in subiculum (Kõljalg
1996). T. ochraceo-olivacea Litsch. is similar to T. olivacea
by having ochraceous to yellowish olivaceous baisidocarps,
pale brown subiculum and irregular globose basidiospores
of approximately the same size. However, it differs by its
adherent, crustaceous to membranaceous basidiocarps and
basidia frequently with transverse septa (Litschauer 1933).
Tomentella olivaceobrunnea H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555766; Facesoffungi number: FoF 05637; Figs. 220, 221, 222
Etymology: Refers to the olive brown hymenophoral
surface.
Holotype: IFP 019311.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
13
218
Fig. 213 Microscopic structures
of Tomentella longiechinuli
(IFP 019305, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
Fig. 214 A basidiocarp of Tomentella megaspora (IFP 019307, holotype)
Fig. 215 SEM of basidiospores of Tomentella megaspora (IFP
019307, holotype)
219
0.3–0.6 mm thick, continuous. Hymenophoral surface
granulose, brownish grey to olive brown (4D2–4E5) and
concolorous with subiculum when dry. Sterile margin often
indeterminate, farinaceous, concolorous with hymenophore.
Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped and rarely simple septate, slightly
thick- to thick-walled, occasionally branched, 3–6 μm diam,
without encrustation, pale brown in KOH, cyanophilous,
inamyloid. Subhymenial hyphae clamped and rarely simple
septate, thin-walled, occasionally branched, 4–7 μm diam;
hyphal cells short and inflated, hyaline to pale brown in
KOH, acyanophilous, inamyloid. Cystidia absent. Basidia
15–40 μm long and 5–8 μm diam at apex, 4–5 μm at base,
with a clamp connection at base, clavate, not stalked, sinuous, rarely with transverse septa, hyaline in KOH and in
distilled water, 4-sterigmate; sterigmata 2–8 μm long
and 1.5–2 μm diam at base. Basidiospores slightly thickwalled, (6–)6.5–7 (–7.5) × (5–)5.5–6(–6.5) μm, L = 6.68 μm,
W = 5.66 μm, Q = 1.15–1.23 (n = 60/2), subglobose to bi-,
tri- or quadra-lobed in frontal and lateral views, echinulate,
pale brown in KOH and distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2
or more, up to 1 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 3 August 2016, Yuan 11194 (IFP
019311, holotype); on rotten angiosperm wood debris, 31
July 2017, Yuan 12148 (IFP 019312).
GenBank numbers: ITS: MK211728, MK211729; LSU:
MK446399, MK446400.
Notes: Tomentella griseocastanea and T. olivaceobrunnea formed a clade in the phylogeny (Fig. 111), and they
share the same morphological and anatomical characteristics: mucedinoid and continuous basidiocarps adherent to
the substrate, a farinaceous sterile margin, short and inflated
subhymenial hyphal cells, clavate basidia and the absence
of rhizomorphs and cystidia. But, T. griseocastanea is differentiated by greyish brown basidiocarps and subglobose to
bi-, tri- or quadra-lobed basidiospores. T. coerulea is similar
to T. olivaceobrunnea by having continuous basidiocarps
13
220
Fig. 216 Microscopic structures
of Tomentella megaspora (IFP
019307, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
221
adherent to the substrate, a granulose hymenophoral surface,
a farinaceous sterile margin, the absence of cystidia, and
subglobose to bi-, tri- or quadra-lobed basidiospores. However, T. coerulea differs by its greyish brown to dark brown
basidiocarps, pale greyish brown and whitish sterile margin
and the presence of rhizomorphs (Kõljalg 1996; Daemmrich
2006; Melo et al. 2006).
Fig. 217 Basidiocarps of Tomentella olivacea (IFP 019309, holotype)
Fig. 218 SEM of basidiospores of Tomentella olivacea (IFP 019309,
holotype)
Tomentella pallidobrunnea H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555767; Facesoffungi number: FoF 05638; Figs. 223, 224, 225
Etymology: Referring to the species having pale brown
sterile margin.
Holotype: IFP 019313.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.5–1.5 mm thick, continuous. Hymenophoral surface
smooth, pale brown to dark brown (6D8–6F7) and concolorous with subiculum when dry. Sterile margin often
indeterminate, lighter than hymenophore, byssoid, pale
brown (6D8). Rhizomorphs absent. Subicular hyphae monomitic; generative hyphae clamped, thick-walled, frequently
branched, 4–7 μm diam, without encrustation, pale brown
in KOH, acyanophilous, inamyloid. Subhymenial hyphae
clamped, thin-walled, occasionally branched, 5–8 μm diam;
hyphal cells short and inflated, pale brown in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia 15–40 μm long
and 4–10 μm diam at apex, 3–6 μm at base, with a clamp
connection at base, utriform, not stalked, sinuous, without
transverse septa, pale brown in KOH and in distilled water,
4-sterigmate; sterigmata 3–6 μm long and 1.5–2.5 μm diam
at base. Basidiospores thick-walled, (7–)7.5–8.5(–9) × (6.5–
)7–8(–8.5) μm, L = 8.03 μm, W = 7.37 μm, Q = 1.05–1.12
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and in distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 2 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 10 August
13
222
Fig. 219 Microscopic structures
of Tomentella olivacea (IFP
019309, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 220 A basidiocarp of Tomentella olivaceobrunnea (IFP 019311,
holotype)
Fig. 221 SEM of basidiospores of Tomentella olivaceobrunnea (IFP
019311, holotype)
223
2016, Yuan 11493 (IFP 019313, holotype), Yuan 11481
(IFP 019314).
GenBank numbers: ITS: MK211731, MK211730; LSU:
MK446402, MK446401.
Notes: Tomentella bryophila resembles T. pallidobrunnea by having basidiocarps adherent to the substrate, the
absence of rhizomorphs, clamped hyphae, thick-walled
subicular hyphae, thin-walled subhymenial hyphae and
echinulate basidiospores of approximately the same shape.
But, it differs by having dark blonde to raw umber basidiocarps and thin-walled, bigger basidiospores (7.5–10.5 μm,
Kõljalg 1996). T. megaspora is similar to T. pallidobrunnea
by having brown to dark brown basidiocarps adherent to
the substrate, clamped hyphae, utriform basidia, globose to
subglobose basidiospores and the absence of rhizomorphs.
However, it differs by its pelliculose basidiocarps and bigger
basidiospores (9–11 × 8–10 μm).
Tomentella pallidomarginata H.S. Yuan, X. Lu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF555768; Facesoffungi number: FoF 05639; Figs. 226, 227, 228
Etymology: Refers to the pale yellow sterile margin of
basidiocarps.
Holotype: IFP 019315.
Basidiocarps annual, resupinate, adherent to the substrate, crustose, without odour or taste when fresh, 0.6–1.2
mm thick, continuous. Hymenophoral surface smooth, dark
blonde to brown (5D4–6E4) and turning darker than subiculum when dry. Sterile margin often determinate, byssoid,
paler than hymenophore, pale yellow (4A4). Rhizomorphs
present at the margin, 30–50 μm diam; rhizomorphic surface more or less smooth; hyphae in rhizomorph monomitic,
undifferentiated, of type A, loosely interwoven and uniform,
several hyphae growing out of the margin; single hyphae
clamped, thick-walled, occasionally branched, 5–6 μm diam,
pale brown in KOH, cyanophilous, inamyloid. Subicular
hyphae monomitic; generative hyphae clamped and rarely
simple septate, slightly thick- to distinctly thick-walled,
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224
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Fig. 222 Microscopic structures
of Tomentella olivaceobrunnea (IFP 019311, holotype). a
Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
occasionally branched, 5–7 μm diam, without encrustation,
pale brown in KOH, cyanophilous, inamyloid. Subhymenial
hyphae clamped and rarely simple septate, thin- to slightly
thick-walled, rarely branched, 5–9 μm diam; hyphal cells
short and inflated, pale brown in KOH, cyanophilous, inamyloid. Cystidia absent. Basidia 20–70 μm long and 5–13
μm diam at apex, 4–8 μm at base, with a clamp connection
at base, utriform, not stalked, sinuous, rarely with transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 4–8 μm long and 2–3 μm diam at base.
Basidiospores slightly thick-walled, (6.5–)7–8.5(–9) × (6–)
13
6.5–7(–7.5) μm, L = 7.54 μm, W = 6.85 μm, Q = 1.07–1.13
(n = 60/2), subglobose to bi- to tri-lobed in frontal and lateral
views, echinulate, pale brown in KOH and distilled water,
cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up to 1.5 μm long.
Material examined: CHINA, Jilin Province, Changbaishan Nature Reserve, on fallen angiosperm branch, 10 August
2016, Yuan 11474 (IFP 019315, holotype), 9 August 2016,
Yuan 11404 (IFP 019316).
GenBank numbers: ITS: MK211733, MK211732; LSU:
MK446404, MK446403.
Fungal Diversity (2020) 104:1–266
Fig. 223 A basidiocarp of Tomentella pallidobrunnea (IFP 019313,
holotype)
Fig. 224 SEM of basidiospores of Tomentella pallidobrunnea (IFP
019313, holotype)
225
Notes: Tomentella ellisii is similar to T. pallidomarginata by having continuous basidiocarps adherent to the
substrate, a smooth hymenophoral surface, the presence of
rhizomorphs, short and inflated subhymenial hyphal cells,
utriform basidia, the absence of cystidia, and subglobose to
bi- or tri-lobed basidiospores. However, T. ellisii differs by
its yellowish sterile margin and thin-walled subicular and
subhymenial hyphae (Stalpers 1993; Kõljalg 1996). T. hjortstamiana Suvi & Kõljalg resembles T. pallidomarginata by
having continuous basidiocarps adherent to the substrate, a
smooth hymenophoral surface, the presence of rhizomorphs,
short and inflated subhymenial hyphal cells, the absence of
cystidia, and subglobose to bi- or tri-lobed basidiospores.
However, the former species is differentiated by yellowish
grey and arachnoid or mucedinoid basidiocarps, clavate
basidia and thin-walled subicular and subhymenial hyphae.
Tomentella parvispora H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555769; Facesoffungi number: FoF 05640; Figs. 229, 230, 231
Etymology: Refers to the small basidiospores.
Holotype: IFP 019317.
Basidiocarps annual, resupinate, adherent to the substrate, pelliculose, without odour or taste when fresh, up to
0.1 mm thick, continuous. Hymenophoral surface smooth
to indistinctly granulose, yellowish brown (5E5–5F8) and
darker than subiculum when dry. Sterile margin often indeterminate, byssoid, light brown (5D5–8). Subiculum mostly
pale yellowish brown. Rhizomorphs present in subiculum,
10–20 μm diam; rhizomorphic surface more or less smooth;
hyphae in rhizomorph monomitic, undifferentiated, of type
B, compactly arranged and of uniform; single hyphae
clamped, thick-walled, unbranched, 1.5–2.5 μm diam, pale
orange in KOH, cyanophilous, inamyloid. Subicular hyphae
monomitic; generative hyphae clamped, thick-walled, occasionally branched, 4.5–6 μm diam, occasionally collapsed,
without encrustation, pale orange in KOH, cyanophilous,
inamyloid. Subhymenial hyphae clamped, thin- to slightly
thick-walled, occasionally branched, 3–6 μm diam; hyphal
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226
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Fig. 225 Microscopic structures
of Tomentella pallidobrunnea (IFP 019313, holotype). a
Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
cells more or less uniform, pale orange in KOH, cyanophilous, inamyloid. Cystidia absent. Basidia 20–40 μm long and
6–9 μm diam at apex, 5–7 μm at base, with a clamp connection at base, clavate, stalked, sinuous, without transverse
septa, pale orange in KOH, yellowish brown in distilled
water, 4-sterigmate; sterigmata 2–5 μm, 1–2 μm diam at
base. Basidiospores thick-walled, (5.5–)5.7–6.7(–7) × (5–)
5.3–6.4(–7) μm, L = 6.14 μm, W = 5.74 μm, Q = 1.04–1.09
(n = 60/2), irregularly globose or lobed in frontal view and
ellipsoid in lateral view, echinulate, pale brown in KOH,
pale brown in distilled water, cyanophilous, inamyloid; echinuli usually grouped in 2 or more, up to 0.6 μm long.
13
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on rotten angiosperm branch, 2 August 2016, Yuan 11144 (IFP 019317,
holotype); Huanren County, Laotudingzi Nature Reserve,
on rotten angiosperm wood debris, 20 October 2015, Yuan
10645; 21 October 2015, Yuan 10703; Qingyuan County,
Qingyuan Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 3 August 2016, Yuan 11196
(IFP 019318), 11162, 11176, 11181, 11182, 11185, 11188,
11192, 11193, 11214 & 11215; on rotten angiosperm
branch, Yuan 11179 & 11220.
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227
Notes: Tomentella minispora Yorou is similar to T. parvispora by having continuous basidiocarps adherent to the substrate, clamped, thin- to slightly thick-walled subhymenial
hyphae, the absence of cystidia, and small, echinulate basdiospores with echinuli of approximately equal size. However,
the former species has an absence of rhizomorphs, short
inflated externally encrusted subhymenial hyphae and a farinaceous basidiocarp margin (Yorou et al. 2013). T. olivacea
resembles T. parvispora by having pelliculose and continuous basidiocarps, a smooth and granulose hymenophore
with byssoid sterile margin, the presence of rhizomorphs,
clamped and thick-walled hyphae without encrustation and
irregular, echinulate basidiospores. However, it differs from
T. parvispora by having olive brown basidiocarps and bigger
basidiospores (6.5–7.9 × 5.5–6.8 μm).
Fig. 226 A basidiocarp of Tomentella pallidomarginata (IFP
019315, holotype)
Fig. 227 SEM of basidiospores of Tomentella pallidomarginata (IFP
019315, holotype)
GenBank numbers: ITS: KY686226, KY686227; LSU:
MK446405, MK446406.
Tomentella pertenuis H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555771; Facesoffungi number: FoF 05641; Figs. 232, 233, 234
Etymology: Refers to the very thin basidiocarps.
Holotype: IFP 019319.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.1–0.15 mm thick, continuous. Hymenophoral surface
smooth, yellowish brown (5F7–8) and concolorous with
subiculum when dry. Sterile margin often indeterminate,
farinaceous, concolorous with hymenophore. Rhizomorphs
absent. Subicular hyphae monomitic; generative hyphae
clamped, thick-walled, occasionally branched, 6–7 μm diam,
without encrustation, yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, slightly thick-walled,
occasionally branched, 7–9 μm diam; hyphal cells more or
less uniform, yellow in KOH, acyanophilous, inamyloid.
Cystidia absent. Basidia 40–60 μm long and 7–10 μm diam
at apex, 6–9 μm at base, with a clamp connection at base,
utriform, not stalked, not sinuous, without transverse septa,
yellow in KOH, yellow in distilled water, 4-sterigmate; sterigmata 5–8 μm long and 2–3 μm diam at base. Basidiospores
thick-walled, (7–)7.5–8.5(–9) × (6.5–)6.7–7.9(–8.5) μm,
L = 7.99 μm, W = 7.41 μm, Q = 1.06–1.11 (n = 60/2), globose, subglobse or irregularly globose in frontal and lateral
views, echinulate, yellow in KOH, yellow in distilled water,
cyanophilous, inamyloid; echinuli usually grouped in 2 or
more, up to 1.2 μm long.
13
228
Fig. 228 Microscopic structures
of Tomentella pallidomarginata (IFP 019315, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 229 A basidiocarp of Tomentella parvispora (IFP 019317, holotype)
Fig. 230 SEM of basidiospores of Tomentella parvispora (IFP
019317, holotype)
229
Material examined: CHINA, Liaoning Province, Xifeng
County, Binglashan National Forest Park, on bark of rotten angiosperm stump, 2 August 2016, Yuan 11131 (IFP
019319, holotype); on rotten angiosperm branch, 1 August
2016, Yuan 1,045 & 11064 (IFP 019320); Qingyuan County,
Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 3 August 2016, Yuan 11189 & 11204.
GenBank numbers: ITS: KY686241, KY686240; LSU:
MK446408, MK446407.
Notes: Tomentella tenuis Suvi & Kõljalg is similar to T.
pertenuis by having very thin and continuous basidiocarps
adherent to the substrate, clamped and thick-walled subicular
hyphae, the absence of cystidia, and ellipsoid to subglobose
basidiospores of approximately the same size. However, T.
tenuis differs by its greyish yellow basidiocarps, the presence of rhizomorphs in subiculum and margins, short and
inflated subhymenial hyphae with thin-walls and bi- to trifurcate echnuli (Suvi et al. 2010). T. terrestris resembles
T. pertenuis by having continuous and mucedinoid basidiocarps adherent to the substrate, a smooth hymenophore,
indeterminate sterile margin, the absence of rhizomorphs,
and echinulate basidiospores of approximately the same
size. However, the former species is differentiated by reddish brown basidiocarps, thin-walled subhymenial hyphae
and transverse septate basidia (Kõljalg 1996).
Tomentella qingyuanensis H.S. Yuan & Y.C. Dai, sp. nov.
Index Fungorum number: IF555772; Facesoffungi number: FoF 05642; Figs. 235, 236, 237
Etymology: Refers to where the type specimen was
collected.
Holotype: IFP 019321.
Basidiocarps annual, resupinate, adherent to the substrate, arachnoid, without odour or taste when fresh, 0.1–0.3
mm thick, continuous. Hymenophoral surface smooth or
indistinctly granulose, brown to dark brown (7E8–7F8) and
concolorous with subiculum when dry. Sterile margin often
indeterminate, farinaceous, concolorous with hymenophore.
Rhizomorphs present in subiculum and margin, 10–35 μm
diam; rhizomorphic surface more or less smooth; hyphae in
rhizomorph monomitic, differentiated, of type C, compactly
13
230
Fig. 231 Microscopic structures
of Tomentella parvispora (IFP
019317, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
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Fungal Diversity (2020) 104:1–266
Fig. 232 A basidiocarp of Tomentella pertenuis (IFP 019319, holotype)
Fig. 233 SEM of basidiospores of Tomentella pertenuis (IFP
019319, holotype)
231
arranged; hyphae in central rhizomorph with both clamps
and simple septa, thick-walled, unbranched, 3.5–4.5 μm
diam; hyphae at outer part of rhizomorph with both clamps
and simple septa, thick-walled, unbranched, 1.5–2.5 μm
diam; all the hyphae pale yellow in KOH, cyanophilous,
inamyloid. Subicular hyphae monomitic; generative hyphae
clamped and simple septate, thin- to slightly thick-walled,
occasionally branched, 2–3.5 μm diam, occasionally collapsed, without encrustation, pale yellow in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, thinwalled, occasionally branched, 2.5–4.5 μm diam; hyphal
cells more or less uniform, pale yellow in KOH, acyanophilous, inamyloid. Cystidia absent. Basidia 20–60 μm long and
5–10 μm diam at apex, 4–7 μm at base, with a clamp connection at base, clavate, stalked, sinuous, without transverse
septa, pale yellow in KOH, pale yellow in distilled water,
4-sterigmate; sterigmata 2–5 μm long and 0.3–0.5 μm diam
at base. Basidiospores thick-walled, (6–)6.5–7.7(–8) × (5.5–
)6–6.9(–7.5) μm, L = 6.92 μm, W = 6.25 μm, Q = 1.07–1.15
(n = 60/2), irregularly globose or lobed in frontal view and
ellipsoid in lateral view, echinulate, orange in KOH, deep
yellow in distilled water, cyanophilous, inamyloid; echinuli
usually grouped in 2 or more, up to 1.2 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm branch, 10 October 2015, Yuan 10616 (IFP
019321, holotype); Xifeng County, Binglashan National
Forest Park, on rotten angiosperm wood debris, 2 August
2016, 11109 (IFP 019322) & 11145.
GenBank numbers: ITS: KY686223, KY686222; LSU:
MK446409, MK446410.
Notes: Tomentella maroana Yorou is similar to T. qingyuanensis by having dark brown, continuous basidiocarps
adherent to the substrate, a smooth or granulose hymenophore, the presence of rhizomorphs; generative hyphae with
both clamped and simple septa and echinulate basidiospores
of approximately the same shape. However, the former
species is differentiated by crustose basidiocarps, thickwalled subhymenial hyphae and bigger basidiospores with
short echinuli (8.5–11 × 7.5–8.5 μm, Yorou et al. 2011). T.
13
232
Fig. 234 Microscopic structures
of Tomentella pertenuis (IFP
019319, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
13
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Fungal Diversity (2020) 104:1–266
233
fuscogranulosa resembles T. qingyuanensis by having continuous basidiocarps adherent to the substrate, a smooth
or granulose hymenophore, the presence of rhizomorphs,
thin- to thick-wall subicular hyphae with both clamped and
simple septa and irregularly globose or lobed basidiospores.
However, it differs from T. qingyuanensis by having undifferentiated rhizomorphs (type B) and bigger basidiospores
(8.0–9.4 × 7.1–9.0 μm).
Fig. 235 A basidiocarp of Tomentella qingyuanensis (IFP 019321,
holotype)
Fig. 236 SEM of basidiospores of Tomentella qingyuanensis (IFP
019321, holotype)
Tomentella segregata H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555773; Facesoffungi number: FoF 05643; Figs. 238, 239, 240
Etymology: Refers to the isolated echinuli of the
basidiospores.
Holotype: IFP 019323.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.8–1.5 mm thick, discontinuous. Hymenophoral surface
smooth, pale brown to dark brown (6D8–6F6) and concolorous with subiculum when dry. Sterile margin often indeterminate, farinaceous, concolorous with hymenophore. Rhizomorphs absent. Subicular hyphae monomitic; generative
hyphae clamped and rarely simple septate, slightly thick- to
thick-walled, frequently branched, 3–5 μm diam, without
encrustation, pale brown in KOH, cyanophilous, inamyloid.
Subhymenial hyphae clamped and rarely simple septate,
thin-walled, occasionally branched, 4–6 μm diam; hyphal
cells short and inflated, pale brown in KOH, acyanophilous,
inamyloid. Cystidia absent. Basidia 15–40 μm long and 5–8
μm diam at apex, 4–6 μm at base, with a clamp connection
at base, utriform, not stalked, sinuous, without transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 4–7 μm long and 1.5–2 μm diam at base.
Basidiospores slightly thick-walled, (7.5–)8–9(–9.5) × (7–)
7.5–8.5(–9) μm, L = 8.65 μm, W = 7.7 μm, Q = 1.08–1.15
(n = 60/2), globose to subglobose in frontal and lateral
views, echinulate, pale brown in KOH and distilled water,
cyanophilous, inamyloid; echinuli usually isolated, up to 2.5
μm long.
13
234
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Fig. 237 Microscopic structures
of Tomentella qingyuanensis
(IFP 019321, holotype). a
Hyphae from a rhizomorph. b
Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
Material examined: CHINA, Liaoning Province, Huanren County, Laotudingzi National Nature Reserve, on fallen
angiosperm branch, 20 October 2015, Yuan 11256 (IFP
019323, holotype); Jilin Province, Changbaishan Nature
Reserve, on living tree root, 6 August 2016, Yuan 10650
(IFP 019324).
GenBank numbers: ITS: MK211735, MK211734; LSU:
MK446412, MK446411.
Notes: Tomentella globospora is similar to T. segregata by
the basidiocarps adherent to the substrate, a smooth hymenophoral surface, a farinaceous sterile margin, globose to
13
subglobose basidiospores and the absence of rhizomorphs
and cystidia. However, T. globospora differs from T. segregata by having crustose basidiocarps and more or less
uniform subhymenial hyphal cells. T. pertenuis is similar
to T. segregata by having basidiocarps adherent to the substrate with a smooth hymenophoral surface and farinaceous
sterile margin, utriform basidia, globose to subglobose
basidiospores and the absence of rhizomorphs and cystidia.
However, T. pertenuis differs from T. segregata by having
slightly thick-walled subicular hyphae and more or less uniform subhymenial hyphal cells.
Fungal Diversity (2020) 104:1–266
Fig. 238 A basidiocarp of Tomentella segregata (IFP 019323, holotype)
Fig. 239 SEM of basidiospores of Tomentella segregata (IFP
019323, holotype)
Tomentella separata H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555780; Facesoffungi number: FoF 05644; Figs. 241, 242 243
235
Etymology: Refers to the basidiocarps easily separable
from the substrate.
Holotype: IFP 019325.
Basidiocarps annual, resupinate, separable from the substrate, pelliculose, without odour or taste when fresh, 0.6–1.2
mm thick, continuous. Hymenophoral surface smooth,
honey yellow to yellowish brown (5D6–5F8) and turning
darker than subiculum when dry. Sterile margin often indeterminate, byssoid, concolorous with hymenophore. Rhizomorphs present in subiculum and margins, 5–45 μm diam;
rhizomorphic surface more or less smooth; hyphae in rhizomorph monomitic, undifferentiated, of type B, compactly
arranged and of uniform; single hyphae clamped and simple
septate, thick-walled, unbranched, 2–4 μm, pale brown in
KOH, cyanophilous, inamyloid. Subicular hyphae monomitic; generative hyphae clamped and rarely simple septate,
thin- to slighty thick-walled, frequently branched, 1.5–3.5
μm diam, without encrustation, pale brown in KOH, acyanophilous, inamyloid. Subhymenial hyphae clamped and
simple septate, thin-walled, frequently branched, 2.5–4.5
μm diam; hyphal cells short and inflated, pale brown in
KOH, inamyloid. Cystidia absent. Basidia 15–55 μm long
and 3.5–6 μm diam at apex, 2–3.5 μm at base, with a clamp
connection at base, clavate, stalked, sinuous, with transverse
septa, pale brown in KOH and in distilled water, 4-sterigmate; sterigmata 4–8 μm long and 1.5–2 μm diam at base.
Basidiospores slighty thick-walled, (6.5–)7–8(–8.5) × (5.5–
)6–7.5(–8) μm, L = 7.37 μm, W = 6.74 μm, Q = 1.08–1.25
(n = 60/2), subglobose to bi-, tri- or quadra-lobed in frontal and lateral views, echinulate, pale brown in KOH and
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 1 μm long.
Material examined: CHINA, Liaoning Province, Huanren County, Laotudingzi National Nature Reserve, on fallen
angiosperm trunk, 20 October 2015, Yuan 10664 (IFP
019325, holotype); on fallen angiosperm branch, 20 October 2015, Yuan 10654 (IFP 019326).
GenBank numbers: ITS: MK211737, MK211736; LSU:
MK850196, MK850197.
Notes: Tomentella botryoides is similar to T. separata by
the continuous basidiocarps, with byssoid sterile margin,
the presence of rhizomorphs and the absence of cystidia.
However, T. botryoides differs from T. separata by having
fuscous black, mouse grey or greyish sepia basidiocarps,
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236
Fungal Diversity (2020) 104:1–266
Fig. 240 Microscopic structures
of Tomentella segregata (IFP
019323, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
more or less uniform subhymenial hyphal cells and subglobose to bi- or tri-lobed basidiospores (Kõljalg 1996; Daemmrich 2006). T. brunneoflava resembles T. separata by having a byssoid sterile margin, the presence of rhizomorphs,
clamped and simple septate hyphae, short and inflated subhymenial hyphal cells, clavate basidia, and the absence of
cystidia. But, it differs by the brownish orange, arachnoid
basidiocarps and subglobose to bi-, tri-lobed basidiospores.
13
Tomentella stipitata H.S. Yuan, X. Lu & Y.C. Dai, sp. nov.
Index Fungorum number: IF555774; Facesoffungi number: FoF 05645; Figs. 244, 245, 246
Etymology: Refers to the stalked basidia.
Holotype: IFP 019327.
Basidiocarps annual, resupinate, adherent to the substrate, mucedinoid, without odour or taste when fresh,
0.5–0.8 mm thick, continuous. Hymenophoral surface
smooth, dark blonde to yellowish brown (5D4–5F8) and
Fungal Diversity (2020) 104:1–266
Fig. 241 A basidiocarp of Tomentella separata (IFP 019325, holotype)
Fig. 242 SEM of basidiospores of Tomentella separata (IFP 019325,
holotype)
237
turning lighter than subiculum when dry. Sterile margin
often indeterminate, farinaceous, dark blonde (5D4). Rhizomorphs absent. Subicular hyphae monomitic; generative
hyphae clamped, frequently branched, thick-walled, 5–8 μm
diam, without encrustation, pale brown in KOH, acyanophilous, inamyloid. Subhymenial hyphae clamped, thin-walled,
occasionally branched, 5–8 μm diam; hyphal cells short and
inflated, pale brown in KOH, acyanophilous, inamyloid.
Cystidia absent. Basidia 20–50 μm long and 6–9 μm diam at
apex, 5–6 μm at base, with a clamp connection at base, utriform, stalked, sinuous, without transverse septa, pale brown
in KOH and in distilled water, 4-sterigmate; sterigmata 6–10
μm long and 1.5–2 μm diam at base. Basidiospores thickwalled, (7–)7.5–8.5(–9) × (6–)6.5–8(–8.5) μm, L = 7.83 μm,
W = 6.88 μm, Q = 1.06–1.15 (n = 60/2), subglobose to bi- or
tri-lobed in frontal and lateral views, echinulate, pale brown
in KOH and distilled water, cyanophilous, inamyloid; echinuli usually isolated, sometimes grouped in 2 or more, up
to 2.5 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on
fallen angiosperm branch, 3 August 2016, Yuan 11160 (IFP
019327, holotype), 30 July 2017, Yuan 12143 (IFP 019328).
GenBank numbers: ITS: MK211740, MK211741; LSU:
MK446413, MK446414.
Notes: Tomentella bryophila is similar to T. stipitata by
the dark blonde to raw umber basidiocarps adherent to the
substrate, a smooth hymenophoral surface, a farinaceous
sterile margin, clamped hyphae, short and inflated subhymenial hyphal cells, utriform basidia and the absence of rhizomorphs and cystidia. However, T. bryophila differs from
T. stipitata by having globose to subglobose basidiospores
(Kõljalg 1996). T. asiae-orientalis resembles T. stipitata by
basidiocarps adherent to the substrate, a farinaceous sterile
margin, clamped hyphae, thick-walled basidiospores and the
absence of rhizomorphs and cystidia. However, T. asiaeorientalis differs from T.stipitata by having more or less uniform subhymenial hyphal cells, utriform basidia and globose
to subglobose basidiospores.
Tomentella storea H.S. Yuan & Y.C. Dai, sp. nov.
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238
Fungal Diversity (2020) 104:1–266
Fig. 243 Microscopic structures
of Tomentella separata (IFP
019325, holotype). a Hyphae
from a rhizomorph. b Section through a basidiocarp. c
Basidiospores in frontal view. d
Basidiospores in lateral view
Index Fungorum number: IF555775; Facesoffungi number: FoF 05646; Figs. 247, 248, 249
Etymology: Refers to the mat-like basidiocarps.
Holotype: IFP 019330.
Basidiocarps annual, resupinate, separable from the substrate, mat-like, without odour or taste when fresh, 0.8–1.2
mm thick, continuous. Hymenophoral surface smooth to
indistinctly granulose, dark brown (6F5–8) and concolorous
13
with subiculum when dry. Sterile margin often indeterminate, byssoid, dark brown. Rhizomorphs absent. Subicular
hyphae monomitic; generative hyphae mostly clamped and
occasionally simple septate, thick-walled, rarely branched,
4–5 μm diam, occasionally collapsed and encrusted, yellowish orange in KOH, cyanophilous, inamyloid. Subhymenial hyphae clamped, thin-walled, occasionally branched,
3–6 μm diam; hyphal cells more or less uniform, yellowish
Fungal Diversity (2020) 104:1–266
Fig. 244 A basidiocarp of Tomentella stipitata (IFP 019327, holotype)
Fig. 245 SEM of basidiospores of Tomentella stipitata (IFP 019327,
holotype)
orange in KOH, acyanophilous, inamyloid. Cystidia absent.
Basidia 20–35 μm long and 4–9 μm diam at apex, 5–8 μm
at base, with a clamp connection at base, clavate, stalked,
239
sinuous, without transverse septa, yellowish orange in KOH,
yellowish brown in distilled water, 4-sterigmate; sterigmata
2–4 μm long and 1–1.5 μm diam at base. Basidiospores
thick-walled, (5.5–)6.1–7.1(–7.5) × (5.5–)5.7–6.6(–7) μm,
L = 6.66 μm, W = 6.21 μm, Q = 1.06–1.12 (n = 60/2), irregularly globose or lobed in frontal view and ellipsoid in lateral
view, echinulate, greyish yellow in KOH, golden yellow in
distilled water, cyanophilous, inamyloid; echinuli usually
isolated, sometimes grouped in 2 or more, up to 0.8 μm long.
Material examined: CHINA, Liaoning Province, Qingyuan County, Experimental Station of Forest Ecology, on rotten angiosperm wood debris, 22 October 2015, Yuan 10749
(IFP 019330, holotype); Huanren County, Laotudingzi
Nature Reserve, on rotten angiosperm wood debris, 20 October 2015, Yuan 10623 (IFP 019329); on fallen angiosperm
branch, 20 October 2015, Yuan 10638; 21 October 2015,
Yuan 10702; on fallen angiosperm twig, 21 October 2015,
Yuan 10692; Kuandian County, Baishilazi Nature Reserve,
on bark of rotten angiosperm branch, 22 October 2015, Yuan
10717; on rotten angiosperm wood debris, Yuan 10729.
GenBank numbers: ITS: KY686245, KY686244; LSU:
MK446416, MK446415.
Notes: Tomentella baroensis Yorou is similar to T. storea
by having thick basidiocarps (up to 2 mm thick), a smooth
to indistinctly granulose hymenophoral surface, the absence
of rhizomorphs and cystidia, and echinulate basidiospores of
approximately the same shape. However, the former species
is differentiated by dull red to greyish brown basidiocarps
adherent to the substrate, the absence of simple septa in
subicular hyphae, and bigger basidiospores (8–11 × 7.5–9
μm, Yorou et al. 2013). T. agbassaensis Yorou resembles T.
storea by having brown, continuous basidiocarps separable
from the substrate, a smooth and granulose hymenophore,
clamped, thick-walled subicular hyphae and echinulate
basidiospores of approximately the same shape and size.
However, it differs from T. storea by having rhizomorphs in
the subiculum and basidiospores with short echinuli (Yorou
et al. 2012).
Trechisporales K.H. Larsson
Notes: The order Trechisporales was proposed by Larsson (2007) and accepted in the Agaricomycetes based on
phylogenetic analysis (Binder et al. 2005; Larsson et al.
2004; Matheny et al. 2007). It includes stipitate, clavariod
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240
Fungal Diversity (2020) 104:1–266
Fig. 246 Microscopic structures
of Tomentella stipitata (IFP
019327, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
and resupinate genera with smooth, hydnoid, grandiniod
or poroid hymenia. Hyphae are clamped, forming a monomitic system, and the subicular hyphae may or may not have
ampullate septa. Basidia have four to six sterigmata, and
the basidiospores are smooth or ornamented. Some species
have cystidia. They grow mostly on wood or soil (Hibbett
et al. 2007).
13
Hydnodontaceae Jülich, Biblthca Mycol. 85: 372 (1981)
Notes: Hydnodontaceae was proposed by Jülich (1981)
and includes a variety of fruitbody shapes: coralloid, hydnoid, stipitate and resupinate. Currently accepted genera in
the family are: Brevicellicium, Dextrinocystis, Dextrinodontia, Hydnodon (Type), Litschauerella, Luellia, Porpomyces,
Scytinopogon, Sistotremastrum, Sphaerobasidium, Subulicystidium, Trechispora and Tubulicium (He et al. 2019).
Fungal Diversity (2020) 104:1–266
241
alutaceous, tan, tinged pink or purple, and have flat branches
in one plane, dilatating before branching, polychotomous in
the first divisions and dichotomous near to the tips, also it
can be dichotomous in slim specimens. This genus includes
10 species (www.indexfungorum.org).
Fig. 247 A basidiocarp of Tomentella storea (IFP 019330, holotype)
Fig. 248 SEM of basidiospores of Tomentella storea (IFP 019330,
holotype)
Scytinopogon Singer, Lloydia 8(3): 139 (1945)
Notes: The genus Scytinopogon was proposed by Singer
(1945). The clavarioid fruitbodies may be cream, white,
Scytinopogon minisporus J. Alvarez-Manjarrez, M. Villegas & R. Garibay-Orijel, sp. nov.
MycoBank number: MB829209; Facesoffungi number:
FoF 05672; Fig. 250
Etymology: Refers to the small-sized basidiospores.
Holotype: MEXU 28300.
Basidiocarps clavarioid 15–40 mm tall, branches are
15–17 mm long, stipe 5–15 × 1–3 mm long, acute to subounded, flat, and whitish or even grey (5A2–5B3), axils
rounded. Branching near to the stipe is polytomical and
near to the tips can be dichotomical to polytomical. Stipe
cylindrical slightly flattened, whitish to pale orange-brown
(5A2–5A3), base covered by numerous short hyphae looking
plushy. Surface of all the fruitbody, except the base of the
stipe, looks smooth to the naked eye, and dusty under the stereoscopic microscope. Consistency is cartilaginous to leathery. Flesh of middle portion has the same color as the surface.
Odor indistinguishable, taste slightly astringent. Basidiospores 4.2–5(–5.6) × 2.1–2.8(–3) µm [ x = 4.6 ± 0.4 × 2.4 ± 0.3
µm, Q = 1.3–1.6, n = 30], dacryoid, slightly elliptical in lateral
view, hyaline, thin-walled, verrucose, sometimes the warts
merge, without conical spines; plage non ornamented, with
lateral hilar appendix, slightly cyanophylic. Basidia mostly
tetrasporic, scarce, (10–)20.3–28 × 4.9–5.6 µm, cylindrical
to subcylindrical, hyaline, thick -and smooth-walled, base
with clamp connection. Sterigmata 2.8–4.2 × 1.4 µm, hyaline, straight with acute apex. Cystidia clavate, incrusted on
the tip, non dissolving in KOH, very scarce. Subhymenium
monomitic; generative hyphae of 1.4–3.5 × 50–56 µm, with
thickened wall (< 1 µm), septae with clamp connections, and
H connections between hyphae. Tramal hyphae strictly parallel, (2.1–)2.8–3.5 µm, thickened wall (< 1 µm), septate with
clamp connections, and hyphae with H connections. Stipe
composed of generative hyphae, 1.4–2.1 µm diam, hyaline,
wall slightly thickened, frequent septa with clamp connections, and abundant crystals on their surface forming irregular plates.
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Fungal Diversity (2020) 104:1–266
Fig. 249 Microscopic structures of Tomentella storea (IFP
019330, holotype). a Section through a basidiocarp. b
Basidiospores in frontal view. c
Basidiospores in lateral view
Habitat and known distribution: Gregarious or solitary,
growing on soil or debris of tropical dry forest from the
Pacific coast of Jalisco, Mexico.
Material examined: MEXICO, Jalisco, municipality La
Huerta, Estación de Biología de Chamela (EBCH), pathway Camino Antiguo, 9 October 2005, Villegas Ríos M.
2630 (FCME 26014); pathway Chachalaca, 11 August 2006,
Villegas Ríos M. 2679 (FCME 26015); pathway Camino
Antiguo Sur, 11 August 2006, Aguirre, Bautista and Pulido
II-40 (MEXU 26345); 1 October 1977, A. Pérez J. and A.
Solís M. (MEXU 11923); pathway Buho, 10 October 2015,
Alvarez-Manjarrez AM170 (MEXU 28300, holotype);
13
pathway Buho, 18 October 2015, Alvarez-Manjarrez AM176
(MEXU 28301, isotype).
GenBank numbers: ITS: MK328885, MK328886; LSU:
MK328894, MK328895.
Notes: Phylogenetic analyses of the combined ITS and
LSU dataset reveal Scytinopogon minisporus as sister to a
clade comprising two undescribed Scytinopogon species
from Australia and Taiwan island and S. aff. pallescens from
Mexico (Fig. 251). S. minisporus differs from other species of Scytinopogon in having clavate cystidia with crystals
on the tip. Microscopically it is very similar to S. scaber
(Berk. & M.A. Curtis) D.A. Reid, however S. minisporus has
Fungal Diversity (2020) 104:1–266
Fig. 250 Scytinopogon minisporus. a Fresh fruitbody (FCME
26015). b Dry fruitbody (MEXU 28300, holotype). c Basidiospores
with verrucose ornamentation, showing the plage without ornamentation (MEXU 28300, holotype). d Ornamentation of spores (MEXU
243
28301). e Immature basidiospores attached to a tetrasterigmata
basidia. f Clavate cystidia with crystals on the tip. Scale bars: b = 5
mm, c–e = 1.5 µm, f = 4 µm
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244
Fig. 251 Bayesian analysis combined dataset of ITS and LSU
sequence data of the Trechispora-Scytinopogon clade. The tree is
rooted with Brevicellicium atlanticum and B. olivascens. The node
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Fungal Diversity (2020) 104:1–266
support is indicated as maximum likelihood bootstrap and Bayesian
posterior probabilities respectively. New sequences are indicated in
blue
Fungal Diversity (2020) 104:1–266
Fig. 252 A basidiocarp of Phaeotremella yunnanensis (BJFC
019764, holotype)
verrucose spores, bigger basidia and encrusted cystidia. S.
papillosus Corner also is similar in spore size but has minute
papillae on the surface of the fruitbody. It is important to
note that Trechispora and Scytinopogon are paraphyletic in
the same clade (Birkebak et al. 2013; Desjardin and Perry
2015). The type sequences of both genera are necessary to
determine if the resupinate Trechispora and coralloid Scytinopogon are synonyms (Desjardin and Perry 2015). S.
minisporus is allocated to this genus, based on the current
concept while waiting for phylogenetic studies including
sequences of type species.
Tremellomycetes
Tremellales Fr.
Notes: The order was set up by Fries (1821), based on
Tremellaceae Fr. Most species of the order have gelatinous
basidiocarps with transverse or oblique basidial septa. However, some teleomorphic species apparently lack basidiocarps and some species grow as yeasts in their haploid stages
(Weiss et al. 2014; Liu et al. 2015b).
Phaeotremellaceae A.M. Yurkov & Boekhout, Stud. Mycol.
81: 137 (2015)
Notes: Phaeotremellaceae was introduced to accommodate the genus Phaeotremella Rea and the newly described
monotypic genus Gelidatrema Yurkov et al. in Liu et al.
(2015a). Phaeotremella was treated as the type genus of the
family and twelve species were accepted in the genus (Liu
et al. 2015a; Spirin et al. 2017). Gelidatrema was proposed
for the single species, Cryptococcus spencermartinsiae V.
de García et al. by Liu et al. (2015a), and a new species, G.
psychrophila M. Tsuji et al. was described in the genus by
Tsuji et al. (2017).
245
Phaeotremella Rea, Trans. Br. Mycol. Soc. 3: 377 (1912)
Notes: The genus was established by Rea (1912) with
one species, Phaeotremella pseudofoliacea Rea, but it was
later considered a synonym of Tremella foliacea Pers. (Donk
1966; Roberts 1999). The viewpoint was accepted by some
researchers and some species similar to T. foliacea were
described in Tremella Pers. (Donk 1966; Chen 1998; Pippola
and Kotiranta 2008; Boekhout et al. 2011; Weiss et al. 2014).
The genus Tremella was shown to be polyphyletic based on
molecular data and these species similar to T. foliacea (the
T. foliacea group) formed a single clade in the phylogenies
(Boekhout et al. 2011). Liu et al. (2015a) resurrected the
generic name Phaeotremella for the T. foliacea group, which
was recognized in later studies (Wedin et al. 2016; Spirin
et al. 2017).
Phaeotremella yunnanensis L.F. Fan, F. Wu & Y.C. Dai,
sp. nov.
Index Fungorum number: IF556263; Facesoffungi number: FoF 05842; Figs. 252, 253
Etymology: Referring to the distribution of the species in
Yunnan, China.
Holotype: BJFC 019764.
Basidiocarps gelatinous and no odor when fresh, foliaceous, 0.4–1.8 cm high, 1.6–4.0 cm diam, greyish brown to
fuscous, sessile, lobes with single-layer structure and undulate caespitose lobes arising directly from the central point
of attachment, becoming firmly gelatinous and deep brown
when dry. Hyphae clamped, thin- to slightly thick-walled,
2–9 μm diam, hyaline to brownish, inflated, anastomosing,
branched and slightly interwoven. Haustoria, hyphidia and
vesicles absent. Swollen cells rarely present. Basidia globose
to subglobose or widely ellipsoid, colorless, thin-walled,
longitudinally septate, rarely obliquely septate, with a basal
clamp connection, obvious oil drops and 1–4 long sterigmata, (11.0–)12.0–15.0(–22.0) × (11.0–)12.0–16.0(–18.0)
μm, L = 13.7 µm, W = 13.8 µm, Q = 1.07(n = 30/1), occasionally wider than long. Basidiospores globose or broadly
ellipsoid, hyaline, thin-walled, smooth, apiculate, with oil
drops, (6.0–)7.0–8.0(–8.9) × 6.0–7.3(–9.0) μm, L = 7.4 µm,
W = 6.9 µm, Q = 1.07 (n = 30/1), sometimes broader than
long, germinating and producing secondary spores by germ
tubes. Conidia usually plentiful, spherical or ovoid to ellipsoid, hyaline, thick-wall, smooth, usually originating from
inflated conidiophores interspersed among basidia in the
hymenium, (3.0–)3.8–5.0(–6.0) × (2.0–)2.5–3.8(–4.0) µm,
L = 4.5 µm, W = 3.0 µm, Q = 1.50 (n = 30/1).
Material examined: CHINA, Yunnan Province, Yongde
County, Daxueshan Nature Reserve, on dead angiosperm
branch, 27 August 2015, Dai 15660 (BJFC 019764, holotype); Jingdong County, Ailaoshan Nature Reserve, on
fallen angiosperm branch, 24 August 2015, Dai 15621
(BJFC 019725).
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246
Fig. 253 Microscopic structures
of Phaeotremella yunnanensis
(BJFC 019764, holotype). a
Basidiospores. b Conidia. c
Basidia. d Conidiophores with
conidia. e Hyphae from context
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Fungal Diversity (2020) 104:1–266
247
Fig. 254 Phylogenetic position
of Phaeotremella yunnanensis inferred from the ITS,
LSU and TEF1-α sequences.
Bootstrap support values (ML
and MP) ≥ 50% and Bayesian
posterior probabilities ≥ 0.90 are
given near nodes respectively.
The tree is rooted with Gelidatremea spencermartinsiae
(CBS10760T). The new isolates
are in bold
GenBank numbers: ITS: MK559397, MK559396; LSU:
MK559399, MK559398; TEF1-α: MK559401, MK559400.
Notes: In the phylogeny (Fig. 254), the new species is
closely related to Phaeotremella roseotincta (Lloyd) V.
Malysheva and P. fuscosuccinea (Chee J. Chen) Spirin &
Yurkov, but the former differs from P. yunnanensis by its
pinkish basidiocarps when fresh, slightly larger basidia
(16–20 × 11–18 µm) and basidiospores (7–10 × 7–9 µm); the
latter differs by its habits on fallen spruce associated with S.
sanguinolentum (Malysheva et al. 2018) and smaller basidia
(11–15 × 8–10.5 µm) (Chen 1998). Macro-morphologically,
P. yunnanensis resembles P. foliacea (Pers.) Wedin et al. in
the brownish tint, but differs from the new species by its
slightly narrower basidiospores (5.3–9.1 × 4.7–8.5 µm) and
the presence of swollen cells producing ellipsoid or subglobose conidial cells. In addition, P. foliacea has slightly
thick-walled basidia and grows on conifers associated with
Stereum sanguinolentum (Malysheva et al. 2018).
Acknowledgements Hai-Sheng Yuan would like to thank the National
Nature Science Foundation of China for the financial support (Project
Nos. 31770028, 31970017 and 31470148), the Special Funds for the
Young Scholars of Taxonomy of the Chinese Academy of Sciences
(Project No. ZSBR-015), and the Qingyuan Forest CERN (Chinese
Academy of Sciences) for supporting sampling in various ways. YuCheng Dai would like to thank the National Natural Science Foundation of China (Project Nos. U1802231) and the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (Grant No.
2019QZKK0503). Kevin D. Hyde would like to thank the Thailand
Research Fund for the grant “Macrofungi diversity research from the
Lancang-Mekong Watershed and surrounding areas contract” (No.
DBG6280009) for supporting this work. He also thanks Chiang Mai
University for the award of Visiting Professor. Ivana Kušan, Neven
Matočec and Margita Jadan are much obliged to Dušan Mrvoš for his
assistance during fieldwork in Gorski kotar region, Croatia, and to
Nedim Jukić (Mycological Society MYCOBH) for help during field
research in Bosnia and Herzegovina. Our studies were supported in
part by the Croatian Science Foundation under the project ForFungiDNA (IP-2018-01-1736). Bao-Kai Cui and Shuang-Hui He thank the
National Natural Science Foundation of China (Project Nos. 31750001
and 31670016) and Beijing Forestry University Outstanding Young
Talent Cultivation Project (No. 2019JQ03016) for financial support.
Monika C. Dayarathne would like to acknowledge the projects, viz.
National Natural Science Foundation of China (Project Nos. 31560489
and 31972222), Science and technology basic work of MOST
[2014FY120100], National Key Technology Research and Development Program of the Ministry of Science and Technology of China
(2014BAD23B03/03), Talent project of Guizhou science and technology cooperation platform ([2017]5788-5, [2019]5641) and Guizhou
science, technology department international cooperation base project
([2018]5806). Guangyu Sun would like to thank the National Natural
Science Foundation of China (Project Nos. 31772113, 31972220 and
31170015) and China Agriculture Research System (CARS-27) for
the financial support. Guangyu Sun also thanks Miss’s Hongcai Chen,
Wenhan Li, Wanyu Zhao for collecting samples. We also express our
thanks to Prof. Zide Jiang, South Agricultural University, for giving
helps in sampling. Shu-Yan Liu would like to thank the National Natural Science Foundation of China (Project Nos. 31670022, 31470153
and 31970019) and “111” Project (No. D17014) for financial support. Kamal C. Semwal and Vinod K. Bhatt would like to thank the
Uttarakhand State Council for Science and Technology (UCoST) for
financial support (Project No.UCSandT/RandD/LS-1/12-13/4912) on
“Collection, identification, documentation of wild edible and medicinal
mushrooms of Garhwal Himalaya of Uttarakhand”. Roberto Garibay
Orijel thanks the CONACYT for funding fieldwork and DNA sequencing through the project CONACYT Ciencia Basica 239266. Jadson
D.P. Bezerra, Camila M Gonçalves and C.M. Souza-Motta thank
FACEPE, CNPq and CAPES (Finance code 001). Fang Wu and LongFei Fan would like to thank the National Natural Science Foundation
13
248
of China (Project No: 31701978). Tatiana Baptista Gibertoni would
like to thank the staff of URM for helping with the exsiccates; Renato
Alvarenga for preparing the images of the Brazilian specimens; Adriana Gugliotta from the Instituto de Botânica (SP), Daniel Lindner and
Beatriz Ortiz-Santana from the Center for Forest Mycology Research
(CFMR), Matthew Smith and Barry Kaminsky from the University
of Florida (FLAS), Matthew Pace from the New York Botanical Gardens (NY), Jordan Teisher from the Academy of Natural Sciences
(PH), and Alan Rockefeller for sharing the images of the materials;
Elaine Malosso, Gladstone Silva, Juliano Baltazar and Leonor Maia
for suggestions; CNPq, CAPES, FACEPE and the Pós-Graduação em
Biologia de Fungos (UFPE) for financing this research and providing
scholarships to the authors. Tie-Zheng Wei would like to thank the
National Natural Science Foundation of China (Project No. 31270072),
the Special Funds for the Young Scholars of Taxonomy of the Chinese
Academy of Sciences (ZSBR-001) and National Key Basic Research
Special Foundation of China (2013FY110400). The authors extend
the appreciation to the International Scientific Partnership Program
ISPP at King Saud University for funding this research work through
ISPP#0089. Yusufjon Gafforov thanks the financial research support by
the Ministry of Innovative Development of the Republic of Uzbekistan
(Project No. P3-2014-0830174425 and PЗ-20170921183), CAS President’s International Fellowship Initiative (PIFI) for Visiting Scientist
(Grant No. 2018VBB0021) and German Academic Exchange Service
(DAAD) for a Visiting Fellowship (Grant No. 57314018). LWZ thanks
to the financial support by the National Natural Science Foundation of
China (Project No. 31970012) and the Youth Innovation Promotion
Association of Chinese Academy of Sciences (Project No. 2017240).
Jun-Zhi Qiu thanks the National Natural Science Foundation of China
(Project Nos. U1803232 and 31670026) and National Key R & D Program of China (2017YFE0122000) for financial support. Carlos AF de
Souza, Diogo X Lima, André LCM de Azevedo Santiago express their
gratitude to the Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES) and Fundação de Amparo à Ciência e Tecnologia
do Estado de Pernambuco (FACEPE) for postgraduate scholarships
provided to Diogo X. Lima and Carlos A. F. de Souza, respectively.
We also thank the Conselho Nacional de Desenvolvimento Científico
e Tecnológico (CNPq) for the Master scholarship provided to André
Luiz C. M de A. Santiago. This manuscript was financed by the project
‘Diversity of Mucoromycotina in different ecosystems of the Pernambuco’s Atlantic Rainforest’ (FACEPE—APQ 0842-2.12/14). Rungtiwa
Phookamsak thanks CAS President’s International Fellowship Initiative (PIFI) for Young Staff 20 19-2021 (Grant No. 2019FY0003), the
Research Fund from China Postdoctoral Science Foundation (Grant No.
Y71B283261), the Yunnan Provincial Department of Human Resources
and Social Security (Grant No. Y836181261), and the National Science
Foundation of China (Project No. 31850410489) for financial research
support. The work of Bálint Dima was partly supported by the ELTE
Institutional Excellence Program financed by the National Research,
Development and Innovation Office (NKFIH-1157-8/2019-DT).
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Affiliations
Hai‑Sheng Yuan1,2 · Xu Lu1,2 · Yu‑Cheng Dai3 · Kevin D. Hyde4,5,6,7,8 · Yu‑He Kan1,2 · Ivana Kušan9 · Shuang‑Hui He3 ·
Ning‑Guo Liu5,10 · V. Venkateswara Sarma11 · Chang‑Lin Zhao12 · Bao‑Kai Cui3 · Nousheen Yousaf13 · Guangyu Sun14 ·
Shu‑Yan Liu15,31 · Fang Wu3 · Chuan‑Gen Lin5,6,7 · Monika C. Dayarathne5,27 · Tatiana Baptista Gibertoni16 ·
Lucas B. Conceição17 · Roberto Garibay‑Orijel18 · Margarita Villegas‑Ríos19 · Rodolfo Salas‑Lizana19 ·
Tie‑Zheng Wei20 · Jun‑Zhi Qiu21 · Ze‑Fen Yu22 · Rungtiwa Phookamsak4,5,8 · Ming Zeng4,5 · Soumitra Paloi23 ·
Dan‑Feng Bao5,24,25 · Pranami D. Abeywickrama5,6,26 · De‑Ping Wei4,5,7,25 · Jing Yang5 · Ishara S. Manawasinghe5,26 ·
Dulanjalee Harishchandra5,6,26 · Rashika S. Brahmanage5,6,26 · Nimali I. de Silva4,8,28,29 · Danushka S. Tennakoon4,5,6 ·
Anuruddha Karunarathna5,30 · Yusufjon Gafforov1,32,33 · Dhandevi Pem5 · Sheng‑Nan Zhang5,30 ·
André L. C. M. de Azevedo Santiago34 · Jadson Diogo Pereira Bezerra35 · Bálint Dima36 · Krishnendu Acharya23 ·
Julieta Alvarez‑Manjarrez18,58 · Ali H. Bahkali37,38 · Vinod K. Bhatt39 · Tor Erik Brandrud40 · Timur S. Bulgakov41 ·
E. Camporesi42,43,44 · Ting Cao1,2 · Yu‑Xi Chen21 · Yuan‑Yuan Chen45 · Bandarupalli Devadatha11 ·
Abdallah M. Elgorban37,38 · Long‑Fei Fan3 · Xing Du22 · Liu Gao14 · Camila Melo Gonçalves46 · Luis F. P. Gusmão17 ·
Naruemon Huanraluek5 · Margita Jadan9 · Ruvishika S. Jayawardena5 · Abdul Nasir Khalid47 · Ewald Langer33 ·
Diogo X. Lima34 · Nelson Correia de Lima‑Júnior48 · Carla Rejane Sousa de Lira16 ·
Jian‑Kui (Jack) Liu49 · Shun Liu3 · Saisamorn Lumyong28,29,50 · Zong‑Long Luo24 · Neven Matočec9 · M. Niranjan11 ·
José Ribamar Costa Oliveira‑Filho16 · Viktor Papp51 · Eduardo Pérez‑Pazos19,52 · Alan J. L. Phillips53 ·
Peng‑Lei Qiu15 · Yihua Ren14 · Rafael F. Castañeda Ruiz54 · Kamal C. Semwal55 · Karl Soop56 · Carlos A. F. de Souza34 ·
Cristina Maria Souza‑Motta46 · Li‑Hua Sun57 · Meng‑Le Xie31 · Yi‑Jian Yao20 · Qi Zhao4 · Li‑Wei Zhou1
11
* Yu-Cheng Dai
yuchengd@yahoo.com
Department of Biotechnology, School of Life
Sciences, Pondicherry University, R.V. Nagar, Kalapet,
Pondicherry 605014, India
12
* Kevin D. Hyde
kdhyde3@gmail.com
College of Biodiversity Conservation, Southwest Forestry
University, Kunming 650224, People’s Republic of China
13
Department of Botany, Govt. College University, Katchery
Road, Lahore, Pakistan
14
State Key Laboratory of Crop Stress Biology in Arid Areas
and College of Plant Protection, Northwest A&F University,
Yangling, Shaanxi, People’s Republic of China
* Hai-Sheng Yuan
hsyuan@iae.ac.cn
1
CAS Key Laboratory of Forest Ecology and Management,
Institute of Applied Ecology, Chinese Academy of Sciences,
Shenyang 110164, People’s Republic of China
2
University of the Chinese Academy of Sciences,
Beijing 100049, People’s Republic of China
15
3
School of Ecology and Nature Conservation, Beijing
Advanced Innovation Center for Tree Breeding By Molecular
Design, Beijing Forestry University, Beijing 100083,
People’s Republic of China
Laboratory of Plant Pathology, College of Plant Protection,
Jilin Agricultural University, Changchun 130118,
Jilin Province, People’s Republic of China
16
CAS Key Laboratory for Plant Diversity and Biogeography
of East Asia, Kunming Institute of Botany, Chinese
Academy of Science, Kunming 650201, Yunnan,
People’s Republic of China
Centro de Biociências, Departamento de Micologia,
Universidade Federal de Pernambuco, Cidade Universitária,
Avenida da Engenharia s/n, Recife, Pernambuco 50740-600,
Brazil
17
5
Departamento de Ciências Biológicas, Universidade
Estadual de Feira de Santana, Av. Transnordestina s/n, Novo
Horizonte, Feira de Santana, BA 44036-900, Brazil
Centre of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
18
6
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
Laboratory of Systematics and Ecology of Mycorrhizae,
Institute of Biology, Universidad Nacional Autónoma de
México, 04510 Mexico City, Mexico
19
7
Mushroom Research Foundation, 128 M.3 Ban Pa Deng
T. Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
Laboratories of Mycology, School of Sciences, Universidad
Nacional Autónoma de México, 04510 Mexico City, Mexico
20
8
East and Central Asia Regional Office, World Agroforestry
Centre (ICRAF), Kunming 650201, Yunnan, China
State Key Laboratory of Mycology, Institute
of Microbiology, Chinese Academy of Sciences,
Beijing 100101, People’s Republic of China
9
Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb,
Croatia
21
10
Faculty of Agriculture, Natural Resources and Environment,
Naresuan University, Phitsanulok 65000, Thailand
State Key Laboratory of Ecological Pest Control for Fujian
and Taiwan Crops, College of Life Sciences, Fujian
Agriculture and Forestry University, Fuzhou 350002, Fujian,
People’s Republic of China
4
13
266
Fungal Diversity (2020) 104:1–266
22
State Key Laboratory for Conservation and Utilization
of Bio-Resources in Yunnan, Yunnan University,
Kunming 650091, People’s Republic of China
23
Molecular and Applied Mycology and Plant Pathology
Laboratory, Department of Botany, University College
of Science and Agriculture, University of Calcutta,
Kolkata 700019, West Bengal, India
College of Agriculture& Biological Sciences, Dali
University, Dali 671003, Yunnan, People’s Republic of China
24
25
Department of Entomology and Plant Pathology, Faculty
of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
40
Norwegian Institute for Nature Research (NINA),
Gaustadalléen 21, 0349 Oslo, Norway
41
Department of Plant Protection, Federal Research
Centre the Subtropical Scientific Centre of the Russian
Academy of Sciences, Yana Fabritsiusa Street 2/28, Sochi,
Krasnodar Region, Russia 354002
42
A.M.B, Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314,
Brescia, Italy
43
A.M.B. Gruppo, Micologico Forlivese ‘‘Antonio Cicognani’’,
Via Roma 18, Forlí, Italy
44
Società Per Gli Studi Naturalistici Della Romagna, C.P. 143,
Bagnacavallo, RA, Italy
45
College of Forestry, Henan Agricultural University,
Zhengzhou 450002, Henan, People’s Republic of China
46
Departamento de Micologia Prof. Chaves Batista,
Universidade Federal de Pernambuco, Cidade Universitária,
Av. da Engenharia, s/n, Recife, PE, Brazil
47
Department of Botany, University of the Punjab, Lahore,
Pakistan
48
Laboratório Bioquímica Geral, Centro Acadêmico de
Vitória, Universidade Federal de Pernambuco, Rua Alto
Do Reservatório S/N, Bela Vista, Vitória de Santo Antão,
Pernambuco 55600-000, Brazil
26
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences,
Beijing 100097, People’s Republic of China
27
Department of Plant Pathology, Agriculture College,
Guizhou University, Guiyang 550025, Guizhou,
People’s Republic of China
28
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
29
Research Center of Microbial Diversity and Sustainable
Utilization, Faculty of Science, Chiang Mai University,
Chiang Mai 50200, Thailand
30
Department of Entomology and Plant Pathology, Faculty
of Agriculture, Chiang Mai University, Chiang Mai, Thailand
49
31
Engineering Research Center of Chinese Ministry
of Education for Edible and Medicinal Fungi,
Jilin Agricultural University, Changchun 130118,
People’s Republic of China
School of Life Science and Technology, University
of Electronic Science and Technology of China,
Chengdu 611731, People’s Republic of China
50
Academy of Science, The Royal Society of Thailand,
Bangkok 10300, Thailand
51
Department of Botany, Faculty of Horticultural Science,
Szent István University, P.O. Box 53, 1518 Budapest,
Hungary
52
Posgrado en Ciencias Biológicas, Universidad Nacional
Autónoma de México, 04510 Mexico City, Mexico
53
Faculdade de Ciências, Biosystems and Integrative Sciences
Institute (BioISI), Universidade de Lisboa, Campo Grande,
1749-016 Lisbon, Portugal
54
Instituto de Investigaciones Fundamentales en Agricultura
(INIFAT), Tropical‘Alejandro de Humboldt’, OSDE,
Grupo Agrícola, Calle 1 Esq. 2, Santiago de Las Vegas,
C. Habana C.P. 17200, Cuba
55
Department of Biology, College of Sciences, Eritrea Institute
of Technology, Mai Nafhi, Asmara, Eritrea
56
Department of Botany, Swedish Museum of Natural History,
P.O. Box 50007, 104 05 Stockholm, Sweden
57
Faculty of Biological Sciences & Technology, Baotou
Teachers’ College, Baotou 014030, Inner Mongolia,
People’s Republic of China
58
Posgrado en Ciencias Biológicas. Edificio B, 1º
Piso. Circuito de Posgrados, Ciudad Universitaria. Coyoacán,
04510 Mexico City, Mexico
32
Laboratory of Mycology, Institute of Botany, Academy
of Sciences of the Republic of Uzbekistan, 32 Durmon Yuli
Street, Tashkent, Uzbekistan 100125
33
Department of Ecology, University of Kassel,
Heinrich-Plett-Strasse, 40, 34132 Kassel, Germany
34
Department of Mycology, Federal University of Pernambuco,
Av. Prof. Nelson Chaves, s/n, Recife, PE 50670-420, Brazil
35
Setor de Micologia, Departamento de Biociências E
Tecnologia, Instituto de Patologia Tropical E Saúde Pública
(IPTSP), Universidade Federal de Goiás (UFG), Rua 235,
s/n, Setor Universitário, Goiânia, Goiás CEP 74605-050,
Brazil
36
Department of Plant Anatomy, Institute of Biology,
Eötvös Loránd University, Pázmány Péter sétány 1/C,
1117 Budapest, Hungary
37
38
39
Center of Excellence in Biotechnology Research, King Saud
University, P.O. Box. 2455, Riyadh 11451, Saudi Arabia
Department of Botany and Microbiology, College
of Sciences, King Saud University, P.O. Box. 2455,
Riyadh 11451, Saudi Arabia
Navdanya, 105, Rajpur Road, Dehradun, Uttarakhand, India
13