MYCOTAXON
THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY e& NOMENCLATURE
VOLUME 130(2) APRIL-JUNE 2015
5 um
Mycorrhaphium pusillum (redescribed) (Tervonen, Spirin, & Halme— Fie. 3, p. 553) VIACHESLAV SPIRIN artist
ISSN (PRINT) 0093-4666 http://dx.doi.org/10.5248/130-2 ISSN (ONLINE) 2154-8889 MYXNAE 130(2): 307-600 (2015)
EDITORIAL ADVISORY BOARD
SABINE HUHNDORE (2011-2016) , Chair Chicago, Illinois, U.S.A.
Scott A. REDHEAD (2010-2015), Past Chair Ottawa, Ontario, Canada
PETER BUCHANAN (2011-2017) Auckland, New Zealand BRANDON MATHENY (2013-2018) Knoxville, Tennessee, U.S.A.
KAREN HANSEN (2014-2019) Stockholm, Sweden
Published by MycoTaxon, LTD. P.O. BOX 264, ITHACA, NY 14581-0264, USA
www.mycotaxon.com & www.ingentaconnect.com/content/mtax/mt © Mycotaxon, LTp, 2015
MYCOTAXON
THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY & NOMENCLATURE
VOLUME 130(2)
APRIL-JUNE 2015
EDITOR-IN-CHIEF
LORELEI L. NORVELL
editor@mycotaxon.com Pacific Northwest Mycology Service 6720 NW Skyline Boulevard Portland, Oregon 97229-1309 USA
NOMENCLATURE EDITOR
SHAUN R. PENNYCOOK
PennycookS@LandcareResearch.co.nz Manaaki Whenua Landcare Research Auckland, New Zealand
CONSISTING OF PAGES I-XII + 307-600, INCLUDING FIGURES
ISSN 0093-4666 (PRINT) http://dx.doi.org/10.5248/130-2.cvr ISSN 2154-8889 (ONLINE)
© 2015. MycoTAxon, LTD.
IV ... MYCOTAXON 130(2)
MYCOTAXON
VOLUME ONE HUNDRED THIRTY (TWO) — TABLE OF CONTENTS
COVER SECTION
PREVICW CHES 4p 2b, lade Aee deed asicck PROM ich PA ce told ob toon oA GALS Mn Meehan ic vii 1 ey ETH LD NRE Rel eG ae Sa nS A ee pe ee Slat, Aa Sh See OS eet at, RAS eS viii BOW! Tie TE HOO. 34s 2 us Sak Ses GRO oye WES oo Bid oa ea Ka a cae CMe na ix SUD SSIONE DRO CCU NCS x EA Ny en, Sy Mee Pe AR ER oly Se age ERS xi
RESEARCH ARTICLES
A new species of Neosporidesmium from Hainan, China XIANG-YU LI, SHU-YAN LIU, & XIU-GUO ZHANG A new species of Matsushimiella from submerged leaves in the Brazilian Amazon Forest JOSIANE SANTANA MONTEIRO, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL FE. CASTANEDA-RUIZ Amazonian phalloids: new records for Brazil and South America TIARA S. CABRAL, CHARLES R. CLEMENT, & IURI G. BASEIA On the generic names Kriegeria MERJE TOOME, SHAUN R. PENNYCOOK, & M. CATHERINE AIME New records of crustose lichens and a lichenicolous Arthonia
from Vietnam SANTOSH JOSHI, DALIP KUMAR UPRETI, SOON-OK OH, Tut THuy NGUYEN, ANH DzuNG NGUYEN, & JAE-SEOUN HuR First records of some Asian macromycetes in Africa PaBLo P. DANIELS,
OuMmAROU HAMA, ALFREDO JUSTO FERNANDEZ, FELIX INFANTE GARCIA-PANTALEON, Moussa BARAGE, DAHIRATOU IBRAHIM, & MARIA Rosas ALCANTARA Contribution to the knowledge of Inonotus baumii in Thailand ANON AUETRAGUL, ORATAI EUATRAKOOL, Maria LETIZIA GARGANO, & GIUSEPPE VENTURELLA Taxonomic studies in Chrysoderma, Corneromyces, Dendrophysellum, Hyphoradulum, and Mycobonia Karen K. NAKASONE A new Graphilbum species from western hemlock (Tsuga heterophylla) in Canada JAMES REID & GEORG HAUSNER Molecular analyses reveal a new species in Melanoderma from tropical China HaI-SHENG YUAN & YU-HE Kan Graphis hongkongensis sp. nov. and other Graphis spp. new to Hong Kong WEI GUO & JAE-SEOUN Hur Two new conidial fungi from Chapada Diamantina, Brazil CAROLINA RIBEIRO SILVA, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL F. CASTANEDA-RUIZ
307
371
315
321
329
337
361
369
399
421
429
437
APRIL-JUNE 2015... V
Anaexserticlava caatingae, a new conidial fungus from the
semi-arid Caatinga biome of Brazil TasciaNo Dos SANTOS SANTA IZABEL, Davi AUGUSTO CARNEIRO DE ALMEIDA, JOSIANE SANTANA MONTEIRO, Marcos FABIO OLIVEIRA MARQUES, Luis FERNANDO PASCHOLATI GUSMAO,
& RAFAEL E CASTANEDA-RUIZ 445 A new species of Endophragmiella from Guizhou, China YING-Rut Ma, JI- WEN XIA, & XIU-GUO ZHANG 451 Peziza succosella and its ectomycorrhiza associated with Cedrus deodara from Himalayan moist temperate forests of Pakistan SANA JABEEN, TAYIBA ASHRAF, & ABDUL NASIR KHALID 455 Cryptocoryneum parvulum, a new species on Araucaria angustifolia (Brazilian pine) SILVANA SANTOS DA SILVA, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL FE. CASTANEDA-RUIZ 465 Two new species and a new record of Leptogium from China Hua-JrE Lru, MAN-QING XI, JIAN-SEN Hu, & QING-FENG WU 471 Digicatenosporium polyramosum, a new hyphomycete from Brazil SHEILA MIRANDA LEAO-FERREIRA, Luis FERNANDO PASCHOLATI GUSMAO, Davi AUGUSTO CARNEIRO DE ALMEIDA, & RAFAEL FE CASTANEDA-RUIZ 479 Duportella lassa sp. nov. from Northeast Asia VIACHESLAV SPIRIN & JIRE KouT 483 Brachycephala exotica, a new hyphomycete from Brazil JOSIANE SANTANA MONTEIRO, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL EF CASTANEDA-RUIZ 489 A new species of Diplococcium from the Brazilian semi-arid region Davi AUGUSTO CARNEIRO DE ALMEIDA, TASCIANO Dos SANTOS SANTA IZABEL, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL F, CASTANEDA-RUIZ 495 Distophragmia, a new genus of microfungi to accommodate Endophragmiella rigidiuscula RAFAEL FE, CASTANEDA-RUIZ, SHEILA MIRANDA LEAO-FERREIRA, & LUIS FERNANDO PASCHOLATI GUSMAO 499 A new species of Chaetochalara on decaying leaves from Brazil SILVANA SANTOS Da SILVA, CAROLINA RIBEIRO SILVA, Luis FERNANDO PASCHOLATI GUSMAO, & RAFAEL FE. CASTANEDA-RUIZ 505 Two new species of Spadicoides and Sporidesmiella from Yucatan, Mexico GABRIELA HEREDIA, MARCELA GAMBOA-ANGULO, Rosa M. ARIAS, & RAFAEL F. CASTANEDA-RUIZ 511 Pholiota olivaceophylla, a forgotten name for a common snowbank fungus, and notes on Pholiota nubigena NOAH SIEGEL, NHU H. NGUYEN, & ELSE C. VELLINGA 517 Leucoagaricus lahorensis, a new species of L. sect. Rubrotincti T. Qasim, T. Amir, R. Nawaz, A.R. NIAzI, & ALN. KHALID 533
v1 ... MYCOTAXON 130(2)
Aspicilia volcanica, a new saxicolous lichen from Northeast China GULBOSTAN ISMAYIL, ABDULLA ABBAS, & SHOU-YU GUO Redescription of Mycorrhaphium pusillum, a poorly known hydnoid fungus Kaisa TERVONEN, VIACHESLAV SPIRIN, & PANU HALME Micropsalliota pseudoglobocystis, a new species from China Li WEI, YONG-HE L1, KEvIN D. HyDE, & Ru1-LIN ZHAO Two new Rosellinia species from Southwest China QztrRuI LI, JICHUAN KANG, & KEVIN D. HYDE Some new records of Uredinales from Khyber Pakhtunkhwa, Pakistan M. Fraz, A. Hasrs, N.S. AFSHAN, & A.N. KHALID The genus Allocetraria (Parmeliaceae) in China Ru1-FANG WANG, XIN-LI WEI, & JIANG-CHUN WEI BOOK REVIEWS AND NOTICES ELsE C. VELLINGA (EDITOR) NOMENCLATURAL NOVELTIES AND TYPIFICATIONS PROPOSED IN MYCOTAXON 130(2)
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577 593
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APRIL-JUNE 2015 ...
REVIEWERS — VOLUME ONE HUNDRED THIRTY (TWO)
The Editors express their appreciation to the following individuals who have, prior to acceptance for publication, reviewed one or more of the papers
prepared for this issue.
N.S. AFSHAN
JOE AMMIRATI JULIANO M. BALTAZAR UwE BRAUN
H.H. BuRDSALL, JR.
R. E CASTANEDA-RUIZ Yu-CHENG Dal KANAD Das
DENNIS E. DESJARDIN MarTIN ESQUEDA WALTER GAMS ZAI-WEI GE
J. GINNS
ToM GRAFENHAN Luis EP. GusMAOo Lru-Fu HAN SHUANGHUI HE JAE-SEOUN Hur OMOANGHE ISHIKHUEMHEN UDENI JAYALAL ZE-FENG JIA
SANTOSH JOSHI
BRYCE KENDRICK
S. KONDRATYUK
HEIKKI KOTIRANTA
SHAMBHU KUMAR
T. K. ARUN KUMAR DeE-WEI LI
LAszL6 LOxK6s THORSTEN LUMBSCH Eric H.C. McKENZIE Davin J. MCLAUGHLIN GABRIEL MORENO ABDUL REHMAN NIAZI TUOMO NIEMELA LORELEI L. NORVELL CLARK L. OVREBO OMAR PAiNno PERDOMO SHAUN R. PENNYCOOK DONALD H. PFISTER ScoTT REDHEAD
Amy Y. ROSSMAN INDREK SELL RAGHVENDRA SINGH JAN VONDRAK
Norou SULEIMEINE YOURO ZE-FEN YU
EUGENE O. YURCHENKO GEorRGIOS I. ZERVAKIS X1u-Guo ZHANG
VII
vill ... MYCOTAXON 130(2)
ERRATA FROM PREVIOUS VOLUMES
VOLUME 123
p. 479, line 2 for: NGUYEN ANH DZUNG
VOLUME 124
p. 51, lines 2-3 for: NGUYEN ANH DZUNG p. 309, line 2 for: NGUYEN ANH DZUNG
VOLUME 125
p. 69, line 3 for: NGUYEN ANH DZUNG
VOLUME 130(1)
p. 22, line 11 FOR: Q = 1.29-1.39 p. 61, line 9 FOR: School of Life Science p. 61, line 13 FOR: Academia of Agriculture
read: ANH DZUNG NGUYEN
read: ANH DzZUNG NGUYEN read: ANH DZUNG NGUYEN
read: ANH DZUNG NGUYEN
READ: Q = 1.33 READ: College of Life Science READ: Academy of Agriculture
PUBLICATION DATE FOR VOLUME ONE HUNDRED THIRTY (ONE)
MYCOTAXON for JANUARY-MARCH, (I-v1 + 1-306) was issued on April 22, 2015
APRIL-JUNE 2015 ... IX
FROM THE EDITOR-IN-CHIEE
MYCOTAXON 130(2) contains 32 research papers by 112 authors (representing 18 countries) and reviewed by 50 expert reviewers. Within its pages are four new genera (Anaexserticlava, Brachycephala, Distophragmia, and Phaeoschizotrichum) and 26 species new to science representing Allocetraria, Aspicilia, Endophragmiella, Graphis, Leptogium, Melanoderma, Micropsalliota, Neosporidesmium, & Rosellinia from China; Anaexserticlava, Brachycephala, Cryptocoryneum, Chaetochalara, —_ Digicatenosporium, —_ Diplococcium, Distophragmia, Matsushimiella, Phaeoschizotrichum, & _ Pyriculariopsis from Brazil; Duportella from Northeast Asia; Graphilbum from Canada; Leucoagaricus from Pakistan; and Spadicoides & Sporidesmiella from Mexico.
In addition to range extensions (in Brazil, Niger, Pakistan, Thailand, & Vietnam) and/or new hosts for previously named taxa, we also offer 6 new combinations (in Corneromyces, Distophragmia, Polyporus, Pseudolagarobasidium, & Vararia), a convincing argument that Kriegeria is a legitmate basidiomycete (not ascomycete) name, a revival of a ‘forgotten’ name for acommon western North American pholiota, and a re-description of a rare Mycorrhaphium.
As we noted in MycoTaxon 129(2) (the 2014 October-December issue), our journal is temporarily without a permanent book review editor. Nonetheless, Else Vellinga has generously returned to serve as a guest editor in this issue. Thanks yet again, Else!
NOMENCLATURE OF AUTHORS AND AUTHORITIES— As an international journal that publishes papers in English, MycoTAxon is not immune to nomenclatural confusion across international borders, particularly when it comes to distinguishing the given or generational names from the surname—that all-important ‘family’ name that determines which of several names will be spelled in full in abbreviated bibliographic references. We usually manage to decipher the author names correctly, although it usually takes us one to two months to override the stubborn Ingenta algorithm that insists on listing all Brazilian and Portuguese names following the Spanish convention.
Asian names — in particular Chinese, Korean, and Vietnamese — present a special challenge. For over two years, we have incorrectly displayed one author’s name, unfortunately for Nguyen Anh Dzung (as it appears in proper Vietnamese). Prof. Nguyen first served as co-author in MycoTAxoOn 117, where his name appeared as Anh Dzung Nguyen (properly abbreviated and indexed
x ... MYCOTAXON 130(2)
as Nguyen AD). Unaccustomed to seeing his name in the ‘English’ style, Prof. Nguyen contacted us to say that his name was Nguyen Anh Dzung. Although well aware that Nguyen is a common Vietnamese surname, we did not know whether it could also serve as a given name. Thus MycotTaxon followed the author’s wishes by displaying his name in his next four papers in the Vietnamese order and indexing his name as Dzung NA. With this issue, however, we establish that Nguyen is, in fact, his surname. With past sins corrected in the ERRATA On p. Vit, Prof. Nguyen has kindly agreed to follow our ‘English’ policy in all future papers.
Other confusions surround how to abbreviate or decode the name of a nomenclatural authority. MycoTAXxoNn asks its authors to adhere to the INDEX FUNGORUM list of author abbreviations covering researchers who have previously named fungi. This web-accessible list is handy both for authors and for readers needing to decode the abbreviations of the authorities that stand after the names of taxa in a full scientific name. An author proposing a new taxon for the first time need use initials before the surname ONLY if previous authorities have had the same surname. [I consider being known only as ‘Norvell’ both simpler and a bit of an honor!] Those who find that their surname has already been ‘taken’ should follow the established abbreviation (e.g., “Sm? for ‘Smith, and add their initials. In instances where two authors share the same initials, then one of the given names should be spelled out. Where nomenclatural authors share all the same names, they are separated by ‘bis’ (for 2), ‘tertia’ (for 3), and so on. Be forewarned — as botanists, phycologists, and mycologists are governed by the same code, each surname can stand alone only once. Which explains why the late Meinhard Moser is designated by ‘M.M. Moser’ and not simply Moser, which refers to the economic botanist Heinrich Christoph Moser, active around 1794.
What’s in a name, indeed? Warm regards,
Lorelei L. Norvell (Editor-in-Chief) 30 June 2015
APRIL-JUNE 2015 ... XI
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ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
- MYCOTAXON
http://dx.doi.org/10.5248/130.307 Volume 130, pp. 307-310 April-June 2015
A new species of Neosporidesmium from Hainan, China
XIANG-YU LI*?, SHU-YAN Liu?“ & XIU-GUO ZHANG? ®
' Department of Plant Pathology, Jilin Agricultural University, Changchun, 130118, China ? Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China
*CORRESPONDENCE TO: “liussyan@163.com; *zhxg@sdau.edu.cn, sdau613@163.com
ABSTRACT — Neosporidesmium diaoluoshanense sp. nov., collected on rotten branches from tropical forest of Hainan Province, China, is described and illustrated. It differs from previously described Neosporidesmium species in producing conidia that possess a colourless, globose, mucilaginous sheath at the apex.
KEY worps — asexual fungi, key, taxonomy
Introduction
The genus Neosporidesmium was established by Mercado & Mena (1988) for a single species, N. maestrense. Subsequently seven other species have been described: N. sinensis and N. microsporum (Wu & Zhuang 2005); N. antidesmatis, N. malloti, and N. xanthophylli (Ma et al. 2011a); N. micheliae (Zhang et al. 2011a); and N. vietnamense (Melnik & Braun 2013). The genus is characterized by effuse, dark brown to black, hairy colonies, unbranched conidiophores aggregating into erect, cylindrical, dark brown to black synnemata and integrated, terminal, monoblastic, lageniform or doliiform, determinate or percurrently proliferating conidiogenous cells that produce solitary, pale brown to brown, obclavate or cylindrical, smooth, distoseptate conidia. The criteria used for species delimitation are mainly based on proliferations of conidiogenous cells, conidial morphological characteristics, and conidiomatal overall size (Mercado & Mena 1988, Wu & Zhuang 2005).
The tropical forests of southern China have a rich mycobiota, and many new species have been discovered there (Zhang et al. 2009a,b, 2011b, Ma et al. 2011a,b, 2012). During a continuing investigation on saprobic microfungi from tropical forest of Hainan Province, a morphologically distinct species of Neosporidesmium was found. It is proposed here as a new species.
308 ... Li, Liu, & Zhang
Neosporidesmium diaoluoshanense Xiang Y. Li & X.G. Zhang, sp. nov. FIG. 1 MycoBank MB 812118 Differs from other Neosporidesmium species in its conidia having an apical, colourless,
globose, mucilaginous sheath.
Type: China, Hainan Province: Diaoluoshan Nature Reserve, on rotten branches of an unidentified tree, 10 Apr. 2014, H.X. Dong (Holotype HSAUP H7562; isotype, HMAS 243460).
EryMo_oey: in reference to the type locality.
Co.ontgs on the natural substrate effuse, solitary, dark brown, hairy. Mycelium partly superficial, partly immersed in the substratum, composed of dark brown to black, septate, cylindrical hyphae. ConIDIOMATA synnemata, indeterminate, scattered, erect, up to 605 um high, 40-60 um wide, composed of straight, unbranched, septate, smooth, brown to dark brown conidiophores 3-6 um wide, diverging terminally and laterally. CoNIDIOGENOUS CELLS monoblastic, integrated, determinate, smooth, cylindrical, doliiform or lageniform, brown to dark brown, 4-10 x 9-15 um. Conidial secession schizolytic. Conrp1a solitary, acrogenous, obclavate, base truncate, 8-11-distoseptate, 105-135 x 10.5-13.5 um, 2.5-3.5 um wide at the truncate base, rounded at the apex, dry, smooth- walled, pale brown, but hyaline at the apical cells, with an apical, globose, hyaline, 10.5-13.5 um diam., mucilaginous sheath.
ComMENTsS - Neosporidesmium diaoluoshanense differs from all previously described species in the genus by producing conidia with an apical, globose, mucilaginous, hyaline sheath and having scattered synnemata. The new species resembles N. micheliae, which differs by having conidia with fewer septa (7-8) and lacking a mucilaginous sheath.
Key to Neosporidesmium species
l.. Conidiogenous cells-with percurrent prolifetation: sc .::.32sess eee dee oe 2 Conidiogenous cells without percurrent proliferation ...................04. 4 2h. Wy Cotidiamnearlyoylindricale Wis siot sano ttle ashen busethae Restathoo ack N. microsporum @onidia.obelavate 2.5. bm the Ame ke me Meee Am a GB head 3 3.. gC onidiasvwuth- filiform beak. 90.9 7 IR Ee PS N. antidesmatis GConidiawithoutibedk ss dis... a dete «a dinanecd diaeeg Db peed baer d Dhoede N. maestrense 4. Syritietmata Scattered s Peete dh sabes deh sukea dye pikes dr gpeteg dye peda Ae poder fe seb d inp ckm repeal 5 Syimematanouscatteted At. turk t, ataug ta Aue toh oe ee OY oe eRe one oan. te ae 6 Daw P@Onidia-distoseptate teehee shin gl igh adie dog ad oy N. diaoluoshanense COnidig"eusepiales ei. 5 -Seis xp Hels xa -dlelnge oleae lelage® lett Steiner 4s N. micheliae 6 Gonidrawith beakig . b..idine+ddike edd caegd naegd bard weedd bow dibapews Qarvuls 7 COMATARWVIENOUL DEA Hert eA forte) Atel A Sat lB ah lah ic Bat uleSat 5 Cae N. malloti Fite, PROT TAR CUSE IAL ES acter ote schire e chce lik a ee a mR N. vietnamense
omIIaPIStOSE PALE! Soups droog tr ponses drinsceg dr pee dh preg aye og ayn gedoe ct peta ea gctoe 8
Neosporidesmium diaoluoshanense sp. nov. (China) ... 309
A | B
20 um
Fic. 1. Neosporidesmium diaoluoshanense. A. Synnemata on natural substratum. B. Conidia with an apical, hyaline, globose, mucilaginous sheath. C, D. Synnemata with conidiophores and conidia.
&. ) sConidia with: 6-10 distoséptany. ams ocaniettiaptcly analy ata N. xanthophylli Coridia svat 1O=TLdistosepta soa ipa earnbryoaeetetoa peter te N. sinensis
Acknowledgments
The authors express gratitude to Dr. Eric H.C. McKenzie and Dr. R.F Castafieda Ruiz for serving as pre-submission reviewers and for their valuable comments and suggestions. This project was supported by the National Natural Science Foundation of
310... Li, Liu, & Zhang
China (Nos. 31093440, 31230001) and the Ministry of Science and Technology of the People’s Republic of China (Nos. 2006FY120100).
Literature cited
Ma J, Wang Y, Ma LG, Zhang YD, Castafeda-Ruiz RF, Zhang XG. 2011a. Three new species of Neosporidesmium from Hainan. Mycol. Progress 10: 157-162. http://dx.doi.org/10.1007/s11557-010-0685-2
Ma J, Wang Y, O’Neill NR, Zhang XG. 2011b. A revision of the genus Lomaantha, with the description of a new species. Mycologia 103(2): 407-410. http://dx.doi.org/10.1007/s11557-010-0696-z
Ma J, Zhang YD, Ma LG, Ren SC, Castafeda-Ruiz RF, Zhang XG. 2012. Three new species of Solicorynespora from Hainan, China. Mycol. Progress 11: 639-645. http://dx.doi.org/10.3852/ 10-176
Mercado Sierra A, Mena Portales J. 1988. Nuevos o raros hifomicetes de Cuba. Acta Bot. Cubana 59: 1-6.
Mel'nik VA, Braun U. 2013. Atractilina alinae sp. nov. and Neosporidesmium vietnamense sp. nov. - two new synnematous hyphomycetes from Vietnam. Mycobiota 3: 1-9. http://dx.doi.org/10.12664/mycobiota.2013.03.01
Wu WP, Zhuang WY. 2005. Sporidesmium, Endophragmiella and related genera from China. Fungal Diversity Research Series 15. 351 p.
Zhang K, Ma J, Wang Y, Zhang XG. 2009a. Three new species of Piricaudiopsis from southern China. Mycologia 101(3): 417-422. http://dx.doi.org/10.3852/08-147
Zhang K, Ma LG, Zhang XG. 2009b. New species and records of Shrungabeeja from southern China. Mycologia 101(4): 573-578. http://dx.doi.org/10.3852/09-006
Zhang YD, Ma J, Ma LG, Castafieda-Ruiz R.F, Zhang XG. 2011a. New species of Phaeodactylidium and Neosporidesmium from China. Sydowia 63: 125-130.
Zhang YD, Ma J, Wang Y, Ma LG, Castafeda-Ruiz R.E, Zhang XG. 2011b. New species and record of Pseudoacrodictys from southern China. Mycol. Progress 10: 261-265. http://dx.doi. org/10.1007/s11557-010-0696-z
ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MY COTAXON
http://dx.doi.org/10.5248/130.311 Volume 130, pp. 311-314 April-June 2015
A new species of Matsushimiella from submerged leaves in the Brazilian Amazon Forest
JOSIANE SANTANA MONTEIRO’, LuiS FERNANDO PASCHOLATI GUSMAO”™, & RAFAEL E CASTANEDA-RUIZ?
‘Departamento de Ciéncias Biologicas, Laboratorio de Micologia, Universidade Estadual de Feira de Santana, BR116 KM03, 44031-460, Feira de Santana, Brazil
?Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt’ (INIFAT), Académico Titular de la Academia de Ciencias de Cuba, Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
*CORRESPONDENCE TO: lgusmao@uefs. br
AxBsTRACT—Matsushimiella paraensis sp. nov. is described and illustrated from specimens collected on submerged leaves in a stream, from Para, Brazil, in the Amazon rainforest. The fungus is characterized by polyblastic sympodially extending conidiogenous cells and ellipsoid, long ovoid to broadly oblong, 2-4-distoseptate medium to golden brown smooth conidia with a pale brown to subhyaline basal frill left by rhexolytic conidial secession.
KEY worDs— taxonomy, freshwater fungi, tropics
Introduction
During a mycological survey of conidial fungi occurring on submerged decaying plant remnants from the Amazon rainforest in Brazil (Monteiro et al 2014a,b), an interesting fungus was collected on submerged leaves. Its conidiogenesis and conidial features clearly suggest a placement in the hitherto monotypic genus Matsushimiella R.F. Castafieda & Heredia (Castaneda-Ruiz et al. 2001). However, there are remarkable conidial differences distinguishing Matsushimiella queenslandica (Matsush.) R.E. Castafieda & Heredia, the type species of the genus, and the new collections from Brazil.
Material & methods
Samples of submerged litter were placed in paper and plastic bags, taken to the laboratory, and treated according to Castafieda-Ruiz (2005). Mounts were prepared in PVL (polyvinyl alcohol, lactic acid, and phenol) and measurements were taken at x1000.
312 ... Monteiro, Gusmao, & Castafieda-Ruiz
Micrographs were obtained with an Olympus microscope (model BX51) equipped with bright field and Nomarski interference optics. The holotype and an additional specimen are deposited in the Herbarium of Universidade Estadual de Feira de Santana, Brazil (HUEEFS).
Taxonomy
Matsushimiella paraensis J.S. Monteiro, R.F. Castafieda & Gusmao, sp. nov. Fia.1 MycoBank MB 807635
Differs from Matsushimiella queenslandica by its larger, ellipsoid, long ovoid to broad oblong, 2-4-distoseptate conidia.
Type: Brazil, Para State, Belém, Parque Estadual do Utinga, 1°25’S 48°27’W, on submerged leaves of an unidentified plant in a stream, 9 Jan. 2013, coll. J.S. Monteiro (Holotype: HUEFS 210425).
EryMo_oey: Latin paraensis refers to Para State, where the type specimen was collected.
CoLonies on the natural substrate effuse, hairy, smooth, golden brown. CONIDIOPHORES distinct, unbranched, erect, cylindrical below, geniculate above, 5-10-septate, smooth, wall somewhat thickened, up to 1.5 um wide, dark brown at the base, pale brown toward the apex, smooth, 170-220 x 5-8 um. CONIDIOGENOUS CELLS integrated, terminal, geniculate, with several sympodial extensions, polyblastic, pale brown, 40-70 x 4-5 um. SEPARATING CELLS more or less cylindrical, subhyaline, 3-4 x 2-2.5 um, fracturing and remaining as a persistent short peg. Conidial secession rhexolytic. Conip1a solitary, acropleurogenous, ellipsoid, long ovoid to broadly oblong, 2-4-distoseptate, mostly 4-distoseptate, 20-25 x 8-10 um, wall 1-1.5 um thick, smooth, mid brown or golden brown, fimbriate at the base, frill subhyaline to pale brown, 1-2 um long, 2-2.5 um wide.
ADDITIONAL SPECIMEN EXAMINED: BRAZIL, ParA, Parque Estadual do Utinga, 1°25’S
48°27’W, on submerged leaves of an unidentified plant in a stream, 17 Oct. 2012, coll.
J.S. Monteiro (HUEFS 210427). Note: Matsushimiella is typified by M. queenslandica (Castafieda-Ruiz et al. 2001) based on Pseudospiropes queenslandica Matsush., which was collected from decaying leaves of an unidentified plant in Australia. The existence of the separating cells was mentioned in the brief discussion by Matsushima (1989), but not described in the original description of Matsushimiella (Castafeda- Ruiz et al. 2001). Seifert et al. (2011) characterized the conidiogenous cells as polytretic, which refers to the original description of Matsushimiella (Castafieda-Ruiz et al. 2001). However, conidiogenesis is holoblastic, with mostly polyblastic conidiogenous cells. Matsushimiella queenslandica is readily distinguishable from the new Brazilian species by its obovate, 1-3-, mostly 2-distoseptate, 9-16 x 6-8 um, smooth, pale brown conidia that secede by rhexolytic fracture of cylindrical separating cells.
Matsushimiella paraensis sp. nov. (Brazil) ... 313
Fic. 1. Matsushimiella paraensis (holotype HUEFS 210425): A. Conidia. B. Conidiophore, conidiogenous cell, separating cells, and conidia. C. Conidiogenous cells. D-F. Conidiogenous cells, separating cells (arrow), and conidia. Scale bars = 10 um.
314 ... Monteiro, Gusmao, & Castafieda-Ruiz
Acknowledgments
The authors express their sincere gratitude to Uwe Braun and Eric H.C. McKenzie for their critical reviews of the manuscript. The authors are grateful to the Coordination for the Improvement of Higher Education Personnel (CAPES) for financial support and the ‘Programa de Pés-graduacao em Biologia de Fungos - PPGBF/UFPE: The authors thank the support provided by ‘Programa Ciéncia sem Fronteiras’ RFCR is grateful to Cuban Ministry of Agriculture and ‘Programa de Salud Animal y Vegetal, project P131LH003033 for facilities. We acknowledge the assistance provided by Paul M. Kirk and Vincent Robert and Arthur de Cock through the IndexFungorum and Mycobank websites. Lorelei Norvell’s editorial and Shaun Pennycook’s nomenclatural reviews are greatly appreciated.
Literature cited
Castaneda-Ruiz RF. 2005. Metodologia en el estudio de los hongos anamorfos. Anais do V Congresso Latino Americano de Micologia, Brasilia: 182-183.
Castafieda-Ruiz RF, Heredia G, Reyes M, Arias RM, Decock C. 2001. A revision of the genus Pseudospiropes and some new taxa. Cryptogamie Mycologie 22: 3-18. http://dx.doi.org/10.1016/S0181-1584(01)01057-0
Matsushima T. 1989. Matsushima mycological memoirs no. 6. Matsushima Fungus Collection, Kobe, Japan.
Monteiro JS, Gusmao LFP, Castafieda-Ruiz RF. 2014a. Helicodochium, a new microfungus from submerged wood in Brazil. Mycotaxon 127: 5-9. http://dx.doi.org/10.5248/127.5
Monteiro JS, Gusmao LFP, Castafeda-Ruiz RE. 2014b. Two new microfungi from Brazilian Amazon Forest: Atrogeniculata submersa and Nigrolentilocus amazonicus. Mycotaxon 127: 39-45. http://dx.doi.org/10.5248/127.39
Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS Biodiversity Series 9: 1-997. http://dx.doi.org/10.3767/003158511X617435
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MYCOTAXON
http://dx.doi.org/10.5248/130.315 Volume 130, pp. 315-320 April-June 2015
Amazonian phalloids: new records for Brazil and South America
TIARA S. CABRAL', CHARLES R. CLEMENT’, & IURI G. BASEIA3
‘Programa de Pés-graduacdo em Genética, Conservacdo e Biologia Evolutiva &
?Coordenacao de Tecnologia e Inovacao, Instituto Nacional de Pesquisas da Amazonia, Av. André Araujo, 2936 - Petropolis, Manaus, Amazonas, 69067-375, Brazil
*Departamento de Botanica e Zoologia, Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte, 59072-970, Brazil
* CORRESPONDENCE TO: ttiara@gmail.com
ABSTRACT — Recent field trips in the Amazonian rainforest revealed two uncommon phalloid species, Lysurus arachnoideus (new for Brazil) and Phallus cinnabarinus (new for South America). Detailed morphological descriptions, photos, and taxonomic remarks are presented.
KEY worps — gasteroid fungi, Lysuraceae, Neotropics, new records, Phallaceae
Introduction
Corda (1842) proposed the family Lysuraceae (as “Lysuroidea”) when he segregated the genera Lysurus Fr. and Aseroe Labill. from Clathraceae. Though many authors did not accept Lysuraceae (Cunningham 1944, Zeller 1949, Dring 1980), Hosaka et al. (2006) showed that Lysuraceae is phylogenetically more closely related to Phallaceae than to Clathraceae, and considered it an independent family, comprising the single genus Lysurus Fr., originally proposed to accommodate L. mokusin (Fries 1823).
Kirk et al. (2008) accept five species in Lysurus, although Index Fungorum lists 41 names for the genus. Diagnostic morphological features for Lysurus include a long stipe with a distinct fertile portion consisting of arms or vertical columns that are united or free or that form a network and a gleba lying on the inner surface of each arm or column (Dring 1980, Trierveiler-Pereira et al. 2014).
The genus Phallus Junius ex L. includes species recognized by their phallic shape that comprises an immature basidioma enclosed in a volva that breaks
316 ... Cabral, Clement, & Baseia
with maturity and from which emerges a pseudostipe with a fetid mucilaginous gleba. Phallus species are widely distributed, but Kreisel (1996) indicated that the center of diversity is China and Southeast Asia.
Thirteen Phallus records and five Lysurus species are currently recognized from Brazil (Trierveiler-Pereira & Baseia 2009, Cortez et al. 2011); some doubtful taxa have also been reported. This paper contributes to the taxonomic knowledge of phalloid diversity in Amazonia.
Materials & methods
The specimens were collected during field trips in Brazilian Amazonian rainforests (Careiro, state of Amazonas, Brazil) and French Guiana (Cayenne), where ecological notes and photographs were taken. Morphological measurements were taken from fresh and dried specimens according to Dring (1980) and Cortez et al. (2011). Color descriptions follow Kueppers (1982). Microstructures were measured after rehydration in KOH 5%, with 20 measurements made randomly for each structure using a Leica DM 2500 microscope. Spore and hyphal images were captured with a coupled EC3 camera and Leica Application suite v.2.1.0 software. The specimens were deposited at the Instituto Nacional de Pesquisas da Amazénia Herbarium, Manaus, Amazonas, Brazil (INPA).
Taxonomy
Lysurus arachnoideus (E. Fisch.) Trierv.-Per. & Hosaka, Mycologia 106: 909, 2014. FIG. 1
= Aseroe arachnoidea E. Fisch., Denkschr. Schweiz. Naturf. Ges. WS. Lee, Mycologia 32(1): 76, 1890.
Eces 15-20 mm diam., globose to subglobose, pale yellow (NOOY20MO00), semi-hypogeous, thin rhizomorphs present. EXPANDED BASIDIOMATA 40 x 11 mm. Volva 27-21 mm wide, white (A00M00C00) to pale yellow (NOOY20M00), outer layer composed of filamentous hyphae 4.6-6.4 um wide, septate with clamp connections, inner layer gelatinous, composed of 4-5.5 um wide filamentous hyphae. Pseudostipe 12 x 11 mm, cylindrical, white (AO0M00C00); surface rugose, hollow, ending in a central perforated disc; composed of globose to subglobose pseudoparenchymatous hyphae, 70 um diam., hyaline in 5% KOH. Receptacle formed by a disc with 9 arms developing from the margins. Arms up to 18 mm long, not cylindrical, hollow and consisting of a single tube, white (AO0M00C00), inner surface rugose, covered by the gleba up to 2/3 of the arm length. Gleba olive brown (N99A50M10), fetid. Bastp1osPpoREs 3.5-4.4 x 2.1-2.5 um, ellipsoid, smooth, greenish in 5% KOH.
ECOLOGY & DISTRIBUTION: Hosaka (2010) reports L. arachnoideus from Africa, Thailand, Japan, China, Indonesia, Sri Lanka, and Malaysia.
SPECIMENS EXAMINED — BRAZIL. AMAZONAsS: Careiro, Purupuru community, 3°22.96’S 59°42.63’W, in dung, in an ombrophilous dense forest fragment, 7.112013, TS.
Phalloids new for Brazil ... 317
20 mm eT
Figure 1. Lysurus arachnoideus (INPA 256537). A: Expanded basidioma. B: Basidiospores.
Cabral 41 (INPA 256537; Genbank KJ764820). JAPAN. TorTort: Kokoje, 1.[X.1980, T.
Hongo & E. Nagasawa (TMI 6927); 5.1X.1980, E. Nagasawa (TMI 6929); 9.1X.1980, E.
Nagasawa (TMI 6930); 22.1X.1983, I. Arita (TMI 11622). ComMMENTS — Lysurus arachnoideus sensu lato was previously recorded only once from South America, by Fischer (1928), who described a new variety (Aseroe arachnoidea var. americana) based on a specimen collected in Suriname, which apparently differed from A. arachnoidea var. arachnoidea mainly by the arms being arranged in pairs. Unfortunately Fischer (1928) did not provide a detailed description or include microstructure sizes. Our specimen has unpaired arms (Fic. 1A), and the spore size matches an African specimen described by Dring (1964). The taxonomically significant characters do not vary significantly between the Brazilian and the Japanese specimens. The Japanese material has a pseudostipe and arms longer than the Brazilian specimen, pseudoparenchymatous hyphae that are up to 76 um diam., and smooth spores that are up to 4 um long. Despite these differences, we prefer to maintain our collection as L. arachnoideus (instead of A. arachnoidea var. americana E. Fisch.) based on the morphological characters until further molecular analyses can be carried out.
Our Amazonas specimen represents the second record for South America
and the first for Brazil.
Phallus cinnabarinus (WS. Lee) Kreisel, Czech Mycol. 48: 278, 1996. Fic. 2 = Dictyophora cinnabarina WS. Lee, Mycologia 49: 156, 1957.
EGcs 24-23 mm wide, ovoid to subglobose, pale yellow (N10A10MO00) to brown (N60A90M00), with thick and lilac to pink rhizomorphs. EXPANDED BASIDIOMATA 192 mm high. Receptacle 34 high x 70 mm wide, bright orange (N10A70M70), campanulate, with an apical pore, surface reticulate.
318 ... Cabral, Clement, & Baseia
Figure 2. Phallus cinnabarinus (INPA 255835). A: Expanded basidioma. B: Basidiospores. C: Pseudoparenchymatous hyphae of pseudostipe with pinkish pigment droplets.
Pseudostipe 130 x 23 mm, cylindrical, spongy, pale pink (N10A40M30), composed of globose to subglobose pseudoparenchymatous hyphae, 40-67 um diam., hyaline in 5% KOH, pinkish pigment droplets present. Volva pale yellow (N10A10MO00), outer layer cottony, composed of filamentous hyphae 3-4 um diam., sinuous, septate, clamp connection present; inner layer gelatinous, composed of filamentous hyphae 2.2-3 um diam., hyaline in KOH 5%; basal portion with rhizomorphs. Indusium extending to half of pseudostipe, bright orange (N10A80M50), 69 mm long, polygonal meshes up to 9 mm, formed of globose to subglobose hyphae, 24-66 um diam., pinkish pigment droplets present. Gleba brown (N60A90M40), mucilaginous. BASIDIOSPORES 2.8-4.2 x 1.2-1.9 um, ellipsoid, smooth, olive brown in 5% KOH.
ECOLOGY & DISTRIBUTION — Kreisel (1996) first described Phallus cinnabarinus from Taiwan (Kreisel 1996), but it has since been reported from many parts of the world (Hemmes & Desjardin 2009).
SPECIMENS EXAMINED — BRAZIL, PaRA: Belém, Jardim Botanico Bosque Rodrigues Alves, 1°25.83’S 48°27.30’W, on soil among grasses and at the base of bamboo clumps, 27.X1.2013, T.S. Cabral 81 (INPA 255835; Genbank KJ764821); Museu Paraense Emilio
Phalloids new for Brazil ... 319
Goeldi, 1°27.55’S 48°28.60’W, 28.X1.2013, T.S. Cabral 82 (INPA 255836). FRENCH GUIANA: Cayenne, in grounds of the Herbier de Guyane (CAY), Institut de Recherche pour le Developpement (IRD), 4°56.35’N 52°17.21’W, 11.III.2013, Cabral TS 48 (INPA 255837). CoMMENTS — ‘There is controversy involving two morphologically similar Phallus species: P. multicolor (Berk. & Broome) Cooke and P. cinnabarinus. Phallus multicolor [= Dictyophora multicolor Berk. & Broome] was originally described as having a lemon-yellow indusium, an orange receptacle, and spores up to 5 um (Berkeley & Broome 1883), while P cinnabarinus was described with a cinnabar-red receptacle and indusium, volva with pinkish rhizomorphs at base, and spores up to 4 um (Lee 1957). Considering the original descriptions, we believe that the specimens analysed here represent P. cinnabarinus, based mainly on the cinnabar-red color of the indusium and receptacle, spores smaller than in P multicolor, and divergence in rhizomorph colours. Cunningham (1944) described an Australian specimen of Phallus multicolor, mentioning a salmon-pink indusium, orange receptacle, volva with short soft spines, and basidiospores up to 4.5 um long. Reporting P multicolor from India, Dutta et al. (2012) cited a lemon-yellow indusium, lemon-yellow to yellowish orange receptacle, and different basidiospore size. Although Hemmes & Desjardin (2009) provide good photos of both species that illustrate the primary macroscopic differences (indusium and receptacle colours) between the two species, unfortunately, a detailed description of P. multicolor cannot be found in the recent literature.
There is no previous record of P. cinnabarinus for South America. Cheype (2010) reported P. aff. multicolor from French Guiana, but he does not mention a herbarium voucher, which prevents further comparative analysis. A Phallus specimen collected on a recent field trip to French Guiana enabled a comparative study that confirmed the conspecificity of the French Guianan and Brazilian collections as P cinnabarinus.
Acknowledgments
This study was supported by the Brazilian funding agencies CNPq (473422/2012- 3) and FAPEAM (3137/2012). The authors thank Herbarium CAY, Piero Giuseppe Delprete, and Rémi Girault for support during the field trip in Cayenne. The authors also thank Eiji Nagasawa for cordially lending materials from the Herbarium TMI, and Clark Ovrebo and Martin Esqueda for reviewing the manuscript.
Literature cited
Berkeley MJ, Broome CE. 1883. List of fungi from Brisbane, Queensland; with descriptions of new species. Part II. Transactions of the Linnean Society of London, Botany, 2nd Series, 2: 53-73.
Cheype JL. 2010. Phallaceae et Clathrus récoltés en Guyane Francaise. Bulletin Mycologique et Botanique Dauphiné-Savoie 66: 51-66.
Corda ACJ. 1842. Icones fungorum hucusque cognitorum. 5: 1-92.
320 ... Cabral, Clement, & Baseia
Cortez VG, Baseia IG, da Silveira RMB. 2011. Two noteworthy Phallus from southern Brazil. Mycoscience 52: 436-438. http://dx.doi.org/10.1007/S10267-011-0124-5
Cunningham GH. 1944. The gasteromycetes of Australia and New Zealand. J. McIndoe. 236 p.
Dring DM. 1964. Gasteromycetes of west tropical Africa. Mycological Papers 98. 60 p.
Dring DM. 1980. Contributions towards a rational arrangement of the Clathraceae. Kew Bulletin 35: 1-96. http://dx.doi.org/10.2307/4117008
Dutta AK, Chakraborty N, Pradhan P, Acharya K. 2012. Phallales of West Bengal, India. II. Phallaceae: Phallus and Mutinus Arun. Researcher 4: 21-25.
Fischer E. 1928. Untersuchungen tiber Phalloideen aus Surinam. Vierteljahrsschrift der Naturforschenden Gesellschaft in Zitirich, 73(Beibl. 15): 1-39.
Fries EM. 1823. Systema Mycologicum. 2: 276-620.
Hemmes DE, Desjardin DE. 2009. Stinkhorns of the Hawaiian Islands. Fungi 2: 8-10.
Hosaka K, Bates ST, Beever RE, Castellano MA, Colgan III W, Dominguez LS, Nouhra ER, Gem J, Giachini AJ, Kenney SR, Simpson NB, Spatafora JW, Trappe JM. 2006. Molecular phylogenetics of the gomphoid-phalloid fungi with an establishment of the new subclass Phallomycetidae and two new orders. Mycologia 98: 949-959.
Hosaka K. 2010. Preliminary list of Phallales (Phallomycetidae, Basidiomycota) in Taiwan. Memoirs of the National Science Museum, Tokyo 46: 57-64.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2008. Ainsworth and Bisby’s Dictionary of the Fungi. 10th ed. Surrey: CABI/International Mycological Institute.
Kreisel H. 1996. A preliminary survey of the genus Phallus sensu lato. Czech Mycology 48: 273-281.
Kueppers H. 1982. Color atlas. Barrons Educational Series, NY.
Lee WS. 1957. Two new phalloids from Taiwan. Mycologia 49: 156-158. http://dx.doi.org/10.2307/3755742
Trierveiler-Pereira L, Baseia IG. 2009. A checklist of the Brazilian gasteroid fungi (Basidiomycota). Mycotaxon 108: 441-444. http://dx.doi.org/10.5248/108.441
Trierveiler-Pereira L, Silveira RMB da, Hosaka K. 2014. Multigene phylogeny of the Phallales (Phallomycetidae, Agaricomycetes) focusing on some previously unrepresented genera. Mycologia 106: 904-911. http://dx.doi.org/10.3852/13-188
Zeller SM. 1949. Keys to the orders, families and genera of the gasteromycetes. Mycologia 41(1): 36-58.
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MYCOTAXON
http://dx.doi.org/10.5248/130.321 Volume 130, pp. 321-328 April-June 2015
On the generic names Kriegeria
MERJE TOOME’, SHAUN R. PENNYCOOK? & M. CATHERINE AIME‘*
' Department of Botany and Plant Pathology, Purdue University, West Lafayette, IN 47907, USA ? Landcare Research, Private Bag 92170, Auckland 1072, New Zealand
*CORRESPONDENCE TO: maime@purdue.edu
AsBsTRACT — The basidiomycetous genus Kriegeria Bres. 1891 (Kriegeriaceae), typified by K. eriophori, is legitimate. An ascomycetous genus “Kriegeria” was originally published by Rabenhorst in 1876 as an invalid provisional name based on Ombrophila kriegeriana. Subsequently, the ascomycetous genus was validly published as Kriegeria Hohn. 1914 (Rutstroemiaceae), typified by Peziza elatina; this genus is an illegitimate later homonym, currently treated as a synonym of Rutstroemia. Additionally, Rabenhorst’s invalid name was “revived” (validated) as Kriegeria Rabenh. ex Seaver 1943, typified by O. kriegeriana, to accommodate species currently placed in Chloroscypha. Revised nomenclators are presented for both of the genera, as well as for Ombrophila kriegeriana.
Key worps — Helotiales, Kriegeriales, Leotiomycetes, Microbotryomycetes, Xenogloea, Zymoxenogloea
Introduction
Two different genera have been named as Kriegeria. The citation of one of these, the basidiomycetous genus Kriegeria published by Bresadola (1891), is uncontroversial. However, the correct citation of the ascomycetous genus Kriegeria has been the subject of long-standing debate, attributed either to Rabenhorst (1876, 1878a) or to Hohnel (1914); and consequently, the basidiomycetous Kriegeria Bres. has been considered to be either an illegitimate later homonym, or a legitimate earlier homonym. ‘The basis of this controversy is whether Kriegeria Rabenh. is a validly published alternative name, or an invalid provisional name. We have conducted an extensive literature survey and nomenclatural analysis to resolve this problem.
Kriegeria Hohn. (Rutstroemiaceae, Helotiales, Leotiomycetes)
Kriegeria Hoéhn., Sitzungsber. K. Akad. Wiss., Math.-Naturwiss. Kl, Abt. 1, 123: 114. 1914, nom. illeg., non Bres. 1891. TyPE: Peziza elatina Alb. & Schwein. : Fr., Consp. Fung. Lusat.: 330. 1805.
322 ... Toome, Pennycook, & Aime
= Rutstroemia P. Karst., Bidr. Kann. Finl. Nat. Folk 19: 12. 1871. Tye: Peziza firma Pers. : Fr., Syn. Meth. Fung.: 658. 1801. = Kriegeria Rabenh. ex Seaver, Mycologia 35: 492. 1943, nom. illeg., non Bres. 1891, nec Hohn. 1914. [“Kriegeria” Rabenh., Fungi Europ. Exsicc. 24: no. 2315. 1876, nom. prov.] Type: Ombrophila kriegeriana Rabenh., Fungi Europ. Exsicc. 24: no. 2315. 1876.
Ombrophila kriegeriana Rabenh., Fungi Europ. Exsicc. 24: no. 2315. 1876. [“Kriegeria olivacea” Rabenh., Fungi Europ. Exsicc. 24: no. 2315. 1876, nom. prov.]
= Ciboria kriegeriana (Rabenh.) Rehm, Hedwigia 22: 36. 1883.
= Chlorosplenium kriegerianum (Rabenh.) Sacc., Syll. Fung. 8: 318. 1889.
= Kriegeria kriegeriana (Rabenh.) Seaver, Mycologia 35: 493. 1943.
= Peziza elatina Alb. & Schwein. : Fr., Consp. Fung. Lusat.: 330. 1805.
= Helotium elatinum (Alb. & Schwein. : Fr.) Quél., Ench. Fung.: 309. 1886.
= Chlorosplenium elatinum (Alb. & Schwein. : Fr.) Sacc., Syll. Fung. 8: 318. 1889.
= Rutstroemia elatina (Alb. & Schwein. : Fr.) Rehm, Rabenh. Krypt.-Fl.,
Ed. 2, 1(3): 767. 1893. = Kriegeria elatina (Alb. & Schwein. : Fr.) Héhn., Sitzungsber. K. Akad. Wiss., Math.-Naturwiss. KI., Abt. 1, 123: 114. 1914.
The critical publication concerning the ascomycetous Kriegeria is the protologue of Ombrophila kriegeriana (Rabenhorst 1876). This printed exsiccata label presents a Latin description, Latin collection data [K6nigstein, Saxony, on Abies twigs], a paragraph of German “Obs.’, and the author's initials “L.R.” In the Observations, Rabenhorst noted: (1) that the specimens are immature, creating some doubt whether Ombrophila is the appropriate genus [indicated by inclusion of a question mark after the genus name in the binomial proposal]; (2) that the collector, W. Krieger, will endeavor to collect more material with mature ascomata; (3) that the species may require the proposal of a new genus for which he would suggest the name Kriegeria; (4) that in this potential new genus, he would suggest the epithet olivacea. He did not combine the genus and epithet into an explicit binomial (Seaver 1943 appears to be the first author to assemble the binomial). Considered as a whole, the Observations do not suggest that “Kriegeria olivacea” was being formally proposed as an alternative name for Ombrophila kriegeriana, but that it was merely being suggested as a provisional name to be taken up in the future if mature specimens justified the proposal of a new genus. “Kriegeria” Rabenh. and “Kriegeria olivacea” Rabenh. are therefore invalid names (McNeill et al. 2012: Art. 36.1(b)).
This situation is somewhat obscured in Rabenhorst (1878a), where the Ombrophila kriegeriana protologue is reproduced verbatim, but with the first two sentences of the Observations omitted; ie., there is no mention of the immaturity of the specimens nor of Krieger's intention to look for mature material. (Nevertheless, when Donk (1958) reached his firm decision that
Kriegeria Bres. vs. Kriegeria Hohn. & Kriegeria Rabenh. ... 323
“Kriegeria” Rabenh. was a provisional name, it was this reference that he cited and its abbreviated version of the original Observations that he quoted.) Because Rabenhorst (1878a) has been widely (but erroneously) accepted as the protologue of O. kriegeriana/“K. olivacea’, several mycologists have misunderstood “Kriegeria olivacea” to have been proposed as a valid alternative name. It is significant that the Index of Names for Hedwigia vol. 17 (Rabenhorst 1878b: p. V) does not list either “Kriegeria” or “Kriegeria olivacea”. Rabenhorst died on 24 April 1881, presumably before Krieger had collected or sent any mature specimens.
Winter (1881: 70-71) commented on Ombrophila kriegeriana, reporting that he had often seen it during spring in Zurich. On the basis of his own observations and Rabenhorst’s description, Winter concluded that O. kriegeriana was a synonym of Peziza elatina Alb. & Schwein.; he made no mention of the names “Kriegeria” and “Kriegeria olivacea”.
Ombrophila kriegeriana was recombined in Ciboria by Rehm (1883: 36-37) and in Chlorosplenium by Saccardo (1889: 318). Both authors mentioned Winter's proposed synonymy with Peziza elatina (without accepting it), but neither of them mentioned the names “Kriegeria” and “Kriegeria olivacea”.
Rehm (1893: 767) recombined Peziza elatina in Rutstroemia (the genus in which this species is currently accepted). Rehm’s R. elatina nomenclator included Ombrophila kriegeriana and its Ciboria and Chlorosplenium recombinations as synonyms, but it did not include “Kriegeria olivacea” (even though Rabenhorst’s suggestion of this genus and this epithet are annotated in Rehm’s synopsis of the literature).
Hohnel (1914: 112-114) was the first author to validate the ascomycetous genus Kriegeria, but his treatment is full of errors:
(1) Hohnel attributed the genus to “Winter (Hedwigia, 1878, 17. Bd., p. 32)
(2) he stated that Peziza elatina was transferred to Kriegeria by “Winter (Hedwigia, 1878, 17. Bd., p. 32)”; and
(3) he stated that the generic type was “Kriegeria elatina (A. u. S.) Winter”.
The cited Hedwigia reference has no connection with Winter and is, in fact, Rabenhorst (1878a), which contains no mention of Kriegeria elatina or its basionym Peziza elatina. Rabenhorst edited the first ten issues of Hedwigia vol. 17 (pp. 1-160, January-October 1878), and Winter's editorship did not commence until issue 11 (November 1878). (Likewise, Winter did not assume editorship of Fungi Europaei Exsiccati until late 1881, after Rabenhorst’s death.) We have scrutinized all of Winter’s publications from 1878 until his death in 1887 and can find no mention of either “Kriegeria” or “Kriegeria elatina’; his only relevant publication was Winter (1881) where he synonymized Ombrophila kriegeriana under Peziza elatina (as discussed above). Kriegeria
324 ... Toome, Pennycook, & Aime
Hohn. cannot be treated as a validation of “Kriegeria” Rabenh., because they have different (albeit conspecific) generic types, Peziza elatina [= K. elatina] versus “K. olivacea”. Consequently, Hohnel’s (1914) Kriegeria and K. elatina must be interpreted as a gen. nov. and a comb. nov. attributable to Héhnel alone (not to “Rabenh. ex Hohn. and definitely not to “G. Winter ex Hohn.”). Kriegeria Hohn. 1914 is an illegitimate later homonym of the basidiomycetous Kriegeria Bres. 1891 — a genus that Hohnel himself (1909: 1157-1159) had apparently accepted as legitimate. Hohnel (1914) included a second species, Kriegeria urceolus (Fuckel) Hohn. [= Rutstroemia urceolus (Fuckel) W.L. White (White 1941; Whetzel 1945)].
In his monographic treatment of Rutstroemia, White (1941) included the erroneous synonym “Kriegeria Wint. Hedw. 17: 32, 1878” (copied from Hoéhnel 1914) in his generic nomenclator. He accepted the full R. elatina nomenclator published by Rehm (1893), including O. kriegeriana and its recombinations, but he added the fictitious “Kriegeria elatina Wint. Hedwigia 17: 32, 1878” (also copied from Hohnel 1914).
Seaver (1943) proposed to “revive” the Rabenhorst genus “Kriegeria’, typified by Ombrophila kriegeriana, which he recombined as Kriegeria kriegeriana. Because of its different type, Seaver’s genus must be interpreted as an additional later homonym, Kriegeria Rabenh. ex Seaver, a heterotypic synonym of Kriegeria Hohn. Seaver’s paper has several puzzling aspects. Inexplicably, he cited two different “protologue” references: “Rab. Fungi Eur. 2315. 1878 [sic]” (i.e., Rabenhorst 1876) for Ombrophila kriegeriana and “Hedwigia 17: 32. 1878” (i.e., Rabenhorst 1878a) for the genus “Kriegeria” and for the species “Kriegeria olivacea”. Seaver’s failure to observe that all three names were actually published simultaneously by Rabenhorst (1876) suggests that he had not seen the O. kriegeriana exsiccata label. Significantly, Seaver (1943) did not mention Hohnel’s treatment of Kriegeria nor Winter’s and Rehm’s synonymization of O. kriegeriana with Rutstroemia elatina, and he explicitly rejected White's (1941) placement of Kriegeria in Rutstroemia. Seaver (1943) synonymized his genus Kriegeria with Chloroscypha Seaver, and cited the protologue description of Chloroscypha (Seaver 1931) as the validating description of Kriegeria. In addition to the type (Rabenhorst’s immature exsiccata material), Seaver (1943) recombined five other species that had all previously been included in Chloroscypha (Seaver 1931, 1938), and two further species were recombined in Kriegeria by Seaver (1951). Apart from Kriegeria kriegeriana (on a pinaceous host: Abies), all of Seaver’s Kriegeria species (on cupressaceous or taxodiaceous hosts: Calocedrus, Chamaecyparis, Cryptomeria, Juniperus, Thuja, Sequoia) are currently accepted in Chloroscypha (Petrini 1982). Seaver seems to have been unaware of the incongruence between Kriegeria sensu Hohnel/White [= Rutstroemia] and Kriegeria sensu Seaver [= Chloroscypha].
Kriegeria Bres. vs. Kriegeria Hohn. & Kriegeria Rabenh. ... 325
Kriegeria Bres. (Kriegeriaceae, Kriegeriales, Microbotryomycetes)
Kriegeria Bres., Rev. Mycol. (Toulouse). 13: 14. 1891. = Xenogloea Syd. & P. Syd., Ann. Mycol. 17: 44. 1919, nom. illeg. (superfluous). TyPE: Kriegeria eriophori Bres., Rev. Mycol. (Toulouse). 13: 14. 1891. = Zymoxenogloea D.J. McLaughlin & Doublés, Mycologia 84: 671. 1992. Type: Zymoxenogloea eriophori D.J. McLaughlin & Doublés, Mycologia 84: 671. 1992.
Contemporary material of the sole species was distributed as exsiccata specimens (Pazschke 1895; Kohlmeyer 1962). Sydow & Sydow (1919) thought that Kriegeria “Winter (cfr. Hedwigia 1878, p. 32)” was a valid name, making the basidiomycetous Kriegeria Bres. an illegitimate later homonym, for which they proposed Xenogloea as a replacement name. Their miscitation of Winter as the author and Rabenhorst (1878a) as the protologue suggests that they were relying on Hohnel’s (1914) treatment rather than carefully considering the original Rabenhorst (1876) protologue. With “Kriegeria” Rabenh. being an invalid provisional name, Kriegeria Bres. is legitimate and Xenogloea is superfluous and illegitimate. Doublés & McLaughlin (1992) treated Kriegeria Bres. as a legitimate generic name, and described a new genus Zymoxenogloea for the yeast-like anamorphic state. Toome et al. (2013) proposed Kriegeriaceae and Kriegeriales as a new family and order typified by Kriegeria Bres.
Discussion The acomycetous genus Kriegeria has been published three times (once invalidly and twice illegitimately):
(1) as “Kriegeria” Rabenh. 1876, an invalid provisional name that no mycologist during the next 38 years ever treated as a formal name: its invalid status has been argued by several eminent mycologists, e.g., by Dennis (1954, 1956) and, most persuasively, by Donk (1958), and it is not listed by Kirk et al. (2008) nor by the Index Nominum Genericorum website (ING 2015);
(2) as Kriegeria Hohn. 1914, a valid generic name based on a different type, but an illegitimate later homonym (of Kriegeria Bres.; ING 2015): it was not taken up by any subsequent mycologist but almost universally accepted as a synonym of Rutstroemia (e.g., Rehm 1893, White 1941, Whetzel 1945, Dennis 1956, Kirk et al. 2008) — major errors in Hohnel’s protologue have created citational confusion that still persists 100 years later;
(3) as Kriegeria Rabenh. ex Seaver 1943, an illegitimate later homonym based on Rabenhorst’s invalid name: Seaver’s genus was rejected by subsequent mycologists (e.g., Dennis 1954, 1956; Kobayashi 1965; Petrini 1982), who argued that the species included in Kriegeria by
326 ... Toome, Pennycook, & Aime
Seaver (1943, 1951) were not congeneric with the type (Ombrophila kriegeriana [= Rutstroemia elatina]) but instead belonged in Chloroscypha. Seaver's Kriegeria homonym is not mentioned by Kirk et al. (2008) nor by ING (2015).
There is no legitimately published ascomycetous genus Kriegeria, and this name is not in current use.
In contrast, the basidiomycetous Kriegeria Bres. 1891 is a legitimate monotypic genus that is predated only by the invalid “Kriegeria” Rabenh. 1876. Saccardo (1892: 497) tentatively identified the type as a Septogloeum, and Hoéhnel (1909: 1157-1159) recombined it in Platygloea, but subsequently Kriegeria Bres. has been accepted as a distinct genus, either under the superfluous replacement name Xenogloea (Sydow & Sydow 1919: 44; Weiss 1950: 295; Martin 1952: 88-89; Kao 1956; Bandoni 1957: 838) or as a legitimate Kriegeria Bres. (e.g., Davis 1922: 410; Donk 1958: 206; Oberwinkler & Bandoni 1982: 1744; Farr et al. 1989: 753; Doublés & McLaughlin 1992; Fell et al. 2001; Swann et al. 2001; Sampaio & Oberwinkler 2011; Toome et al. 2013). Kriegeria is accepted as legitimate, and Xenogloea as superfluous, by Kirk et al. (2008) and ING (2015).
Acknowledgments
This study was supported by Assembling the Fungal Tree of Life (AFTOL) project NSF DEB-0732968. The authors are grateful to Paul Kirk for numerous discussions and to Drs. David McLaughlin and Dennis Desjardin for acting as expert referees.
Literature cited
Bandoni RJ. 1957 [“1956”]. A preliminary survey of the genus Platygloea. Mycologia 48: 821-840. http://dx.doi.org/10.2307/3755709
Bresadola G. 1891. Sur un nouveau genre de Tuberculariée. Revue Mycologique 13: 14-15, t.113.
Davis JJ. 1922. Notes on parasitic fungi in Wisconsin — VII. Transactions of the Wisconsin Academy of Sciences, Arts, and Letters 20: 399-411.
Dennis RWG. 1954. Chloroscypha Seaver in Britain. Kew Bulletin 1954: 410. http://dx.doi.org/10.2307/4108803
Dennis RWG. 1956. A revision of the British Helotiaceae in the Herbarium of the Royal Botanic Gardens, Kew, with notes on related European species. Mycological Papers 62. 216 p.
Donk MA. 1958. The generic names proposed for Hymenomycetes—VIII. Auriculariaceae, Septobasidiaceae, Tremellaceae, Dacrymycetaceae. Taxon 7: 193-207. http://dx.doi.org/10.2307/1216401
Doublés JC, McLaughlin DJ. 1992. Basidial development, life history, and the anamorph of Kriegeria eriophori. Mycologia 84: 668-678. http://dx.doi.org/10.2307/3760376
Farr DF, Bills GF, Chamuris GP, Rossman AY. 1989. Fungi on plants and plant products in the United States. APS Press, St. Paul, MN. 1252 p.
Fell JW, Boekhout T, Fonseca A, Sampaio JP. 2001. Basidiomycetous yeasts. 3-35, in: DJ McLaughlin et al. (eds). The Mycota, Vol. VII, Part B. Springer-Verlag, Berlin, Heidelberg.
Kriegeria Bres. vs. Kriegeria Hohn. & Kriegeria Rabenh. ... 327
Hohnel F von. 1909. Fragmente zur Mykologie (VIII. Mitteilung, Nr. 354 bis 406). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abt. 1, 118: 1157-1245.
Hohnel F von. 1914. Fragmente zur Mykologie (XVI. Mitteilung, Nr. 813 bis 875). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abt. 1, 123: 49-155.
ING. 2015. Index Nominum Genericorum. http://botany.si.edu/ing/ [accessed 3 March 2015].
Kao CJ. 1956. The cytology of Xenogloea eriophori. Mycologia 48: 288-301. http://dx.doi.org/10.2307/3755477
Kirk PM, Cannon PF, Minter DW, Stalpers JA (eds). 2008. Dictionary of the Fungi, 10th edition. Wallingford, UK: CABI. 771 p.
Kobayashi T. 1965. Taxonomic notes on Chloroscyphae causing needle blight of Japanese conifers. Bulletin of the Government Forest Experiment Station, Tokyo, 176: 55-77.
Kohlmeyer J. 1962. Index alphabeticus Klotzschii et Rabenhorstii Herbarii Mycologici. Beihefte zur Nova Hedwigia 4. 231 p.
Martin GW. 1952. Revision of the north central Tremellales. University of lowa Studies in Natural History 19(3). 122 p. [Reprint - Bibliotheca Mycologica 22. 1969.]
McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Marhold K, Prado J, Prud’homme van Reine WF, Smith GF, Wiersema JH, Turland NJ. 2012. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code), adopted by the Eighteenth International Botanical Congress, Melbourne, Australia, July 2011. Regnum Vegetabile 154. 208 p. http://www.iapt-taxon.org/nomen/main.php
Oberwinkler FE, Bandoni RJ. 1982 A taxonomic survey of gasteroid, auricularioid Heterobasidiomycetes. Canadian Journal of Botany 60: 1726-1750. http://dx.doi.org/10.1139/b82-221
Pazschke FO. 1895. Kriegeria Eriophori Bres. Fungi Europaei et Extraeuropaei Exciccati, Editio nova, Series secunda, Century 41: no. 4084.
Petrini O. 1982. Notes on some species of Chloroscypha endophytic in Cupressaceae of Europe and North America. Sydowia 35: 206-222.
Rabenhorst L. 1876. Ombrophila ? Kriegeriana Rabenhorst n. sp. Fungi Europaei Exciccati, Editio nova, Series secunda, Century 24: no. 2315.
Rabenhorst L. 1878a. L. Rabenhorst, Fungi europaei exciccati. Cent. 24. Dresdae, 1877. Hedwigia 17(2): 31-32.
Rabenhorst L (ed.). 1878b. Namens-Verzeichniss. Hedwigia 17: HI-VII.
Rehm H. 1883. Dr. Rehm: Ascomyceten fasc. XIV. Hedwigia 22: 33-41.
Rehm H. 1893. Ascomyceten: Hysteriaceen und Discomyceten. CCLXXXIX. Rutstroemia Karst. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz, 2. Aufl., 1(3): 763-769.
Saccardo PA. 1889. Sylloge Discomycetum et Phymatosphaeriacearum. Sylloge Fungorum 8: 3-859.
Saccardo PA. 1892. Supplementum universale. Pars II. Discomyceteae - Hyphomyceteae. Sylloge Fungorum 10. 964 p.
Sampaio JP, Oberwinkler F. 2011. Kriegeria Bresadola (1891). 1477-1479, in: CP Kurtzman et al. (eds). The Yeasts, a Taxonomic Study, 5th Ed., Vol. 3. Elsevier, Amsterdam.
Seaver FJ. 1931. Photographs and descriptions of cup-fungi—XIV. A new genus. Mycologia 23: 247-251. http://dx.doi.org/10.2307/3753827
Seaver FJ. 1938. Photographs and descriptions of cup-fungi—XXIX. The genus Chloroscypha. Mycologia 30: 594-596. http://dx.doi.org/10.2307/3754353
328 ... Toome, Pennycook, & Aime
Seaver FJ. 1943. Photographs and descriptions of cup-fungi—XXXVII. The genus Kriegeria. Mycologia 35: 492-493. http://dx.doi.org/10.2307/3754603
Seaver FJ. 1951. The North American cup-fungi (inoperculates). New York. 428 p.
Swann EC, Frieders EM, McLaughlin DJ. 2001. Urediniomycetes. 37-56, in: DJ McLaughlin et al. (eds). The Mycota, Vol. VII, Part B. Springer-Verlag, Berlin, Heidelberg.
Sydow H, Sydow P. 1919. Mycologische Mitteilungen. Annales Mycologici 17: 33-47.
Toome M, Roberson RW, Aime MC. 2013. Meredithblackwellia eburnea gen. et sp. nov., Kriegeriaceae fam. nov. and Kriegeriales ord. nov.—toward resolving higher-level classification in Microbotryomycetes. Mycologia 105: 486-495. http://dx.doi.org/10.3852/12-251
Weiss FE. 1950. Index of plant diseases in the United States. Part II. Plant Disease Survey Special Publication 1(2): 219-381.
Whetzel HH. 1945. A synopsis of the genera and species of the Sclerotiniaceae, a family of stromatic inoperculate discomycetes. Mycologia 37: 648-714. http://dx.doi.org/10.2307/3755132
White WL. 1941. A monograph of the genus Rutstroemia (Discomycetes). Lloydia 4: 153-240.
Winter G. 1881. Notizen tiber einige Discomyceten. Hedwigia 20: 65-72.
ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MY COTAXON
http://dx.doi.org/10.5248/130.329 Volume 130, pp. 329-336 April-June 2015
New records of crustose lichens and a lichenicolous Arthonia from Vietnam
SANTOSH JOSHI’, DALIP KUMAR UPRETI’, SOON-OK OH’, Tur THuy NGUYEN?, ANH DZUNG NGUYEN 3, & JAE-SEOUN HurR’*
' Lichenology Laboratory, CSIR-National Botanical Research Institute,
Rana Pratap Marg, Lucknow (UP)-226001, India ? Korean Lichen Research Institute, Sunchon National University, Suncheon-540 950, Korea > Institute of Biotechnology & Environment, Tay Nguyen University,
567 Le Duan, Buon Ma Thuot City, Lak Province, Vietnam
*CORRESPONDENCE TO: jshur1@sunchon.ac.kr
ABSTRACT — New records from Vietnam of the crustose lichen species Arthonia excipienda, Chiodecton leptosporum, Graphidastra multiformis, Pertusaria pycnothelia, P. thwaitesii, and Phlyctis uncinata are presented together with the second world report of the non-lichenized lichenicolous fungus, Arthonia diorygmae. All species were collected from the Bidoup Nui Ba National Park located in the central highlands in Vietnam. A taxonomic description of each species is accompanied by distributional and ecological data and illustrations.
KEY worps — corticolous, Da Lat city, evergreen forest, tropical
Introduction
A lichenological expedition was organized in 2014 by the National Research Foundation of Korea and the Korea National Research Resource Center Program. During fieldwork two of the authors collected a large number of crustose lichens growing predominantly on trees in evergreen forests of the Bidoup Nui Ba National Park in Vietnam. Among the more interesting collections, we report and describe six corticolous species new for the country. New Vietnamese records include Arthonia excipienda, first representatives of the genera Chiodecton (C. leptosporum), Graphidastra (G. multiformis), and Phlyctis (P. uncinata), and two new Pertusaria species (P pycnothelia, P. thwaitesii) to be added the previously reported P. asiana Vain. and P. pertusa (L.) Tuck. (Aptroot & Sparrius 2006, Joshi et al. 2014).
The lichenicolous non-lichenized Arthonia diorygmae was found parasitizing a thallus of Diorygma, where it restricted the growth of the
330 ... Joshi & al.
host ascocarps. Although Arthonia accolens Stirt., A. antillarum (Fée) Nyl., A. cinnabarina (DC.) Wallr., A. cyanea Mill. Arg., and A. microcephala Vézda have been previously recorded from Vietnam as corticolous and foliicolous species (Aptroot & Sparrius 2006, Nguyen et al. 2011), no lichen-inhabiting fungus has ever been reported from Vietnam in Arthonia or any other genus.
Materials & methods
Morphological features of specimens collected from tropical rain forests of Da Lat city in Lam Dong province of Vietnam were examined under Magnus Zoom Stereo Trinocular (MSZ-TR) dissecting microscope and anatomical characters were observed under a compound microscope (Leica DM 500). Chemical spot tests and TLC (using solvent system A) were conducted according to Orange et al. (2010). The studied material is deposited in the herbarium of KoLRI (Korean Lichen Research Institute), South Korea.
Taxonomy
Arthonia diorygmae S. Joshi & Upreti, Lichenologist 45: 323, 2013. PL.1A
Thallus absent, non-lichenized, lichenicolous on Diorygma; ascomata dot- like, round, irregular to shortly lirellate, prominent; 0.2-0.3 x 0.1-0.2 mm; proper exciple completely carbonized to dark brown, 15-20 um wide, in continuation with hypothecium, undulating, V-shaped; epihymenium brown, 15-25 um high; hymenium hyaline, inspersed with oil droplets, gelatinous, <100 um high, I+ wine red; paraphyses indistinct, 1-1.5 um thick; asci broadly clavate, 8-spored, 40-50 x 13-15 um, epiplasm I+ wine red; ascospores hyaline becoming dark brown and warty, 2-celled, ovoid to slipper-shaped, 10-16 x 4—5 um, I-.
CHEMISTRY: Ascomata (Arthonia): K-, PD-, C-. Thallus (Diorygma host species): K+ yellow, PD+ yellow-orange, C-; norstictic and stictic acid chemosyndrome detected in TLC of host species.
DISTRIBUTION & ECOLOGY: This is the second report of A. dorygma, previously known only from the type locality in India (Joshi et al. 2013b). In Vietnam it was collected at ca. 1400 m from a thick- and smooth-barked tree, where it was growing luxuriantly and spread largely in association with Cryptothecia, Diorygma, and other graphidaceous taxa.
SPECIMENS EXAMINED: VIETNAM. LAM DONG PROVINCE: Bidoup Nui Ba National
Park, Da Lat city, 12°10°55.4”N 108°40’50.1”E, alt. 1454 m, on tree bark, 7 January 2014,
Hur & Oh VN140150, VN140098 (KoLRI). REMARKS: This species is distinctive in having non-lichenized lichenicolous life form. Though, our specimen differs from the A. diorygmae type material in also producing stictic acid by the host, all taxonomic characters were similar to those of the holotype described from the Western Ghats in India (Joshi et al. 2013b).
New crustose lichen records (Vietnam) ... 331
Arthonia excipienda (Nyl.) Nyl., Lich. Scand.: 261, 1861. PL. 1B
Thallus corticolous, crustose, endoperidermal, smooth, continuous, shiny, pale green to yellowish, thin, 40-60 tm in cross section, corticate; cortex indistinct to 30 um; photobiont cells chlorococcoid, aggregate to disperse, layer discontinuous, <25 um; medulla indistinct; ascomata lirelliform, sessile; lirellae 0.5-1 x 0.1-0.2 mm; disc slightly open, black, epruinose; thalline margin absent to indistinct; proper exciple laterally carbonized, 25-30 um wide, convergent, internally lined by 20-60 um periphysoids; epihymenium brown, 10-15 um high; hymenium hyaline, clear, gelatinous, 60-70 um high, I+ blue; hypothecium hyaline to brownish 20-25 um high; asci broadly clavate, 8-spored, 60-65 x 20-22 um; ascospores hyaline to brown, 19-21 x 7-8 um, I-.
CHEMISTRY: K-, PD-, C-; no lichen compounds detected in TLC.
DISTRIBUTION & ECOLOGY: The species was recorded from West European countries and South Korea (Joshi et al. 2013a); in Vietnam, it was found growing poorly at 1400-1500 m in association with other crustose corticolous lichens on smooth-barked trees.
SPECIMENS EXAMINED: VIETNAM. Lam DONG PROVINCE: Bidoup Nui Ba National Park, Da Lat city, 12°10’55.4”N 108°40’50.1”E, alt. 1454 m, on tree trunk, 7 January 2014, Hur & Oh VN140080 (KoLRI); 12°10’54.6”N 108°40’45.1’E, alt. 1426 m, on tree trunk, 8 January 2014, Hur & Oh VN140167 (KoLRI). REMARKS: Arthonia excipienda, an uncommon species in Asia, apparently resembles A. punctiformis Ach. and A. radiata (Pers.) Ach., which differ in having multiseptate ascospores. Arthonia didyma Korb. also produces 2-celled ascospores, but they are brown and warty on maturity.
Chiodecton leptosporum Miill. Arg., Flora 65: 332, 1882. PL. 1c
Thallus corticolous, crustose, epiperidermal, tightly attached to the substrate, smooth to slightly verruculose, pale green, with a white pruina, 200-400 um thick in cross section, ecorticate; photobiont cells Trentepohlia-like, layer <200 um; medulla whitish with few calcium oxalate crystals, 200-250 um, mostly endoperidermal; prothallus usually distinct, brownish; ascomata perithecioid, dot-like to round, solitary or rarely united, aggregated into distinctly elevated stroma-like structures usually with more than 10 ascocarps, 0.1-0.2 mm in diameter; stroma round to oval, 0.5-2 mm in diameter; proper exciple dark brown to completely carbonized, 25-45 um; epihymenium brown, indistinct to 15 um high; hymenium hyaline, clear, 135-140 um high, I+ blue; hypothecium extending down to the substrate, hard and black, <200 um high; paraphyses 1-1.5 um thick; asci clavate, 8-spored, 80-100 x 9-12 um, I-; ascospores obovate, hyaline, 3-septate, 30-40 x 3-4 um, I-; locules cylindrical, 7-9 x 1-2 um.
CHEMISTRY: K-, PD-, C-; roccellic acid detected in TLC.
332 ... Joshi & al.
DISTRIBUTION & ECOLOGY: Chiodecton leptosporum is widely distributed in Asia, Australia, and scattered localities in New Caledonia, Fiji, and Guam (Thor 1990); in Vietnam, it was luxuriantly growing at 1400-1500 m on hard- barked trees in an evergreen forest. The other lichens growing in association were unknown leprose taxa and Cryptothecia.
SPECIMENS EXAMINED: VIETNAM. Lam DONG PROVINCE: Bidoup Nui Ba National Park, Da Lat city, 12°10’55.4”N 108°40’50.1”E, alt. 1454 m, on tree bark, 7 January 2014, Hur & Oh VN140043, VN140065, VN140067, VN140078, VN140090 (KoLRI); 12°10’38.9"N 108°40’37.9’E, alt. 1426 m, on tree bark, 8 January 2014, Hur & Oh VN140149 (KoLRI). REMARKS: Chiodecton leptosporum can easily be confused with another most commonly distributed C. congestulum Nyl., which differs in a thallus containing a yellow pigmented medulla and a hymenium with a feeble iodine reaction. A comparatively rougher thallus and mostly united ascocarps further distinguish C. congestulum from C. leptosporum.
Graphidastra multiformis (Mont. & Bosch) G. Thor, Opera Bot. 103: 82, 1991. Pu, Lp
Thallus corticolous, crustose, epiperidermal, tightly attached to the substrate, cretaceous, pale green to greenish grey, 200-300 um thick in cross section, epinecral layer resembling cortex is present, 80-85 um; photobiont cells Trentepohlia-like, layer <120 um; medulla whitish, with few to numerous calcium oxalate crystals, 100-120 um; prothallus whitish in inner part and brownish in outer part; ascomata usually lirelliform, but sometimes apothecioid, solitary; apothecioid and lirelliform structures distinctly elevated with a slightly to distinctly constricted base, 1-2 x 0.5-1 mm; disc black, 5-10 mm thick; thalline margin 130-330 um thick; proper exciple thinly carbonized laterally, 15-20 um, basally continuous with hypothecium; epihymenium brown, 25-35 um; hymenium hyaline, clear, 60-85 um, I+ reddish; hypothecium extending down to the substrate, hard, black, + V-shaped, 0.3-0.5 mm; paraphyses 1-2 um thick; asci clavate, 55-60 x 5-6 um, I+ blue; ascospores hyaline, spermatoid, 3-septate, 35-40 x 2-3 um (including tail of 20-22 um), I-.
CHEMISTRY: K-, PD-(?), C-; roccellic acid and traces of protocetraric acid detected in TLC.
DISTRIBUTION & ECOLOGY: Graphidastra multiformis has previously been reported from India, Sri Lanka, The Philippines, Australia, West Samoa, and Tahiti (Thor 1990); in Vietnam it was collected at ca. 1400 m from thick- and hard-barked trees, where it was growing luxuriantly along with members of Pannariaceae and Chiodecton leptosporum.
SPECIMEN EXAMINED: VIETNAM. LAM DONG PROVINCE: Bidoup Nui Ba National Park, Da Lat city, 12°10°55.4”N 108°40’50.1”E, alt. 1454 m, on tree bark, 7 January 2014, Hur & Oh VN140075 (KoLRI).
New crustose lichen records (Vietnam) ... 333
REMARKS: Graphidastra multiformis is close to G. byssiseda (Mill. Arg.) G. Thor in having spermatoid ascospores and a thallus containing roccellic acid, but G. byssiseda differs in producing bi-clavate larger ascospores and lacking protocetraric acid.
Pertusaria pycnothelia Nyl., Bull. Soc. Linn. Norm., ser. 2, 2: 70, 1868. PL. 1E
Thallus corticolous, crustose, greenish grey to green or pale green, shiny, + rimose, verrucose, <500 um thick in cross section, corticate; cortex 20-25 um; photobiont Trebouxia, layer 85-90 um; medulla 180-320 um; prothallus blackish; ascomata verruciform, numerous, scattered, sometimes confluent and forming an almost continuous crust, concolorous with the thallus, flattened, hemispherical, 0.5-0.8 mm in diam; ostiole conspicuous, brown, dark brown in apothecioid to blackish in verruciform, 0.3-0.5 mm in diam; thalline margin 300-370 um wide; proper exciple hyaline to yellowish, 25-50 um wide; epihymenium pale brown, 45-47 um high; hymenium hyaline, clear, 300-400 um high; subhymenium hyaline, 80-120 um high; paraphyses 1-2 um thick; asci broadly clavate, 2-spored; ascospores hyaline, ellipsoid to subfusiform, double walled, wall smooth, 50-75 x 17-25 um, I+ blue.
CHEMISTRY: K-, KC-, C-, PD-; 4,5-dichlorolichexanthone and 2’-O-methylperlatolic acid detected in TLC.
DISTRIBUTION & ECOLOGY: Pertusaria pycnothelia was previously reported from Australia, New Caledonia, and Papua New Guinea (Archer 1997); in Vietnam, it was collected at 1400-1500 m, where it was growing in small patches over tree bark in an evergreen forest.
SPECIMEN EXAMINED: VIETNAM. LAM DONG PROVINCE: Bidoup Nui Ba National Park, Da Lat city, 12°10°55.4”N 108°40’50.1”E, alt. 1454 m, on tree bark, 7 January 2014, Hur & Oh VN140096 (KoLRI). REMARKS: The chemically similar Pertusaria trachyspora A.W. Archer is distinguished from P. pycnothelia by ascospores with a rough inner wall.
Pertusaria thwaitesii Mill. Arg., Flora 67: 470, 1884. PL. 1F
Thallus corticolous, crustose, epiperidermal, wrinkled, cracked, uneven, shiny, off-white to pale grey, 100-150 um thick in cross section, corticate; cortex 10-15 um; photobiont Trebouxia, layer 50-80 um; medulla white, 70-90 um; prothallus jet black; ascomata verruciform, conspicuous, concolorous with the thallus, numerous, very flattened hemispherical or irregular in outline, 1-2 mm wide; ostioles numerous, conspicuous, pale yellow becoming black, punctiform, becoming somewhat sunken, 0.05-0.07 mm in diam., 4-7 per verruca; proper exciple hyaline to yellowish, indistinct to 30 um wide; epihymenium indistinct; hymenium hyaline, clear, <430 um high; subhymenium indistinct; asci broadly
334 ... Joshi & al.
clavate, 2-spored, 290-300 x 45-50 um; ascospores elongate ellipsoid, double walled, inner wall ornamented and outer wall smooth, 140-165 x 35-45 um. CHEMISTRY: K-, PD+ orange-red, C-; protocetraric acid detected in TLC. DISTRIBUTION & ECOLOGY: Known from Australia, Papua New Guinea, Sri Lanka (Archer 1997); in Vietnam, Pertusaria thwaitesii was growing in association with Graphis at 1700-1800 m in trees in an evergreen forest. SPECIMEN EXAMINED: VIETNAM. LAM DONG PROVINCE: Mt Langbian, Da Lat city, 12°02’18.6’N 108°25’35.1”E, alt. 1779 m, on tree bark, 9 January 2014, Hur & Oh VN140281 (KoLRI). REMARKS: Pertusaria hartmannii Mull. Arg., which superficially resembles P. thwaitesii, differs in producing smoother ascospores and a thallus containing norstictic acid (Archer 1997).
Phlyctis uncinata Stirt., J. Linn. Soc., Bot. 14: 464, 1875. PL. 1G
Thallus corticolous, crustose, rough, epiperidermal, subleprose, + rimose due to substrate, greenish grey, whitish green to greyish green, 100-130 um thick in cross section, ecorticate; photobiont cells green protococcoid, layer 50-70 um; medulla white, 40-50 um; prothallus white; ascomata round to irregular, solitary to aggregate, immersed, 0.4-0.5 mm; disc blackish, finely pruinose, 0.2-0.3 mm in diameter; thalline margin recurved, 165-180 um wide; proper exciple hyaline to brownish, apically dark, 35-55 um wide; epihymenium granular, brownish 12-15 um high; hymenium hyaline, clear, 65-80 um high, I+ wine red; subhymenium <40 um high; asci clavate, 8-spored, 110-165 x 15-25 um, I+ wine red; ascospores hyaline, fusiform, crescent shape, transversely septate, 50-54 x 5-7 um, 8-locular, I+ wine red.
CHEMISTRY: K+ yellow turning red, PD+ yellow-orange, C-; norstictic acid detected in TLC.
DISTRIBUTION & ECOLOGY: Known from New Zealand and Thailand (Galloway 1985, 2007); in Vietnam, the species was collected at 1700-1800 m, where it was spread in short patches on rough-barked trees.
SPECIMEN EXAMINED: VIETNAM. LAM DONG PROVINCE: Mt Langbian, Da Lat city,
12°02’18.6”N 108°25’35.1”E, alt. 1779 m, on tree bark, 9 January 2014, Hur & Oh
VN140109 (KoLRI). REMARKS: Phlyctis uncinata is closely related to P. karnatakana S. Joshi & Upreti, which differs in having small (20-30 um long) ascospores (Joshi et al. 2010). The thallus of Phlyctis himalayensis (Nyl.) D.D. Awasthi is morphologically similar but reacts K+ red (Awasthi 1991). Phlyctis, which has poorly delimited species, needs further revision to clarify the status of P uncinata (Joshi & Upreti 2012; Joshi et al. 2010, 2012).
New crustose lichen records (Vietnam) ... 335
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Prate. 1. New records of lichens from Vietnam. A. Arthonia diorygmae (white arrow indicating ascomata of Arthonia and black arrow indicating suppressed growth of ascomata in host); B. Arthonia excipienda (ascomata); C. Chiodecton leptosporum; D. Graphidastra multiformis; E. Pertusaria pycnothelia; F. Pertusaria thwaitesii; G. Phlyctis uncinata. Scale bars: A, F, G = 0.5 mm; B, E=1mm;C,D=2 mm.
Acknowledgments
This work was supported by a grant from the National Research Foundation of Korea (#2011-0031494) and the Korea National Research Resource Center Program. Authors are grateful to Drs. S.Y. Kondratyuk and L. Lék6s for their valuable comments on the manuscript. DKU and SJ thank Director, CSIR-National Botanical Research Institute, Lucknow, India, for providing infrastructure to facilitate the identification of Korean lichen material.
336 ... Joshi & al.
Literature cited
Aptroot A, Sparrius LB. 2006. Additions to the lichen flora of Vietnam, with an annotated checklist and bibliography. Bryologist 109(3): 358-371. http://dx.doi.org/10.1639/0007-2745(2006) 109[358:ATTLFO]2.0.CO;2
Archer AW. 1997. The lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 69. 249 p.
Awasthi DD. 1991. A key to the microlichens of India, Nepal and Sri Lanka. Bibliotheca Lichenologica 40. 337 p.
Galloway DJ. 1985. Phlyctella Krempelh. 385-389, in: Flora of New Zealand Lichens. Government Printer, Wellington, New Zealand.
Galloway DJ. 2007. Phlyctis (Wallr.) Flot. 1184-1191, in: Flora of New Zealand Lichens, 2nd ed., Vol. 2. Manaaki Whenua Press, Lincoln, New Zealand.
Joshi S, Upreti DK. 2012. Lichen genus Phlyctis (Phlyctidaceae) in India. Geophytology 44(3): 363-369.
Joshi S$, Upreti DK, Mishra GK, Divakar PK. 2010. Two new species of lichen genus Phlyctis in India. Bryologist 113(4): 724-727. http://dx.doi.org/10.1639/0007-2745-113.4.724
Joshi S, Upreti DK, Nayaka S. 2012. Two new species in the lichen genus Phlyctis (Phlyctidaceae) from India. Lichenologist 44(3): 363-369. http://dx.doi-org/10.1017/S0024282911000879
Joshi S, Kondratyuk SY, Crisan F, Jayalal U, Oh SO, Hur JS. 2013a. New additions to lichen mycota of the Republic of Korea. Mycobiology 41(4): 177-182. http://dx.doi.org/10.5941/MYCO.2013.41.4.177
Joshi S, Upreti DK, Nayaka S. 2013b. A new lichenicolous Arthonia species (Arthoniaceae) on Diorygma from India. Lichenologist 45(3): 323-327. http://dx.doi.org/10.1017/S0024282913000042
Joshi S, Jayalal U, Oh SO, Nguyen TT, Dzung NA, Hur JS. 2014. A new species of Graphis and new lichen records from Vietnam, including a second worldwide report of Sarcographina cyclospora. Mycobiology 42(1): 17-21. http://dx.doi.org/10.5941/MYCO.2014.42.1.17
Nguyen TTT, Joshi Y, Licking R, Nguyen AD, Wang XY, Koh YJ, Hur JS. 2011. Seven new records of foliicolous lichens from Vietnam. Mycotaxon 117: 93-99. http://dx.doi.org/10.5248/117.93
Orange A, James PW, White FJ. 2010. Microchemical methods for the identification of lichens. British Lichen Society, London.
Thor G. 1990. The lichen genus Chiodecton and five allied genera. Opera Botanica 103. 92 p.
ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MYCOTAXON
http://dx.doi.org/10.5248/130.337 Volume 130, pp. 337-359 April-June 2015
First records of some Asian macromycetes in Africa
PaBLo P. DANIELS’, OUMAROU HAMA?’, ALFREDO JUSTO FERNANDEZ?, FELIX INFANTE GARCIA-PANTALEON’, MoussA BARAGE?’, DAHIRATOU IBRAHIM‘, & MARIA ROSAS ALCANTARA!
"Department of Botany, Ecology and Plant Physiology, University of Cordoba, Ed. Celestino Mutis, Campus Rabanales, Cordoba 14071 Spain
? Department of Plant Production, Faculty of Agronomy, University Abdou Moumouni, Niamey BP-10960 Niger
* Biology Department, Lasry Biological Science Center, Clark University, 950 Main St., Worcester, MA 01610 USA
* Life Sciences and Earth Department, High School of Education, University Abdou Moumouni, Niamey BP-10963 Niger
* CORRESPONDENCE TO: ppdaniels@hotmail.com
ABSTRACT — This paper reports and discusses preliminary data on new Asian macromycete species now recorded on the African continent and collected for the first time in Niger during sampling conducted in the southwestern region from 2008 to 2012. Descriptions and comments on chorology, systematics, and closely related species are given for Hymenagaricus subepipastus, Clitopilus orientalis, Tulostoma evanescens, Termitomyces bulborhizus, and Volvariella cf. sathei.
Key worps — basidiomycetes, fungi, taxonomy
Introduction
The literature available on the macromycetes of West Africa is generally both sparse and highly fragmented. Boa (2004) reported that there appeared to be no data at all for Niger, but since then a handful of references in local publications and international journals have cited a total of 16 fully identified species for this country (TABLE 1). Countries bordering this sub-Saharan region (with the exception of Mali and Chad) have recently been studied by mycologists who are beginning to publish research on the diversity, systematics, ecology, ethnomycology and use of macromycetes. As a result, much more information is now available on fungal species in Benin (De Kesel et al. 2000, 2002; De Kesel & Yorou, 2000; Yorou et al. 2002a,b; Yorou & De Kesel 2002; Yorou, 2010), Burkina Faso (Ganaba et al. 2002, Guissou et al. 2002, 2008, Guissou 2005),
338 ... Daniéls & al.
TABLE 1. Macromycete species previously recorded from Niger.
TAXON REFERENCE
Agaricus augustus Fr. Hama et al. 2008
Agaricus bulbillosus Heinem. & Gooss.-Font. Hama et al. 2008
Agaricus subsaharianus L.A. Parra et al. Hama et al. 2010, 2012 Crinipellis glaucospora (Beeli) Pegler Antonin 2013a
Crinipellis pseudosplachnoides (Henn.) Singer Antonin 2013a
Ganoderma colossus (Fr.) C.F. Baker Hama et al. 2008, 2009, 2012 Ganoderma lucidum (Curtis) P. Karst. Hama et al. 2008, 2009 Itajahya rosea (Delile) E. Fisch. Hama et al. 2008 Leucocoprinus brebissonii (Godey) Locq. Hama et al. 2008 Lyophyllum aggregatum (Schaeff.) Kihner Hama et al. 2008
Marasmius atrorubens (Berk.) Mont. Antonin 2013b
Phellinus allardii (Bres.) S. Ahmad Hama et al. 2012
Podaxis pistillaris (L.) Fr. Hama et al. 2008, 2009, 2012 Termitomyces striatus (Beeli) R. Heim Hama et al. 2008, 2009 Trametes cingulata Berk. Hama et al. 2008
Trametes leonina (Klotzsch) Imazeki Hama et al. 2008
Nigeria (Zoberi 1973; Oso 1975, 1977; Adewusi et al. 1993; Osagualekhor & Okhuoya 2005; Akpaja et al. 2003; Osemwegie et al. 2006; Okhuoya et al. 2010) and, to a lesser extent, Togo (De Kesel et al. 2008, Gardt et al. 2011) and Ivory Coast (Koné et al. 2010a,b). In general, many West African ecosystems are very sparsely explored. However, Yorou et al (2014) recently updated a total list of about 72 edible macromycetes for West Africa (Benin, Burkina Faso, Ivory Coast, and Togo). In West Africa, macromycetes are not only taxonomically poorly documented, but patterns of distributions, ecology, and local use should be addressed especially in the context where forest and savannah ecosystems are disappearing at an alarming rate (FAO 2010) along with the fungal species. As an example, a preliminary Red List of threatened fungi has been published for Benin (Yorou & De Kesel 2011).
Given the lack of information regarding Niger, the University of Cordoba submitted to the Spanish Agency for International Development Cooperation a project to be implemented jointly with the Abdou Moumouni University, Niamey, entitled “Edible and cultivable Macromycetes of Niger (Ethnomycology)”. One of the aims of this project was to draw up a classified inventory of Niger's fungal biodiversity. At an earlier stage, the local project team had already compiled a preliminary inventory, which was expanded and completed as part of the current research and served as a basis for the subsequent ethnomycological survey. A number of collections have been published recently (Hama et al. 2012, Antonin 2013a,b). The findings of the present study relate to the fungal diversity portion of the project and focus on the key taxonomic features of species reported earlier in Asia but which are considered first records in Africa.
New Asian macromycete records for Africa ... 339
Materials & methods
Material collected between 2008 and 2012 came mainly from the West Sudanian savannah (White 1986), deemed the most suitable area for a study of national macromycete biodiversity in view of its abundant vegetation; the study centered primarily on the deep-soil, wooded savannah and gallery forests bordering rivers of the W National Park in southwestern Niger. Mushrooms were mostly collected using routine sampling methods (Halling 1996, Eyi-Ndong et al. 2011); each species was assigned a collection number and photographed with a Canon 400D, Canon PowerShot G10, or Olympus U 700 digital camera. Fresh mushroom organoleptic data - colour, odour, and taste - and information on gross features likely to be modified during drying and required for identification purposes were entered on field records. Collected mushrooms were then dehydrated in situ using a Bunsen burner attached to a folding portable dryer (De Kesel 2001) and then put inside minigrip bags. Exsiccata were kept at the Abdou Moumouni University (Niamey, Niger) and duplicates were placed in the fungal section of the COFC herbarium (Thiers 2014). Some exsiccata and type material from K and BR herbaria were requested as loans to compare with Nigerien samples and also with protologues.
Colour references were coded following Kornerup & Wanscher (1981). Material was then studied under a Nikon Labophot2 light microscope fitted with a drawing tube at a 1000x magnification. Spore measurements were made in 3% KOH mounts (Menzel- Glaser) in profile position and excluding the hilar appendix and ornamentation (Lm = mean length; Wm = mean width, Em = Lm/Wm). Unless otherwise indicated, the specimens were identified by PP. Daniéls. Species citations follow Index Fungorum (www.indexfungorum.org). Collection localities are placed in alphabetic order. For each collection, field data are ordered as follows: Country, Administrative Region, Department, Locality, Site, {Park}, latitude/longitude coordinates, altitude, ecology and substrate, date, collector and identifier (where appropriate), collection accession number, and herbarium accession number. Distribution notes have been added when this was significant compared to other African flora, and observations are included on nomenclature and taxonomy when these differ from known species descriptions.
Standard methods for DNA extraction, PCR amplification, and DNA sequencing were applied (e.g., Justo et al. 2011) to check Nigerien Volvariella samples due to the noteworthy morphological differences found with closely related species. Primer pairs ITS1F and ITS4 (Gardes & Bruns 1993) were used for both PCR and sequencing. The remaining sequences used in the analysis were retrieved from GenBank and come from the studies of Menolli & Capelari (2008), Li et al. (2009), and Justo et al. (2011). GenBank accession numbers are given under the specimens examined section. Volvariella bombycina (Schaeff.) Singer and V. volvacea (Bull.) Singer were used as outgroup taxa in the final dataset. These two species appear to be the sister group to all other Volvariella species sequenced to date (Vizzini et al. 2011). Sequences were aligned with MAFFT (Katoh et al. 2002) using the Q-INS-i strategy. The alignment was examined and manually corrected in MacClade 4.05 (Maddison & Maddison 2002) and it has been deposited in TreeBASE (http://purl.org/phylo/treebase/phylows/study/TB2:S15058). A Maximum Likelihood analysis was run in RAxML servers (Stamatakis et al. 2008) with 100 rapid bootstrap replicates.
340 ... Daniéls & al. Taxonomy
Clitopilus orientalis TJ. Baroni & Watling, Mycotaxon 72: 58, 1999. PLATE 1
MACROCHARACTERS — BASIDIOMATA small, gregarious. PILEUS 1-2.2 cm diameter, at first convex, then flat to centrally depressed; white coloured [A1] and fibrillose; margin involute and radially ridged. St1pe central or slightly eccentric, slightly turbinate, up to 10-19 x 2-4 mm, concolourous with pileus, surface pruinose, base with white mycelial tufts. LAMELLAE subdecurrent to decurrent, with lamellulae, 3-4/mm, white at first [A1], then brownish pink [5A3, 5B5], with whitish crenulate margin. CONTEXT fibrous; odour and taste not recorded.
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Pate 1. Clitopilus orientalis (COFC-F 5160): a) Pileipellis; b) Basal mycelium; c) Pileus context; d) Spores; e) Basidia; f) Cheilocystidia; g) Stipitipellis showing a secretory hyphae.
New Asian macromycete records for Africa ... 341
MICROCHARACTERS — PILEIPELLIS made up of interlaced 3-5 um wide elements forming a trichodermis; hyphae hyaline, thin-walled. STiprT1PELLIs similar to pileipellis but with inner hyphae parallel, 3-13 um wide; hyphae hyaline, thin-walled, with conspicuous septa. CONTEXT HYPHAE subparallel, cylindrical, 3-8 um wide, thin-walled, sometimes with vesicular end, 12-14 um wide, hyaline and thick-walled. MycELi1aL TuFTs with interlaced cylindrical hyphae, 4-6.5 um wide, hyaline, thin-walled. Secretory hyphae filled with refringent content, 5-13.5 um wide. CLAMP CONNECTIONS absent in all structures. CHEILOCYSTIDIA cylindrical to fusiform or subcapitate, 29-37(-42) x 4-8 um. PLEUROCYsTIDIA absent. Basrp1a claviform, 21-30 x 7-9 um, with 4 sterigmata, thin-walled. Sporss ellipsoid, striate with 10-11 longitudinal ribs, hyaline, thin to thick-walled, (6.3-)6.8-7.5(-8.5) x (3.7-)4-4.7(-5) um, [Lm = 7.1 um, Wm = 4.4 um, E = 1.40-1.88; Em = 1.63].
SPECIMEN EXAMINED — NIGER, Dosso, Gaya, Albarkeizé, Saboula, 12°04’55.6”N 3°14’07.5’E, elevation 136 m, fallow land beside the Niger river on a termite mound under Anacardium occidentale L., 13 September 2008, O. Hama, Hama 132 (COFC-F 5160). EcoLoGcy & DISTRIBUTION — On termite mounds in India and Malaysia (Baroni & Watling 1999). Clitopilus orientalis appears not to have been previously reported in Africa.
ComMEnts — ‘This particular Clitopilus species displays cheilocystidia and grows on termite mounds (Baroni & Watling 1999). The specimen examined differs from descriptions of the Asian species only in that cheilocystidia are abundant and sometimes clustered, rather than scattered. The most closely related species are C. apalus (Berk. & Broome) Petch, which has subglobose rather than ellipsoid spores, and C. peri (Berk. & Broome) Petch, which neither grows on termite mounds nor displays cheilocystidia (Pegler 1977, Baroni & Watling 1999).
Hymenagaricus subepipastus Heinem. & Little Flower, Bull. Jard. Bot. Nat. Belg. 54: 168, 1984. PLATES 2, 3 MACROCHARACTERS — BASIDIOMATA small, gregarious. PILEUS 0.5-1.2 cm diameter, at first convex, then trapezoid to flattened displaying dark olive-green [1E7, 30D7] scales on a light-green background [30A4], changes colour slightly during ageing, the background turns yellow [4A4] while the scales — though retaining their colour — become less abundant, since they are easily detached; margin thin, regular. Stipe central, fistulose, cylindrical, concolourous with pileus, whitish towards the base, up to 22 x 0.6 mm, surface pruinose to squamulose with scarce evanescent scales below ring zone, base
342 ... Daniéls & al.
PLATE 2. Hymenagaricus subepipastus (COFC-F 5186): Fresh basidiomata. Scale bar = 1 cm.
with white mycelial tufts. RING absent or powdery and evanescent, seen only in very young specimens. LAMELLAE free, with lamellulae, 3-4/mm, greyish white at first [B2], then grayish yellow [1B2-3, 2B2-3], and finally brown [5C7], with paler crenulate margin. Spore deposit brown [5C7]. ConTExtT thin, fibrose, in stipe; odour and taste not recorded.
MICROCHARACTERS — PILEIPELLIS made up of claviform, pyriform to spherical elements 7-17 x 7-16 um grouped in chains and tufts forming a hymeniodermis, with thick walls and encrusted brownish orange pigments; subpellis made up of cylindrical elements, interwoven to subparallel, 3.5-8 um, also with a thick wall and encrusted brownish orange pigments, becoming thinner, parallel and hyaline inwards. StT1p1TIpELLis formed by parallel cylindrical elements, 3-9 um wide, often with chains of doliiform (barrel-shaped) to short-claviform terminal elements grouped in tufts; hyphae hyaline to yellowish brown, thin- to thick-walled, somewhat constricted at the septa. CONTEXT HYPHAE subparallel, cylindrical, 4-12 tm wide, thin-walled. MYCELIAL TUFTS with parallel cylindrical hyphae of 2-3.5(-5) um wide, hyaline and thin-walled, sometimes with secretory hyphae, filled with refringent contents. CLAMP CONNECTIONS absent in all structures. CHEILOCYSTIDIA lageniform to fusiform, 16-20(-26) x (5-)6-10(-12) um. PLEUROCYSTIDIA absent. Basip1a widely claviform, 12-15 x 5-7 um, with (2)4 sterigmata, thin-walled. Spores ovoid to ellipsoid, smooth, brown coloured, thin to
New Asian macromycete records for Africa ... 343
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PLATE 3. Hymenagaricus subepipastus (COFC-F 5186): a) Pileipellis; b) Basal mycelium; c) Pileus context; d) Spores; e) Basidia; f) Cheilocystidia; g) Stipitipellis.
thick-walled, (4-)4.5-6.3 x (2.7-)3-4(-4.5) um, [Lm = 5.1 um, Wm = 3.4 um, E = 1.29-1.71; Em = 1.49]. SPECIMEN EXAMINED — NIGER, Niamey, Niamey, Karadjé, guest house at Abdou
Moumouni University, 13°29’25.5”N 2°04’64’E, elevation 200 m, in garden soil, 8 August 2010, P. Daniéls, Hama 383 (COFC-F 5186).
344 ... Daniéls & al.
EcoLocy & DISTRIBUTION — Scattered in the grass or garden soils. Hymenagaricus subepipastus was first described in India (Heinemann & Little Flower 1984), but Heinemann (1986) considered its presence in Africa likely. Our report may be the first record of the species in Africa.
COMMENTS — Our specimen fully matches existing descriptions of H. subepipastus by Heinemann & Little Flower (1984). This species is related to H. viridulus Heinem. & Little Flower, but the spores of the studied collection are larger and ovoid rather than cymbiform (wineskin-shaped). Hymenagaricus subaeruginosus (Berk. & Broome) Heinem. & Little Flower has slightly smaller, cymbiform spores and larger basidiomata, with a pileus up to 4.5 cm diameter (Heinemann & Little Flower 1984). Hymenagaricus epipastus (Berk. & Broome) Heinem. & Little Flower differs from H. subepipastus only in spore size, and has been reported in Ceylon (Asia) (Heinemann & Little Flower 1984). For details see TABLE 2.
TABLE 2. Comparison of pileus and spores from species related to Hymenagaricus subepipastus.
SPECIES PILEUS DIAM. (cm) Lm Wm _ SPORE SHAPE H. viridulus type* 0.5-1.5 4.7 3.3 Cymbiform H. subaeruginosus isotype (K) 1.5-2.5°/ 4.5 47 3.2 Cymbiform H. subaeruginosus MGF 5088 (BR) 1.5-2.5> 4.5 3.0 Cymbiform H. epipastus type * 2 42—6 301 Ovoid
H. subepipastus type * 1-1.5 5.0 3.6 Ovoid
H. subepipastus Hama 383 0.5-1.2 5.l- 34 Ovoid
* Heinemann & Little Flower 1984; measurements in exsiccatum
Termitomyces bulborhizus T.Z. Wei, Y.J. Yao, B. Wang & Pegler, Mycol. Res. 108: 1458, 2004. PLATES 4-6
MACROCHARACTERS — BASIDIOMATA medium sized, gregarious. PILEUS 4-12 cm diameter, fleshy, at first convex, then flattened and slightly depressed with obtuse blunt perforatorium 2-3 mm wide; white, isabella to olivaceous brown coloured [A1, 4A4, 4B6, 5A3] pallescent towards the margin and with brownish orange centre [6D5, 5B6-8]; surface smooth and dry or slightly viscose and radially rugose in wet conditions; margin thin, sinuous to sub-lobate, slightly involute when young then uplifted when old. St1px central, cylindrical, straight, white to slightly isabella coloured [A1], (4—)7-11 x 0.5-2 cm, surface squamulose with evanescent recurved 2-3 mm long scales; bulbose at ground level of 2-3.5 cm diameter. PSEUDORHIzOID robust, rugose and grooved longitudinally, orange brown [4B5, 5C5, 6C8, 6B7] to black [5E3, 6F2-5, F1] downwards, of 5-20 cm long, tapering quickly towards the base. LAMELLAE
New Asian macromycete records for Africa ... 345
Piate 4. Termitomyces bulborhizus: fresh basidiomata: a) COFC-F-5301; b) COFC-F-5305; c) COFC-F-5306. Scale bar = 1 cm.
free to almost free, with lamellulae, crowded, 8-12/cm, to 6 mm wide, white to slightly pink [A1, 6A2] with concolourous margin; spore deposit pink [6A2]. ContTEXT solid, fibrous, <1 cm in pileus; odour sometimes of fresh fish (taste not recorded). Termite mold ellipsoid, 10-20 cm long, grayish isabelline [4A5, 5C2] with irregular 0.5-1.5 cm wide holes.
MACROCHEMICAL REACTIONS — KOH- in pileus; HCl+ dark yellow in pileus.
MICROCHARACTERS — PILEIPELLIS made up of doliiform to cylindrical elements 6-17 x 3-9 um, hyaline with thin wall, somewhat constricted at the septa; hyphal width increasing through context; parallel to interwoven forming a lax ixocutis-subtrichodermis. PSsEUDORHIZzOID formed by parallel cylindrical to doliiform elements, 4-15 um wide in the external zone with often claviform to cylindrical terminal elements; hyphal width increasing through context to 15-35 um wide, hyphae brown with thick wall, constricted at the septa. STIPITIPELLIs similar to pseudorhizoid but hyphae hyaline, 3-5 um wide in the external zone and 7-18 um wide through context. CONTEXT HYPHAE cylindrical, 9-21 um wide with thick 0.5-2 um wall, and mixed with cylindrical hyphae 4-7 um wide, with thin wall. CLamp CONNECTIONS absent
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QanaanQn0avoals OOO b
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PiateE 5. Termitomyces bulborhizus (COFC-F 5306): a) Spores; b) Basidia; c) Cheilocystidia; d) Pileus context; e) Pleurocystidia.
in all structures. CHEILocystTip1A claviform, fusiform to pyriform, (25-) 27-42 x (11-)13-24 um. PLEUROCYSTIDIA similar, sometimes with a secondary septum and/or thick wall, 28-56 x (10-)11-20(-35) um. Basrp1a claviform, 18-24(-27) x 6-8.5 um, with (2-)4 sterigmata and thin to, rarely, thick-walled. Spores ellipsoid, smooth, hyaline, thin-walled, (5.5-)6-7(-8) x (3-)3.8-4.4(-5) um, [Lm = 6.4 um, Wm = 4.1 um, E = (1.38-)1.50-1.75(-1.80); Em = 1.56].
SPECIMENS EXAMINED — NIGER, Dosso, Gaya, Gaya, Gorou Bassounga Forest, 11°54’00’N 3°24’25”E, elevation 209 m, savannah, on termite nest, 5 September 2010, O. Hama, Hama 443b (COFC-F 5302). TILLABERY, Say, Tamou, Haoussa, 12°15’18”N 2°22’14’E, elevation 221 m, wooded Afzelia africana Smith ex Pers. savannah, on underground termite nest, 6 August 2010, O. Hama, Hama 286 (COFC-F 5301); 12°14’50’N 2°22’14”E, 24 August 2011, O. Hama, Hama 470 (COFC-F 5303); 25 August 2011, O. Hama, Hama 475 (COFC-F 5304); 12°14’51”N 2°22’15”E, wooded savannah of Afzelia africana, Combretum glutinosum Perr. ex DC., C. collinum Fres., with Andropogon gayanus Kunth, Crossopteryx febrifuga (Afzel. ex G. Don) Benth., Strychnos spinosa Lam., and Flueggea virosa (Willd.) Royle, on active termite nest, 15 August 2012, O. Hama, Hama 519 (COFC-F 5305); Say, Torodi, Gnaktiré, 12°59’01”N 1°44’32”E, elevation 237 m, cropland, under Sclerocarya birrea (A. Rich.) Hochst., on termite mound, 25 August 2010, O. Hama, Hama 405 (COFC-F 5095); 12°59’09.4”N 1°44’08.5”E, elevation 226 m, ina field of Pennisetum glaucum (L.) R. Br. and Sorghum bicolor (L.) Moench, close to Balanites aegyptiaca (L.) Delile, on a termite nest, 19 August 2012, O. Hama, Hama 559 (COFC-F 5306).
EcoLoGcy & DISTRIBUTION — This taxon has been reported growing on termite nests in southern China (Sichuan, Yunnan) and Thailand (Wei et al.
New Asian macromycete records for Africa ... 347
10 um
Pileipellis
10 um
Pseudorhiza
PLaTE 6. Termitomyces bulborhizus: comparison of pseudorhizoid and pileipellis from exsiccata COFC-F 5306 (left) and isotype K 109284 (right).
2004; Sawhasan et al. 2011). The associated termite species is identified as Macrotermes barneyi Light, native to Vietnam and southern China (Wang et al. 2009) and Hypotermes makhamensis Ahmad in the Thailand collection (Sawhasan et al. 2011). In this first African finding, the associated termite was Macrotermes subhyalinus (Rambur) (Koné, pers. comm.).
ComMMENTS — Termitomyces bulborhizus is a large mushroom with a characteristic black, swollen pseudorhizoid; the stipe surface appears slightly scaly due to the presence of fine floccules.
Our material differs slightly from the isotype of T: bulborhizus (K 109284) in the pileipellis structure that can be justified to different development stages of the fruitbodies. Due to the few notes regarding pileipellis and pseudorhizoid structure of this species, we consider showing them from both isotype and COFC-F 5306 (PLATE 6). Descriptions of referred structures in the isotype are as follows: pileipellis made up of doliiform to cylindrical elements 8-45(-78) x 4-17 um, hyaline with thin wall, somewhat constricted at the septa; hyphal width increasing through context; parallel to interwoven forming a subtrichodermis cutis; pseudorhizoid formed by parallel cylindrical to doliiform elements, 15-32 um wide, hyphae brown with thick wall, constricted at the septa. The pseudorhizoid of the isotype was not clean enough for an accurate analysis so
348 ... Daniéls & al.
we do not know enough to ascertain whether it might also have small hairs or claviform caulocystidia-like structures, as in the Nigerien sample. The bulbose stipe width in Nigerien samples was also smaller (<3.5 cm wide) than those given by Wei et al. (2004). Actually, Asian collections are bigger than African ones regarding basidioma size, with a pileus of (5—)10-22 cm (Wei et al. 2004).
Our material differs from T! eurrhizus (Berk.) R. Heim in having a dark- brown color and a swollen pseudorhizoid (Wei et al. 2004). In the most similar African species (T. umkowaan (Cooke & Massee) D.A. Reid and T. subumkowaan Mossebo) the stipe surface is smooth and the pseudorhizoid is not swollen. Also, T. subumkowaan bears two spores per basidium (Mossebo et al. 2002).
PLATE 7. Tulostoma evanescens (COFC-F 5292): Exsiccatum. Scale bar = 1 cm.
Tulostoma evanescens Long & S. Ahmad, Farlowia 3: 235, 1947. PLATES 7, 8
MACROCHARACTERS — BASIDIOMATA small, gregarious. GLOBOSE HEAD 0.7-0.9 cm diameter. ExoPERIDIUM undefined, hyphal, thin, brown [5B4], intermixed with sand grains. ENDOPERIDIUM membranous, smooth, white [A1]. PERISTOME circular and not prominent, white [A1] because of the absence of exoperidium around the mouth. StT1pe cylindrical or tapering upwards, 1-1.3 x 0.2-0.3 cm, surface squamose with evanescent scales and longitudinally grooved, joined in the base to form a volva-like swollen structure in one basidioma; sometimes with rhizomorphs. Socket lacerate, with several small dentate membranes around the stipe. GLEBA brown coloured [6C6]. CONTEXT firm, fibrous; odour absent and taste not recorded.
New Asian macromycete records for Africa ... 349
000000e0000 |: ; S
PLATE 8. Tulostoma evanescens (COFC-F 5292): a) Exoperidium; b) Context of stipe scales; c) Endoperidium; d) Hyphae of rhizomorph; e) Spores; f) Capillitium.
MICROCHARACTERS — EXOPERIDIUM with cylindrical interlaced hyphae, 1.5-4 um wide with, hyaline to brownish, moderate wall. ENDOPERIDIUM similar, with hyphae 3-6(-8) um wide, hyaline and with thick wall. CAPILLITIUM 3-8 tm wide, with lumen and thick wall minutely encrusted with granules, septate and with a few branches. HYPHAE OF STIPE SCALES parallel, 4-6.5 um wide, grouped in tufts; hyphae hyaline to yellowish, with thin wall, somewhat constricted at the septa. HYPHAE OF RHIZOMORPH 2.5-7 um wide, hyaline, often minutely encrusted with granules and with moderate wall.
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CLAMP CONNECTIONS absent in all structures. BAsIDIA not seen. SPORES ovoid to ellipsoid, smooth, brown coloured, thick-walled, some apiculate, (4.7-) 5-5.4(-6) x (3.7-)4-4.5(-5) um, [Lm = 5.2 um, Wm = 4.1 um, E = 1.11-1.38; Em=126]. SPECIMEN EXAMINED — NIGER, TILLABERY, Say, Tamou, Tamou Total Faunal Reserve {Tamou Reserve}, 12°34’21”N 2°18’28”E, elevation 240 m, wooded savannah, humus- containing sandy soil, 7 August 2010, O. Hama, Hama 361 (COFC-F 5292). ECOLOGY & DISTRIBUTION — ‘This species grows in arid sites and appears after rains in copious quantities. According to Wright (1987), this species has been reported in India and Argentina. This appears to be the first report of this species in Africa.
ComMENTs — The material examined here agrees with earlier descriptions, including the following features: 1) Clearly-defined circular, tubular, or flat peristome on a head of <1 cm in diameter; 2) smooth subglobose or ellipsoid spores measuring about 5-5.4 x 4-4.5 um; 3) inconspicuous hyphal exoperidium; and 4) capillitium 4-10 um wide.
Tulostoma operculatum Long & S. Ahmad has smaller (4-4.7 um diam.) spores and a fimbriate mouth. Tulostoma brevistipitatum B. Liu et al. does not havea volvoid structure at the stipe base, and its spores are slightly longer (5.4-6.1 x 4.3-5 um). Tulostoma fusipes Har. & Pat. is very similar, but its spores are globose, the capillitium is coloured, and the stipe is longer (5.5 x 0.4 cm) (Wright 1987).
Volvariella cf. sathei Senthil., Rahul Sharma & S.K. Singh, Mycotaxon 119: 470, 2012: PLATES 9, 10
MACROCHARACTERS — BASIDIOMATA medium sized, gregarious. PILEUS 3.5-12(-15) cm diameter, fleshy, convex; white to isabella [A1l, 4A2]; surface fibrillose; margin exceeding the lamellae, eroded, or striate with triangular tufts. Stipe central, fistulose when aged, cylindrical and tapering towards apex, straight, concolourous with pileus, 5-12 x 1-2 cm, surface glabrous to fibrillose; slightly bulbose. VoLva membranous, 1-5 x 2-4 cm and 1-1.5 mm thick, white [A1], sometimes with grayish brown hue [4C3]. LAMELLAE free, distant from stipe, with lamellulae, crowded, (10-)13-16(-17)/cm, 3-5 mm wide, at first whitish [Al, 3A2] then pink [9A2, 14A3] with paler, dentate to fibrillose margin. Spore deposit pink [9A3, 10B5]. CONTEXT solid, fibrous; odour disagreeable, farinaceous to fruity, and pleasant taste.
MACROCHEMICAL REACTIONS — KOH- or NaOH- in pileus, context or stipe.
MICROCHARACTERS —PILEIPELLIS made up of cylindrical elements 8-21 um, hyaline with thin to thick wall, somewhat constricted at the septa; parallel, forming a regular cutis. STIPITIPELLIS similar to pileipellis, hyphae
New Asian macromycete records for Africa ... 351
PLATE 9. Volvariella cf. sathei, fresh basidiomata: a) COFC-F-5092; b) COFC-F-5036; c) COFC-F-5033. Scale bar = 3 cm.
hyaline, 4-15 um wide in the external zone and up to 35 um wide through context. CONTEXT HYPHAE cylindrical, 8-21(-28) um wide with thin wall. VoLvaA composed mostly of cylindrical hyphae, 3-5 um wide with scattered inflated elements, cylindrical, fusiform or irregularly shaped, up to 20 um wide. BASAL TOMENTUM formed by interlaced cylindrical elements, 5-22 um wide; hyaline with thin to thick wall. CLamp CONNECTIONS absent in all structures. CHEILOCYSTIDIA fusiform to lageniform, 40-88(-95) x 10-38 um, with hyaline thin wall, sometimes with a basal secondary septum. PLEUROCYSTIDIA similar, 39-80 x 11-24 um. SUBHYMENIUM cellular, with ellipsoid to doliiform hyphae, 10-20 um in diameter. Basrp1a claviform, (19-)22-27(-30) x 7-9 um, with (2—)4 sterigmata and thin-walled. Spores globose to ob-triangular, smooth, brownish pink, thick-walled, (5-)5.6-6(-7) x (4.5-)5.1-5.7(-6.7) um, [Lm = 5.8 um, Wm = 5.4 um, E = 0.92-1.40; Em = 1.07].
SPECIMENS EXAMINED: NIGER, NiaMeEy, Niamey, Abdou Moumouni University,
Faculty of Agricultural Sciences, experimental garden, 13°30'00.5”N 2°05’24.5”E,
elevation 178 m, in a disturbed area, growing on plant residue under Prosopis juliflora (Sw.) DC., 6 August 2009, O. Hama, det. A. Justo, Hama 175 (COFC-F 5036); sandy
352 ... Daniéls & al.
99900000000] E
Nh
e
7
g ie eal PiatE 10. Volvariella cf. sathei (Fics a-e, g—h from COFC-F 5448; Fia. f from COFC-F 5033): a) Pileus context; b) Pileipellis; c) Spores; d) Basidia; e) Cheilocystidia; f) Volva, g) Basal tomentum, h) Stipitipellis.
soil with cattle manure under Prosopis juliflora, O. Hama, Hama 176 (COFC-F 5092); Niamey, Abdou Moumouni University, High School of Education, 13°30’07”N 2°05’29’E, elevation 187 m, in a disturbed area, in sandy soil near Azadirachta indica A. Juss., 4 August 2010, O. Hama, det. A. Justo, Hama 390 (COFC-F 5088; Genbank KF926666). TILLABERY, Say, Torodi, Fayra, 13°01’26.7”N 1°45’39.4’E, elevation 232 m, in arable fields with plant residue of Piliostigma reticulatum (DC.) Hochst., 21 August
New Asian macromycete records for Africa ... 353
2008, O. Hama, Hama 38 (COFC-F 5448); Say, Torodi, Gnaktiré, 12°59’02”N 1°44’49’E, elevation 222 m, cropland, growing on organic residue near houses with Pennisetum glaucum and Sorghum bicolor, 16 August 2009, O. Hama, det. A. Justo, Hama 194 (COFC-F 5034; Genbank KF926664); 12°59’01”N 1°44’32”E, elevation 221 m, in arable fields on heaped-up plant residue with Sorghum and Pennisetum, 15 August 2009, O. Hama, det. A. Justo, Hama 193 (COFC-F 5035; Genbank KF926663); Say, Tamou, Mékrou, {W National Park}, 12°15’16”N 2°23’24’E, elevation 218 m, in gallery forests with Cola laurifolia Mast., Diospyros mespiliformis Hochst. ex A. DC., Mitragyna inermis (Willd.) Kuntze, on sandy-clay soils, 22 August 2009, O. Hama, Hama 204, det. A. Justo (COFC-F 5033; Genbank KF926665).
EcoLoGy & DISTRIBUTION — ‘The authentic species, Volvariella sathei, was recently proposed, based on Indian material (Senthilarasu et al. 2012), and has not been reported elsewhere.
ComMENTsS — The large white species, V. sathei, differs from V. nivea 'T.H. Li & Xiang L. Chen in having ovoid to widely ellipsoid spores and larger cystidia (Senthilarasu et al. 2012).
Our samples agree mostly with the description given by Senthilarasu et al. (2012) except for: (1) the presence of fusiform to lageniform cystidia (cylindro- clavate in V. sathei); (2) spores often with a triangular shape; and (3) grayish hue sometimes present in volva. Molecular analysis indicates that the African isolates are in the same clade as the Indian V. sathei but that there are some differences between them (PLATE 11). Additional data are needed to confirm whether the African isolates represent V. sathei or a closely related taxon.
89°>- HM246492 Volvariella hypopithys TOAV137 Sardinia '— JF415139 Volvariella sp LOU18924 Spain af HM246494 Volvariella pusilla Sardinia TOAV139 100 JF415137 Volvariella pusilla AJ51 Spain 98 -—— HM246493 Volvariella strangulata Italy TOAV141 '—— HM562213 Volvariella surrecta AJ55 Spain 95 '—— JF415141 Volvariella terrea LUG 11010 Holotypus France JF415140 Volvariella dunensis SCM3513 Spain 100 JF415138 Volvariella dunensis RFS 07031003 Spain | JF415136 Volvariella dunensis TOAV140 Sardinia 86 HM 246491 Volvariella taylorii TOAV142 Sardinia HM562214 Volvariella lepiotospora AJ155 USA 0.06 |_| -___________ 562211 Volvariella caesiotincta MA54717 Spain HM246500 Volvariella sp TOAV143 Slovenia HM562210 Volvariella taylorii AJ54 Portugal g3)KF 926664 Volvariella cf. sathei Hama194 Niger KF926663 Volvariella cf. sathei Hama193 Niger KF926665 Volvariella cf. sathei Hama204 Niger | erszeee Volvariella cf. sathei Hama390 Niger JN792550 Volvariella sathei AMH 9436 India HM562212 Volvariella bombycina AJ244 Spain FJ379274 Volvariella volvacea VV34 PLaTE 11. Best tree from the Maximum Likelihood analysis of ITS sequences of Volvariella. Nigerien samples of V. cf. sathei are emphasized. Scale bar indicates nucleotide substitutions/site.
100
100
354 ... Daniéls & al.
Discussion
The taxa examined here are species previously recorded in Asia; this is the first time they have been reported on the African continent. Fungal species common to both continents are frequently reported, perhaps due to climatic and/or ecological similarities, spore dispersal capacity, and shared geological history; cases of vicariance and species fragmentation are also found (Berndt 2002, O'Donnell et al. 2011, Geml et al. 2008, Moncalvo & Buchanan 2008). The fragmentation of biogeographical ranges may sometimes be due to human intervention and particularly to the introduction of exotic plant species with their own associated fungal suites (Niveiro et al. 2009, Chen et al. 2006, Read 2000, Pringle et al. 2009); some ectomycorrhizal fungi entered Africa by this means (Duponnois & Galiana 2007, Garbaye et al. 1988). The climate of the West Sudanian savannah is similar to that of the Asian savannah of India, Laos, Thailand, Myanmar, Vietnam, and Cambodia (Young & Solbrig 1993). There are, however, differences in floral composition, with Dipterocarpaceae predominating in Asia and Leguminosae in Africa. This can help to discriminate some tree-restricted ectomycorrhizal species better than saprobes. However, there are some wide-ranging mycorrhizal fungi, which could also suggest a lack of fungal sampling in this ecosystem type (Tulostoma evanescens might represent such a case). Actually, infrequent sampling of the African savannah may have led to a bias in our knowledge of the biogeographical range, substrate versatility, spore dispersal capacity, and ecological similarity of these species. Furthermore, some saprobic species are often unspecific regarding substrate degradation and sometimes the substrate itself is originated by fire or domestic herbivore digestion, reducing substrate differences between distant geographic areas.
Mushrooms associated with termite nests grow only where termites are found. Several termite genera and species (e.g. Macrotermes, Microtermes, and Odontotermes) are common to both continents (Cheng and al. 2011) and this kind of mycological association seems to have originated in West Africa (Nobré et al. 2011). Clitopilus orientalis is a saprobic fungus with an apparent preference for termite nests.
Our discoveries provide a promising basis for further research into Niger’s fungal diversity. Many collections are yet to be reviewed, and their examination will doubtless enhance our knowledge of fungal diversity in the West Sudanian savannah as a whole, and also enable a closer appraisal of its similarity with its Asian counterpart. More samples and DNA-analysis of the studied material are needed to discern cryptic species, and Volvariella cf. sathei may be a geographic variety of the Asian species.
New Asian macromycete records for Africa... 355
Acknowledgments
This study was part of the “Edible and cultivable Macromycetes of Niger (Ethnomycology)” project, financed by the Spanish Agency for International Development Cooperation (AECID-C/023163/09; D/031488/10; A1/039675/11). The authors are grateful to the curators of the COFC, K and BR herbaria for the exsiccata loan management. They would also like to thank the mycologists T.J. Baroni (State University New York, Cortland) and L.A. Parra (Aranda de Duero) for providing assistance, comments, and taxonomic information regarding the identification of certain species; also thanks to Nourou Soulemane Yorou (University of Miinchen) and Gabriel Moreno (University of Alcala de Henares) for their critical revision of the manuscript. We are indebted to N’Golo Abdoulaye Koné (University of Abobo-Adjamé) for the identification of Macrotermes subhyalinus and to M. Robichaud (Ottawa) for the English revision of the manuscript.
Literature cited
Adewusi SRA, Alofe FV, Odeyemi OA, Oke OL. 1993. Studies on some edible wild mushrooms from Nigeria: 1. Nutritional, teratogenic and toxic considerations. Plants Foods for Human Nutrition 43: 115-121. http://dx.doi.org/10.1007/BF01087916
Akpaja EO, Isikhuemhen OS, Okhuoya JA. 2003. Ethnomycology and usage of edible and medicinal mushrooms among the Igbo People of Nigeria. International Journal of Medicinal Mushrooms 5(3): 313-319. http://dx.doi.org/10.1615/InterJ MedicMush.v5.i3.100
Antonin V. 2013a. Monograph of Crinipellis and Chaetocalathus in tropical Africa. Fungus Flora of Tropical Africa 3: 1-41.
Antonin V. 2013b. Supplements to the monograph of tropical African species of Marasmius (Basidiomycota, Marasmiaceae). Cryptogamie Mycologie 34(2): 113-135. http://dx.doi.org/10.7872/crym.v34.iss2.2013.113
Baroni TJ, Watling R. 1999. Taxonomic and mycogeographic notes on some Malaysian fungi IV. Notes on Clitopilus and Rhodocybe. Mycotaxon 72: 57-72.
Berndt R. 2002. Uromyces vankyorum sp. nov. and Uromyces atriplicis, putative vicariant rust species of Patagonian and Australian semi-desert biomes. Mycological Progress 1(2): 179-186. http://dx.doi.org/10.1007/s11557-006-0017-8
Boa E. 2004. Wild edible fungi. A global overview of their use and importance to people. Non- Wood Forest Products 17. FAO, Roma.
Chen YL, Kang LH, Malajczuk N, Dell B. 2006. Selecting ectomycorrhizal fungi for inoculating plantations in south China: effect of Scleroderma on colonization and growth of exotic Eucalyptus globulus, E. urophylla, Pinus elliottii and P. radiata. Mycorrhiza 16(4): 251-259. http://dx.doi.org/10.1007/s00572-006-0039-8
Cheng S, Kirton LG, Panandam JM, Siraj SS, Ng KK-S, Tan S-G. 2011. Evidence for a higher number of species of Odontotermes (Isoptera) than currently known from Peninsular Malaysia from mitochondrial DNA phylogenies. PLoS ONE. http://dx.doi.org/10.1371/journal.pone.0020992
De Kesel A. 2001. A mushroom dryer for the travelling mycologist. Field Mycology 2: 131-133. http://dx.doi.org/10.1016/S1468-1641(10)60534-8
De Kesel A, Yorou NS. 2000. Etude préliminaire des macromycetes associés aux foréts claires du Nord Bénin (Afrique de POuest). Scripta Botanica Belgica 20: 26.
De Kesel A, Codjia JT, Yorou NS. 2000. Connaissances ethnomycologiques des peuples Nagot du centre du Bénin. Scripta Botanica Belgica 20: 27.
356 ... Daniéls & al.
De Kesel A, Codjia JT, Yorou NS. 2002. Guide des champignons comestibles du Bénin. Jardin Botanique National, Meise.
De Kesel A, Guelly AK, Yorou NS, Codjia JT. 2008. Ethnomycological notes on Marasmiellus inoderma from Benin and Togo (West Africa). Cryptogamie Mycologie 29(4): 313-319.
Duponnois R, Galiana A. 2007. Ectomycorrhizal inoculation of fast growing tree species in the tropics: potential benefits in sustainable rehabilitation of degraded soils. 33-50, in: AK Skaggs (ed.). New research on forest ecology. Nova Science Publishers, New York.
Eyi-Ndong H, Degreef J, De Kesel A. 2011. Champignons comestibles des foréts denses d'Afrique centrale. ABC taxa 10: 1-254.
FAO. 2010. FAOSTAT, FAO Statistical Databases. http//faostat.fao.org/ [last access 21-I-2010].
Ganaba S, Ouadba JM, Bognounou O. 2002. Utilisation des ressources végétales spontanées comme complément alimentaire en région sahélienne du Burkina Faso. Annales de Botanique de l'Afrique de Ouest 2: 125-142.
Garbaye J, Delwaulle JC, Diangana D. 1988. Growth response of Eucalyptus in the Congo to ectomycorrhizal inoculation. Forest Ecology and Management 24: 151-157. http://dx.doi.org/10.1016/0378-1127(88)90118-1
Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes — application to the identification of mycorrhizae and rusts. Molecular Ecology 2: 113-118. http://dx.doi.org/10.1111/j.1365-294X.1993.tb00005.x
Gardt S, Yorou NS, Guissou ML, Guelly AK, Agerer R. 2011. Amaurodon angulisporus (Basidiomycota, Fungi), a new species from West Africa identified by molecular and anatomical features. Nova Hedwigia 93(1-2): 237-247. http://dx.doi.org/10.1127/0029-5035/2011/0093-0237
Geml J, Tulloss RE, Laursen GA, Sazanova NA, Taylor DL. 2008. Evidence for strong inter- and intracontinental phylogeographic structure in Amanita muscaria, a wind-dispersed ectomycorrhizal basidiomycete. Molecular Phylogenetics and Evolution 48: 694-701. http:// dx.doi.org/10.1016/j-ympev.2008.04.029
Guissou KML. 2005. Inventaire des macromycétes du Burkina Faso: ethnomycologie, valeurs nutritionnelle et thérapeutique de quelques espéeces. Dissertation, University of Ouagadougou, Burkina Faso.
Guissou KML, Sankara P, Guinko S. 2002. Les champignons macroscopiques de la Réserve de la Mare aux Hippopotames de Bala: diversité et étude ethnomycologique. Etudes de la flore et la végétation de Burkina Faso 7: 45-48.
Guissou KML, Lykke AM, Sankara P, Guinko S. 2008. Declining wild mushroom recognition and usage in Burkina Faso. Economic Botany 62(3): 530-539. http://dx.doi.org/10.1007/s12231-008-9028-5
Halling RE. 1996. Recommendations for collecting mushrooms for scientific study. 135-141, in: MN Alexiades, JW Sheldon (eds). Selected Guidelines for Ethnobotanical Research: a Field Manual. The New York Botanical Garden Press, Bronx.
Hama O, Barage M, Marafa D, Adam T, Saadou M. 2008. Inventaire et caractérisation des macromycetes des rives du moyen Niger. Annales de Université Abdou Moumouni de Niamey (Sér. A) 9: 1-12.
Hama O, Barage M, Marafa D, Adam T, Saadou M. 2009. Détermination de quelques constituants nutritionnels er anti-nutritionnels sur 6 especes de macromycetes collectées sur les rives du moyen Niger. Annales de Université Abdou Moumouni de Niamey (Sér. A) 10: 35-42.
Hama O, Maes E, Guissou ML, Ibrahim D, Barage M, Parra Sanchez LA, Raspé O, De Kesel A. 2010. Agaricus subsaharianus, une nouvelle espece comestible et consommée au Niger, au Burkina Faso et en Tanzanie. Cryptogamie Mycologie 31(3): 221-234.
New Asian macromycete records for Africa ... 357
Hama O, Ibrahim D, Baragé M, Alhou B, Daniéls PP, Infante F. 2012. Utilisations de quelques especes de macromycetes dans la pharmacopée traditionnelle au Niger. Journal of Applied Biosciences 57: 4159-4167.
Heinemann P. 1986. Agariceae (Agaricaceae). Flore Illustrée des Champignons d'Afrique Centrale 12: 249-262.
Heinemann P, Little Flower. 1984. Hymenagaricus (Agaricaceae) de Kerala (Inde) et de Sri Lanka. Bulletin du Jardin Botanique National de Belgique 54: 341-346. http://dx.doi.org/10.2307/366787 1
Justo A, Vizzini A, Minnis AM, Menolli Jr N, Capelari M, Rodriguez O, Malysheva E, Contu M, Ghingnone S, Hibbett DS. 2011. Phylogeny of the Pluteaceae (Agaricales, Basidiomycota): taxonomy and character evolution. Fungal Biology 115: 1-20. http://dx.doi.org/10.1016/j.funbio.2010.09.012
Katoh K, Misawa K, Kuma K, Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acid Research 30: 3059-3066. http://dx.doi.org/10.1093/nar/gkf436
Koné NA, Koné D, Nicot P. 2010a. State of knowledge of fungal diversity in Cote d'Ivoire. 172-177, in: S Konaté, D Kampmann (eds). Biodiversity Atlas of West Africa, Tome III.
Koné NA, Konaté S, Linsenmair EK. 2010b. Socio-economic importance of Termitomyces in Cote d'Ivoire. 177-178, in: S Konaté, D Kampmann (eds). Biodiversity Atlas of West Africa, Tome III.
Kornerup A, Wanscher JH. 1981. Taschenlexikon der Farben, 3rd edn. Muster-Schmidt, Gottingen.
Li TH, Chen XL, Shen YH, Li T. 2009. A white species of Volvariella (Basidiomycota, Agaricales) from southern China. Mycotaxon 109: 255-262. http://dx.doi.org/10.5248/109.255
Maddison DR, Maddison WP. 2002. MacClade4: analysis of phylogeny and character evolution. Sinauer Associates, Sunderland, Massachusetts.
Menolli Jr N, Capelari M. 2008. Records and two new species of Volvariella (Pluteaceae, Agaricales) from Brazil. Mycotaxon 106: 385-398.
Moncalvo JM, Buchanan PK. 2008. Molecular evidence for long distance dispersal across the Southern Hemisphere in the Ganoderma applanatum-australe species complex (Basidiomycota). Mycological Research 112(4): 425-436. http://dx.doi.org/10.1016/j.mycres.2007.12.001
Mossebo DC, Amougou A, Atangana RE. 2002. Contribution a létude du genre Termitomyces (Basidiomycetes) au Cameroun: écologie et systématique. Bulletin de la Société Mycologique de France 118(3): 195-249.
Niveiro N, Popoff OF, Alberto EO. 2009. Edible wild mushrooms: exotic species of Suillus (Boletales, Basidiomycota) and Lactarius (Russulales, Basidiomycota) associated to culture of Pinus elliottii in northeastern Argentina. Bonplandia 18(1): 65-71.
Nobré T, Koné NA, Konaté S, Linsenmair KE, Aanen DK. 2011. On the origin and co-diversification of fungus-growing termites and their fungal symbionts. Molecular Ecology 20: 2619-2627. O’Donnell K, Rooneya AP, Mills GL, Kuo M, Weber NS, Rehner SA. 2011. Phylogeny and historical biogeography of true morels (Morchella) reveals an early Cretaceous origin and high continental endemism and provincialism in the Holarctic. Fungal Genetics and Biology 48:
252-265. http://dx.doi.org/10.1016/j.fgb.2010.09.006
Okhuoya JA, Akpaja EO, Osemwegie OO, Oghenekaro AO, Ihayere CA. 2010. Nigerian mushrooms: underutilized non-wood forest resources. Journal of Applied Science and Environmental Management 14(1): 43-54.
Osagualekhor DO, Okhuoya JA. 2005. Sociocultural and ethnomycological uses of mushrooms among the Esan people of Nigeria. International Journal of Medicinal Mushrooms 7: 442-443. http://dx.doi.org/10.1615/Int)MedMushr.v7.i3.810
358 ... Daniéls & al.
Osemwegie OO, Eriyamremu GE, Abdulmalik J. 2006. A survey of macrofungi in Edo/Delta region of Nigeria, their morphology and uses. Global Journal of Pure and Applied Sciences 12(2): 149-157. http://dx.doi.org/10.4314/gjpas.v12i2.16583
Oso BA. 1975. Mushrooms and the Yoruba people of Nigeria. Mycologia 67: 311-319. http://dx.doi.org/10.2307/3758423
Oso BA. 1977. Mushrooms in Yoruba Mythology and Medicinal practices. Economic Botany 31: 367-371. http://dx.doi.org/10.1007/BF02866888
Pegler DN. 1977. A preliminary agaric flora of East Africa. Kew Bulletin Additional Ser. 6. 615 p.
Pringle A, Adams RI, Cross HB, Bruns TD. 2009. The ectomycorrhizal fungus Amanita phalloides was introduced and is expanding its range on the west coast of North America. Molecular Ecology 18: 817-833. http://dx.doi.org/10.1111/j.1365-294X.2008.04030.x
Read DJ. 2000. The mycorrhizal status of Pinus. 324-340, in: DM Richardson (ed.). Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge.
Sawhasan P, Worapong J, Vinijsanun T. 2011. Morphological and molecular studies of selected Termitomyces species collected from 8 districts of Kanchanaburi Province, Thailand. Thai Journal of Agricultural Science 44(3): 183-196.
Senthilarasu G, Sharma R, Singh SK. 2012. A new species of Volvariella from India. Mycotaxon 119: 467-476. http://dx.doi.org/10.5248/119.467
Stamatakis A, Hoover P, Rougemont J. 2008. A rapid bootstrap algorithm for the RAxML Web servers. Systematic Biology 75: 758-771. http://dx.doi.org/10.1080/10635 150802429642
Thiers B. 2014. Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. http://sciweb.nybg.org/science2/IndexHerbariorum.asp [accessed March 2014].
Vizzini A, Contu M, Justo A. 2011. Additional records of Volvariella dunensis (Basidiomycota, Agaricales): morphological and molecular characterization. Mycotaxon 117: 37-43. http://dx.doi.org/10.5248/117.37
Wang ZY, Mo JC, Lu YJ. 2009. Biology and ecology of Macrotermes barneyi (Isoptera: Termitidae). Sociobiology 54(3): 777-786.
Wei T-Z, Yao Y-J, Wang B, Pegler DH. 2004. Termitomyces bulborhizus sp. nov. from China, with a key to allied species. Mycological Research 108(12): 1458-1462. http://dx.doi.org/10.1017/S0953756204001042
White F. 1986. La végétation de 1’ Afrique. ORSTAM-UNESCO, Paris.
Wright JE. 1987. The genus Tulostoma (Gasteromycetes) - a world monograph. Bibliotheca Mycologica 113. 338 p.
Yorou NS. 2010. Champignons supérieurs/larger fungi. 324-331, in B Sinsin, D Kampmann (eds). Biodiversity Atlas of West Africa. Volume I (Benin) - Current state of plant diversity. BIOTA West Africa, Frankfort/Main.
Yorou NS, De Kesel A. 2002. Connaissances ethnomycologiques des peuples Nagot du centre du Bénin (Afrique de l'Ouest). Systematics and Geography of Plants 71: 627-637. http://dx.doi. org/10.2307/3668707
Yorou NS, De Kesel A. 2011. Larger fungi. 47-60, in: P Neuenschwander et al. (eds). Nature Conservation in West Africa: Red List for Benin. International Institute of Tropical Agriculture. Ibadan.
Yorou NS, De Kesel A, Codjia JTC, Sinsin B. 2002a Biodiversité des champignons comestibles du Bénin. 231-240, in: Proceedings Symposium-Workshop on Biodiversity in Benin. Abomey- Calavi (Benin) October 30th to November 18th 2002.
New Asian macromycete records for Africa ... 359
Yorou NS, De Kesel A, Sinsin B, Codjia JT. 2002b. Diversité et productivité des champignons comestibles de la forét classée de Wari-Maro (Bénin, Afrique de l'Ouest). Systematics and Geography of Plants 71: 613-625. http://dx.doi.org/10.2307/3668706
Yorou NS, Koné NA, Guissou KML, Guelly AK, MAba DL, Ekue MR, De Kesel A. 2014. Biodiversity and sustainable use of wild edible fungi in the Sudanian centre of endemism: a plea for valorization. 241-269, in: AM Ba et al. (eds). Ectomycorrhizal symbiosis in tropical and Neotropical Forests. Taylor and Francis Group, Boca Raton, Florida.
Young MD, Solbrig OT. 1993. The world’s savannas: economic driving forces, ecological constraints and policy options for sustainable land. Man and the Biosphere Series. UNESCO Publishing / Parthenon Publishing Group, Paris.
Zoberi MH. 1973. Some edible mushrooms from Nigeria. Nigerian Field 38: 81-90.
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MY COTAXON
http://dx.doi.org/10.5248/130.361 Volume 130, pp. 361-367 April-June 2015
Contribution to the knowledge of Inonotus baumii in Thailand
ANON AUETRAGUL’, ORATAI EUATRAKOOL?’, MarIA LETIZIA GARGANO}, & GIUSEPPE VENTURELLA?
' AnonBiotec Institute, Department of Biotech, The North Chiangmai University, Talad Thai, Klongsong, Klongluang, Patumthani 12120, Chiangmai, Thailand ? Office of Agricultural Regulation, Department of Agriculture, Chatuchak, Bangkok, 10900 (Thailand) ° Department of Agricultural and Forest Sciences, University of Palermo, Viale delle Scienze, Bld. 4, I-90128 Palermo (Italy)
* CORRESPONDENCE TO: giuseppe.venturella@unipa. it
ABSTRACT —Inonotus baumii in Thailand is reported on three new host-plants: Acacia tomentosa, Shorea robusta, and S. siamensis. A key for the identification of Inonotus species in Thailand accompanies notes on the taxonomy of I. baumii.
Key worps — Basidiomycota, Hymenochaetales, Phellinus, white rot fungus
Introduction
Fungi are one of the most species-rich groups of organisms but compared to plants they are still vastly under-investigated (Hawksworth 1997). In particular there is still a lack of distribution and ecological data on several fungal species (Venturella et al. 2011). This is even more true for wood-inhabiting fungi, which have become increasingly important in recent years not only for their ecological role but also as natural sources of anticancer, antioxidative, and antibacterial agents (Karaman et al. 2010).
Several wood-decaying species are used in Asian folk medicine (Larsson et al. 2006). An example is Inonotus baumii, sometimes erroneously named Phellinus linteus (Berk. & M.A. Curtis) Teng, which is known for its use in traditional Chinese medicine (Ying et al. 1987, Shon et al. 2003). Larsson et al. (2006) have researched the perplexing taxonomy of Hymenochaetales, while Tura et al. (2011) have conducted cultural-morphological investigations on
362 ... Auetragul & al.
I. baumii [sub Phellinus baumii] and P. linteus. According to these authors, the taxonomic delimitation at the generic and species level in Phellinus s.l. and Inonotus s.l. is confusing, and P. linteus and I. baumii [sub P. baumii] show close phylogenetic relationships within the P. linteus species complex.
Wu et al. (2012) investigated the species delimitation of the I. baumii- I. linteus group (including taxa recorded as P. linteus) and included the “sanguangh mushroom, a popular polypore used medicinally in China, Japan, and Korea. They described the “sanguangh mushroom” as a new species (Inonotus sanghuang Sheng H. Wu et al.) growing solely on Morus sp. in China, Japan, Korea, and Taiwan. Wu et al. (2012) also reported I. baumii on living Syringa sp. distributed in temperate NE Asia. Parmasto & Parmasto (2001) highlighted the need for correctly identifying the host species when collecting Phellinus species on trees or bushes in tropical and subtropical areas. The effects of Cambodian P linteus in the wood chemistry and structure of Shorea obtusa Wall. ex Blume were investigated by Srivilai et al. (2013).
The aim of this paper is to contribute to the knowledge of Inonotus baumii in Thailand and to report three new host plants.
Materials & methods
The studied basidiomata were collected in different periods of the year in Chiangmai (Mai Hongson Province) and the northeastern region of Thailand; the morphological characters of basidiomata kept in the personal collection of one of the authors (A. Auetragul) were also examined. The morphology of fresh and dried basidiomata was analyzed in the Laboratory of Mycology of the Department of Agricultural and Forest Sciences (University of Palermo, Italy). The basidiospore length and width, mean number of pores per mm, mean size of setae, and diameter of tramal skeletal hyphae were determined according to Parmasto & Parmasto (2001). Morphological characters were also evaluated according to the identification keys of Wu et al. (2012) and Tian et al. (2013), which consider the hymenial surface pore sizes, basidiospore length and width, basidioma type, pileus surface characters, color of the pileal surface marginal zone, and host-plants. The host-plants on which we collected the basidiomata were identified according to Graf (1992). Fungal nomenclature follows MycoBank. The dried herbarium specimens are kept in the collection of AnonBiotec Institute, Department of Biotech, The North Chiangmai University, Thailand (DBNC) and the Herbarium, Department of Agricultural and Forest Sciences, University of Palermo, Italy (SAF).
Taxonomy
Inonotus baumii (Pilat) T. Wagner & M. Fisch., Mycologia 94: 1009 (2002) PLaTE 1
Basidiomata woody, perennial, sessile, semicircular, ungulate or imbricate, 7 x 15 cm, 5-6 cm thick at the base, pileal surface dark grayish brown to dark gray, densely sulcate, radially rimose and cracking; margin rounded; hymenophores golden yellow when unripe then dark brown with wide and
Inonotus baumii in Thailand ... 363
PLATE 1. Inonotus baumii basidiomata (photos courtesy of AnonBiotec). A, from Shorea robusta (DBNC 021, SAF 038); B. from Acacia tomentosa (DBNC 020).
sterile zone along the margin; pores regular, circular 8-10 per mm; tube layers receding in old basidiomata; context woody. Hyphal system dimitic; generative hyphae simple-septate, hyaline to golden yellow, 2-2.5 um wide; skeletal hyphae golden yellow to golden ferrugineous, thick-walled, up to 4-5 um wide; hymenial setae frequent to scanty, ventricose with an acute apex, thick-walled at the base, 20-27 x 5-13 Um; basidia clavate, 7-12 x 5-8 um, tetrasporic; basidiospores broadly ellipsoid to subglobose, hyaline then pale yellowish brown, thin-walled, 6 x 5 um.
SPECIMENS EXAMINED: THAILAND. NORTHERN THAILAND: CHIANG MAI PROVINCE,
Chiang Mai, UTM 2077469 498244 47Q, 310 m, on living and dead trees of Acacia
tomentosa Willd. (Leguminosae), 20 Oct 2012, A. Auetragul (DBNC 020); Maz Hone
SON ProvincE, Mae Hong Son, UTM 2134467 391785 47Q, 240 m, on living and dead
trees of Shorea robusta C.F. Gaertn. (Dipterocarpaceae), 17 Oct 2013, A. Auetragul
(DBNC 021, SAF 038); LaMpaNG Province, Lampang, UTM 552842 2023449 47Q, 244
m, on living and dead trees of Shorea siamensis Miq. (Dipterocarpaceae), 25 Oct 2013, A. Auetragul (DBNC 022).
Identification key for Inonotus spp. in Thailand (partly modified from Wu et al. 2012) bPorés or hymenial Surface 5/ DANG tee tee eee cote eon «eee I. lonicerinus LpPoresiol- hymemial-siintace! S5/ Ini .a5 Bae wtt Boe Ae AE ae hin eee ae Ai Bi 2 2. On dead branch of Lonicera spp. Basidiospores mostly <4. 1 x 3.1 um I. lonicericola 2. Not on Lonicera spp. Basidiospores mostly >4.1 X 3.LUMm.............. 0... eee 3 3. On fallen trunks of Populus spp. Basidiomata resupinate or pileate. Pileus surface indistinctly or moderately sulcate with furrows <3/cm. Pileus always with a wide yellowsmarginalzone onthe surtace ss: bu 5 aes es ae cae I. vaninii 3. Not on Populus spp. Basidiomata pileate. Pileus surface densely sulcate with furrows >3/cm. Pileus usually lacking a wide yellow marginal zone on the surface in ripe basidioniata: (Asc. tins -ehines Hein aioe eRince tein? Mme + eee + elds om taht om, 4
364 ... Auetragul & al.
4. On Weigela coraeensis. Basidiomata sessile to effused-reflexed. Pileus applanate, frequently becoming decurrent with more or less effused base, surface with a thin and black crust, margin usually thin and acute................ I. weigelae
4. Not on Weigela. Basidiomata always sessile. Pileus applanate to concave or subungulate; surface without a thin crust. Pileus surface sometimes black and crustose in ripe basidiomata. Pileus margin acute or blunt ................... 5
5. On Morus spp. Basidiomata perennial, sessile, pileate. Pileus applanate or applanate with an umbo to convex, adaxially flat, slightly convex or slightly concave. Pore surface golden-yellow, brownish yellow to yellowish brown. Pileus margin turmine-dark=redewithyi@ Flt ie: he biG aphasia aa I. sanghuang
5. On dead and living trees of Syringa sp., Acacia tomentosa, Shorea robusta, S. siamensis. Basidiomata always sessile, applanate to subungulate, adaxially more or less concave, margin acute or blunt, thin crust absent, although pileus surface may become black and crustose in ripe basidiomata, surface densely sulcate, with farrows >3/cm. Aged pileus usually lacking a wide yellow marginal zone on pileus surface. Pileus margin not turning dark-red with KOH. Pore surface yellowish brown. Pores of hymenial surface >5/mm. Basidiospores more or less sphiaerical,6im long:and Siti wide ie . casege seeing eae) ees wee I. baumii
Inonotus baumii (Hymenochaetaceae) belongs to the I. baumii-I. linteus group distributed in temperate NE Asia. It has been recorded on living and dead trees of Syringa (Wu et al. 2012) in China (Dai 2010), Japan (Ito 1955), Korea (Lim et al. 2003), and Far East Russia (Parmasto & Parmasto 2001). Phellinus linteus has been reported to grow on Quercus L., Cassia L., Lonicera L., Morus L., Symplocos Jacq., and Hemiptelea Planch. (Ahmad 1972, Kang et al. 2002, Larsen & Cobb-Poulle 1990, Lim et al. 2003). However, Lim et al. (2003), who investigated basidiomata from China, Korea, Costa Rica and, Mexico, showed that the temperate East Asian species known as “P. linteus” represents I. baumii. Wt et al. (2012) proved that specimens collected on Morus species represent another taxon, I. sanghuang.
Discussion
The difficulties in identifying Inonotus and Phellinus spp. belonging to the P. linteus species complex fuel a wide debate on the taxonomic status of the collected basidiomata growing on different wild and ornamental trees. This is also due to the great interest by scientists and people towards the medicinal use of the mushrooms identified as P. linteus and/or I. baumii. The worldwide scientific movement on medicinal mushrooms and the mushroom industry (mainly in Asian countries) specifically requires an exact identification of mushroom species (Wasser 2013). According to our investigation the host range of the P. linteus species complex is not exhaustive. We report Inonotus baumii for the first time on three new host plants growing in Thailand: Acacia tomentosa (PLATE 2), Shorea robusta (PLATE 2), and S. siamensis (PLATE 3).
Inonotus baumii in Thailand ... 365
PLATE 2. (A) Acacia tomentosa in Chang Mai (Thailand); (B) Shorea robusta in Mae Hong Son.
a
2,
PiaTE 3. Shorea siamensis in Lampang (Thailand).
366 ... Auetragul & al.
These plants are cultivated as ornamentals in Chiang Mai, Mae Hong Son, and also in Lampang. Depending on its host, different vernacular names are attributed to I. baumii in Thailand: the local name of Acacia tomentosa is “Kra-Tin-Piman” and I. baumii on this host is called “Hed Kra-Tin-Piman” (Hed = mushroom); the local name of Shorea robusta is “Jigg” (North Thailand) or “Keng” (North-Eastern Thailand) so I. baumii on this host is known as “Hed Jigg” or “Hed Keng”. Given that I. baumii is collected by a large number of people attracted by the medicinal properties of the basidiomata, the above identification key should enable non-specialists also to identity I. baumii.
Acknowledgments The authors wish to thank Prof. Georgios Zervakis (Greece) and Prof. Omon Isikhuemhen (USA) for carefully reading the manuscript and for English revision.
Literature cited
Ahmad S. 1972. Basidiomycetes of West Pakistan. Monograph Biological Society of Pakistan 6: 1-141.
Dai YC. 2010. Hymenochaetaceae (Basidiomycota) in China. Fungal Diversity 45: 131-343. http://dx.doi.org/10.1007/s13225-010-0066-9
Graf AB. 1992. Tropica. Color cyclopedia of exotic plants and trees. Roehrs Co. 1152 p.
Hawksworth DL. 1997. Orphans in “botanical” diversity. Muelleria 10: 111-123.
Ito S. 1955. Mycological flora of Japan. Vol. 2. Basidiomycetes no. 4. Yokendo Press, Tokyo. 450 p.
Kang HW, Park DS, Park YJ, Lee BM, Cho SM, Kim KT, Seo GS, Go, SJ. 2002. PCR based detection of Phellinus linteus using specific primers generated from universal race primer (URP) derived PCR polymorphic band. Mycobiology 30: 202-207. http://dx.doi.org/10.4489/ MYCO.2002.30.4.202
Karaman M, Jovin E, Malbasa R, Matavuly M, Popovic M. 2010. Medicinal and edible lignicolous fungi as natural sources of antioxidative and antibacterial agents. Phytotherapy Research 24(10): 1473-1481. http://dx.doi.org/10.1002/ptr.2969
Larsen MJ, Cobb-Poulle LA. 1990. Phellinus (Hymenochaetaceae): a survey of the world taxa. Fungiflora, Oslo. 206 p.
Larsson K-H, Parmasto E, Fischer M, Langer E, Nakasone KK, Redhead SA. 2006. Hymenochaetales: a molecular phylogeny for the hymenochaetoid clade. Mycologia 98(6): 926-936. http://dx.doi.org/10.3852/mycologia.98.6.926
Lim YW, Lee JS, Jung HS. 2003. Type studies on Phellinus baumii and Phellinus linteus. Mycotaxon 85: 201-210.
Parmasto E, Parmasto I. 2001. Phellinus baumii and related species of the Ph. linteus group (Hymenochaetaceae, Hymenomycetes). Folia Cryptogamica Estonica 38: 53-61.
Shon M-Y, Kim T-H, Sung N-J. 2003. Antioxidants and free radical scavenging activity of Phellinus baumii (Phellinus of Hymenochaetaceae) extracts. Food Chemistry 82: 593-597.
Srivilai P, Chaiseana W, Loutchanwoot P, Dornbundit P. 2013. Comparison of differences between the wood degradation by monokaryons (n) and dikaryons (2n) of white rot fungus (Cambodian Phellinus linteus). Journal of Biological Sciences 13: 131-138. http://dx.doi.org/10.3923/jbs.2013.131.138
Tian X-M, Yu H-Y, Zhou L-W, Decock C, Vlasak J, Dai Y-C. 2013. Phylogeny and taxonomy of the I. linteus complex. Fungal Diversity 58: 159-169. http://dx.doi.org/10.1007/s13225-012-0202-9
Inonotus baumii in Thailand ... 367
Tura D, Zmitrovich IV, Wasser SP, Spirin WA, Nevo E. 2011. Biodiversity of the Hetero- basidiomycetes and non-gilled Hymenomycetes (former Aphyllophorales) of Israel. Gantner Verlag KG, Ruggell. 566 p.
Venturella G, Altobelli E, Bernicchia A, Di Piazza S$, Donnini D, Gargano ML, Gorjon SP, Granito VM, Lantieri A, Lunghini D, Montemartini A, Padovan F, Pavarino M, Pecoraro L, Perini C, Rana L, Ripa C, Salerni E, Savino E, Tomei PE, Vizzini A, Zambonelli A, Zotti M. 2011. Fungal biodiversity and in situ conservation in Italy. Plant Biosystems 145(4): 950-957. http://dx.doi.org/10.1080/11263504.2011.633115
Ying JZ, Mao XL, Ma QM, Zong SC, Wen HA. 1987. Illustrations of Chinese medicinal fungi. Beijing: Science Press (in Chinese).
Wasser SP. 2013. Modern view on current status, future trends, and unsolved problems in studies of medicinal mushrooms. 17-32, in: Proceedings of the 7 International Medicinal Mushroom Conference, IMMC7, Beijing, China.
Wu S-H, Dai Y-C, Hattori T, Yu T-W, Wang D-M, Parmasto E, Chang H-Y, Shih S-Y. 2012. Species clarification for the medicinally valuable “sanghuang” mushroom. Botanical Studies 53: 135-149.
ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MYCOTAXON
http://dx.doi.org/10.5248/130.369 Volume 130, pp. 369-397 April-June 2015
Taxonomic studies in Chrysoderma, Corneromyces, Dendrophysellum, Hyphoradulum, and Mycobonia
KAREN K. NAKASONE
Center for Forest Mycology Research, U.S. Forest Service One Gifford Pinchot Drive, Madison, WI 53726 USA
*CORRESPONDENCE TO: knakasone@fs.fed.us
ABSTRACT — Eight poorly known or unusual crustose and pileate basidiomycete species were studied. These included the type specimens of three monotypic genera: Chrysoderma alboluteum from Réunion is conspecific with Cerocorticium molle; Dendrophysellum amurense from the Russian Far East is a species of Vararia; and Hyphoradulum conspicuum belongs in Pseudolagarobasidium and is the first representative of the genus from Europe. Corticium murrillii, from Mexico, is congeneric with Corneromyces kinabalui. New combinations Vararia amurensis, Pseudolagarobasidium conspicuum, and Corneromyces murrillii are proposed. Mycobonia flava and M. brunneoleuca are macroscopically similar species with diagnostically distinct basidiospore shape and size. Mycobonia disciformis is accepted in Mycothele, and Mycobonia winkleri represents a species of unknown affinities. Recent molecular phylogenetic studies indicate that Mycobonia is embedded in Polyporus sensu stricto and is a synonym of Polyporus. Transferring M. brunneoleuca and M. flava to Polyporus requires the creation of the replacement names, P. polyacanthophorus and P. epitheloides.
Key worps — Amylocorticiales, cyanophilous basidiospores, Epithele, Favolus curtipes, Polyporales
Introduction
Tremendous advances in the systematics of basidiomycetes have been made in the last twenty years since the advent of molecular phylogenetics. Nevertheless, morphological studies are still essential to advance this discipline. In this paper, eight species of poorly known crustose or pileate basidiomycetes are described and illustrated. Type specimens of Chrysoderma alboluteum, Corticium murrillii, _Dendrophysellum amurense, and Hyphoradulum conspicuum were examined. In addition, the genus Mycobonia (Polyporales) is revisited. Species of Mycobonia produce stipitate, pileate basidiomes with hyphal pegs penetrating a smooth hymenial surface. Four species have been
370 ... Nakasone
included in Mycobonia — M. brunneoleuca, M. disciformis, M. flava, and M. winkleri (Julich 1976). Mycobonia brunneoleuca is considered a synonym of M. flava by some authors, and M. disciformis was transferred to the monotypic genus Mycothele by Jilich (1976). Little is known about M. winkleri, which has not been studied since it was first described in 1911. In this paper, Mycobonia brunneoleuca and M. flava are accepted as distinct species and transferred to Polyporus Adans. The literature concerning M. flava and M. brunneoleuca is critically examined. All the taxa are described and illustrated.
Materials & methods
Thin, freehand sections from basidiomes were mounted in Melzer’s reagent (Kirk et al. 2008) or in 1% (weight/volume) aqueous phloxine and 2% (w/v) aqueous potassium hydroxide. Cyanophily of basidiospore and hyphal walls was observed in 0.1% cotton blue in 60% lactic acid (Kotlaba & Pouzar 1964; Singer 1986). Basidiome sections were mounted in freshly prepared sulfovanillin solution (1 g vanillin, 3 ml distilled water, 8 ml concentrated sulfuric acid); a positive reaction was recorded if contents of the cystidium turned black. Drawings were made with a camera lucida attachment on an Olympus BH2 compound microscope. Q values were obtained from dividing average basidiospore length by width (Kirk et al. 2008). Basidiospores are sometimes scarce in specimens, thus Q values based on less than 30 basidiospores are approximate and indicated with an asterisk (*). Color codes are from Kornerup & Wanscher (1978) except that capitalized color names follow Ridgway (1912). Herbarium code designations follow Index Herbariorum (Thiers 2014). Accepted species names are in boldface.
Taxonomy & discussion
Chrysoderma alboluteum Boidin & Gilles, Cryptog. Mycol. 12: 127. 1991. FIGS 1-2, 10-11
= Corticium molle Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10: 336. 1868 [“1869”].
= Cerocorticium molle (Berk. & M.A. Curtis) Jiilich, Persoonia 8: 219. 1975. = Corticium armeniacum Sacc., Syll. fung. 6: 637. 1888.
= Terana armeniaca (Sacc.) Kuntze, Revis. gen. pl. 2: 872. 1891. = Corticium ceraceum Berk. & Ravenel, in Massee, J. Linn. Soc., Bot. 27: 150. 1890 [“1891”]. = Cerocorticium bogoriense Henn. & E. Nyman, Monsunia 1: 139. 1900 [“1899”]. = Cerocorticium tjibodense Henn., Monsunia 1: 139. 1900 [“1899”]. = Corticium aureolum Bres., Ann. Mycol. 9: 272. 1911.
BASIDIOME resupinate, widely effused, orbicular at first then confluent, <10 x 3 cm, rarely <1 m long, often breaking up into smaller pieces when dried, often slightly detached from substrate, <650 um thick, ceraceous to corneous, smooth to slightly warted, white at first, then dull yellow, light orange (5A4), brownish orange (6C7), brown [7D(6-7)], Warm Buff, Russet, Tawny, Hazel, or Snuff Brown, sometimes slightly cracked on drying. CONTEXT white to cream- colored. MARGIN distinct, abrupt, edges detached or curling slightly away from
Unusual crust and pileate basidiomycetes ... 371
substrate, thickly fibrillose, or thinning out, concolorous with hymenium, white, or dark yellow.
HYPHAL SYSTEM monomitic with clamped generative hyphae. SuBicuLUM <400 um thick, a moderately dense, non-agglutinated tissue of more or less vertical hyphae; subicular hyphae (2-)3-6 um diam, clamped, moderately branched, walls hyaline, thin to 1.5 um thick, smooth, acyanophilous. SUBHYMENIUM <75 um thick, a dense, compact but non-agglutinated tissue of vertically arranged hyphae; subhymenial hyphae 4-5.5 um diam, clamped, frequently branched, walls hyaline, thin, smooth. HyMENtum <100 um thick, a dense, non-agglutinated palisade of hyphidia and basidia sometimes embedded in mucilaginous material HypuHip1A sometimes absent, when present filamentous with slight constrictions or swellings, sometimes with knobby outgrowths at apex, 35-70 x 2-4 um diam, clamped at base, rarely branched, walls hyaline, thin or sometimes basally thickened, smooth. Basip1a narrowly clavate with a long stalk, (40-)65-85(-100) x 7.5-10(-12) um, clamped at base, often containing oil-like inclusions, walls hyaline, thin to slightly thickened, smooth, 4-sterigmate, sterigmata 7-10 x 1.5-2.2 um. BASIDIOSPORES scarce to numerous, narrowly cylindrical to cylindrical, (13-)15-20(-23) x 4.5-8(-9) um, average of five specimens 16.4-19.4 x 5.7-7.9, Q = (2.1-)2.6-3.1, often containing oil-like, cyanophilous materials, walls hyaline, thin, smooth, acyanophilous, not reacting in Melzer’s reagent.
HABITAT & DISTRIBUTION — On bark and wood of various dead angiosperms, occasionally on or under bark of living trees, throughout tropical and subtropical areas of North America, South America, Asia, and Africa.
TYPE SPECIMENS EXAMINED — REUNION. Anse des Cascades, sur branche tombée au bordure de leau, 6 avril 1990, J Boidin (LY 14412, holotype of C. alboluteum). UNITED
STATES. SoutH Caro ina: ad corticeum ramulosque Corni, 1855, H Ravenel — Fungi Caroliniana, Fasc. III no. 29 (BPI 280736, 280735, syntypes of C. ceraceum).
ADDITIONAL SPECIMENS EXAMINED — BRAZIL. Parana: Curitiba, Capao da Imbuia, on decayed, dicot trunk, 16 Nov 1992, AAR de Meijer 2367 (CFMR). SAo LEOPOLDO: in ligno frondoso, 1907, Rick - Fungi Austro-Americani no. 240, as Aleurodiscus alboroseus Bres. (BPI 280723, as C. ceraceum). MEXICO. Orizaba, Rincon Grande and El Barrio, 4000 ft, on decorticate branch, 10-14 Jan 1910, WA & EL Murrill 54615 (BPI 280733, as C. ceraceum). PUERTO RICO. LuquILLo Municipio: Sabana barrio, ridge above chicken farm along Rio Sabana, on decorticated hardwood, 30 Jun 1996, KK Nakasone (CFMR FP 150011). REUNION. St. Gilles I, en partie vivante Casuarina equisetifolia L. (bark), 26 avril 1985, J Boidin (LY 11356, as C. alboluteum); St. Gilles II, ravine, sur branche au sol, 26 avril 1985, J Boidin (LY 11373, as C. alboluteum). SOUTH AFRICA. No location, on bark, no date, P van der Bijl 13 (BPI 280731). UNITED STATES. FLoripa: Highlands Hammock, on bark of Quercus sp., 8 Feb 1937, CL Shear 330 (BPI 28038). Loutsi1Ana: Lafayette Parish, on rotten Cornus florida L., 7 May 1887, AB Langlois 1467 (BPI 289279, as C. armeniacum); St. Martinsville, on bark, 21 Apr 1897, AB Langlois 2389 (BPI 330776, as C. ceraceum). SOUTH CAROLINA: on Cornus sp., no date, H Ravenel, Ellis — North American Fungi no. 607 (BPI 280278).
372 ... Nakasone
Bd ?
: ie 3) ; F : >
Ki be z Fics 1-9. Cerocorticium molle (FP 150011). 1. Basidiome. Chrysoderma alboluteum (LY 11356). 2. Basidiome. Corticium murrillii (isotype HUH 00290581). 3. Basidiome surface; 4. close-up of
basidiome surface; 5. subicular hyphae in Melzer’s reagent; 6. basidiospore with cyanophilous walls; 7. basidiospore with bluish black, amyloid walls in Melzer’s reagent. Dendrophysellum amurense
(holotype TAAM 015561). 8. Basidiome; 9. close-up of hymenial surface. Scale bars: 1, 8 = 10 mm; 2= 2 mm; 3, 4,9 = 1 mm; 5, 6 = 10 um.
Unusual crust and pileate basidiomycetes ... 373
FiGs 10-11. Corticium ceraceum (syntype BPI 0280736). 10. A. Basidiospores; B. basidia; C. hyphidia. Cerocorticium molle. 11. Basidiospores: A. from BPI 0280731; B. from BPI 0280733.
DESCRIPTIONS & ILLUSTRATIONS — Boidin & Gilles (1991, as C. alboluteum), Burt (1926:
216, as C. ceraceum), Hjortstam (1983), Hohnel & Litschauer (1907, as C. ceraceum),
Julich (1975), Maekawa et al. (2003), Nakasone (2008, as C. aureolum), Talbot (1951, as
C. armeniacum), Trierveiler-Pereira et al. (2009), Wu & Chen (1990). COMMENTS — Cerocorticium molle is characterized by yellow, orange, or brown, ceraceous to corneous basidiomes, simple hyphidia, large basidia, and large, cylindrical to narrowly cylindrical basidiospores. The basidiospores are variable in shape and size, sometimes within a single specimen. Hohnel’s (1910) and Julich’s (1975) synonymies of Cerocorticium bogoriense and C. tjibodense, both from Java, with C. molle are accepted.
374 ... Nakasone
There is no doubt that C. alboluteum is conspecific with C. molle as suggested by Hjortstam & Larsson (1995). Although hyphidia were not observed in specimens of C. alboluteum examined, basidiomes and basidiospores are typical for the species. Boidin & Gilles (1991) noted that in C. alboluteum the basidiospore walls were pale yellow, producing a mass basidiospore color of yellowish orange, and calculated the average basidiospore size of three specimens as 12.8-14.6 x 4.4—5.1 um, Q = 2.8-3.
Corticium murrillii Burt, Ann. Missouri Bot. Gard. 13: 289. 1926. FIGS 3-7, 12 = Corneromyces murrillii (Burt) Nakasone, comb. nov. MycoBank MB812353
BASIDIOME resupinate, widely effused, <9 x 3.5 cm, loosely attached, <900 um thick (Burt 1926), soft, fragile, membranous, abhymenial surface fibrous, felty, brown (7E8), turning black in KOH then fading, sometimes hyphal strands found under basidiome or in the substrate. HYMENIAL SURFACE smooth, even, finely farinaceous, orange white to pale orange [5A(2-3)], greyish orange [5B(4-5)], brownish orange [5C4-6)], yellowish brown (5D6), Cream-Buff, or Chamois, black in KOH then fading to light brown; cracks occasional. CONTEXT with a thin, dark brown layer next to substrate and a thicker, cream-colored upper layer. MARGIN thinning out, fibrillose, loosely attached or detached, concolorous with hymenium or developing a dark brown edge, sometimes poorly developed hyphal strands present.
HYPHAL SYSTEM monomitic with clamped generative hyphae. SusicuLuM <750 um thick, a non-agglutinated tissue composed of brown, rough-walled subicular hyphae loosely arranged parallel to substrate, then hyphae turning into hymenium, becoming hyaline; subicular hyphae of two types: (a) next to substrate 3-5.5 um diam, clamped, moderately branched, walls yellow to brown, <1.1 um thick, smooth or ornamented with tiny tubercles; (b) in upper subiculum 3-6 um diam, clamped, moderately branched, walls hyaline, thin, smooth. SUBHYMENIUM a non-agglutinated, scarcely differentiated tissue of upright hyphae; subhymenial hyphae 2.2-5 um diam, clamped, moderately branched, walls hyaline, thin, smooth. HyMENIuM a palisade of hyphidia and basidia. Hypuip1a scarce, filamentous, obclavate, or narrowly clavate, (25-) 35-65 x (3-)4-6.5(-10) um, clamped atbase, walls hyaline, thin, smooth. BAsIDIA collapsing soon after ejecting spores, obclavate at first then clavate to cylindric, usually with a distinct stalk, (35-)40-90(-110) x 10-14 um, clamped at base, walls hyaline, thin or slightly thickened at base, smooth, 4-sterigmate, sterigmata
Fics 12-13. Corticium murrillii (holotype BPI 0282179). 12. A. Basidiospores; B. basidia; C. hyphidia. Dendrophysellum amurense (holotype TAAM 015561). 13. A. Basidiospores; B. basidia; C. gloeocystidia; D. subicular hypha; E. skeletal-like hyphae; EF. dichohyphidia.
Unusual crust and pileate basidiomycetes ... 375
eS 12
|
376 ... Nakasone
<17 x 3.5 um. Basipiospores abundant, narrowly cylindric to allantoid, (25-)28-40(-43) x (7.2-)8-11(-11.5) um, averages of three specimens 28.9-29.6 x 8.7-10.4 um, Q = 2.8-3.4, average of isotype at FH 34.4 + 4.2 x 9.9 + 0.9 um, Q = 3.5 + 0.4, walls hyaline, occasionally pale yellow, thin to 1.3 um thick, smooth, cyanophilous, dark blue-black in Melzer’s reagent.
HABITAT & DISTRIBUTION — On bark and wood of dead angiospermous branches and logs in subtropical and tropical Central and South America.
TYPE SPECIMENS EXAMINED — MEXICO. Jalapa, 5000 ft, on bark of decaying log, 12-20 Dec 1909, WA & EL Murrill 182 (BPI 0282179, holotype; NY 00776553, NY 00562612, and FH HUH-00290581, isotypes).
SPECIMENS EXAMINED — COLOMBIA. Magdalena, Sierra Nevada de Santa Marta, Reserva Forestal San Lorenzo, on wood, 17-19 June 1978, L Ryvarden 16070 (O, as “C. kinabalui”). ECUADOR. SucumBios PROVINCE: Reserva Natural de Cuyabeno, 300 m.a.s.l., on hanging branch, 28 June-15 July 1993, M Nunez 306 (O F-902593, as “C. kinabalui” ).
DESCRIPTIONS & ILLUSTRATIONS — Burt (1926), Liberta (1969).
COMMENTS — Corneromyces murrillii is an uncommon and unusual species characterized by a soft fragile basidiome, hyphidia, large basidia, and large narrowly cylindric to allantoid basidiospores with amyloid and cyanophilous walls. There was a wide range in basidium and basidiospore size among the specimens examined. Corneromyces murrillii is similar to C. kinabalui Ginns from Borneo, which has brown aculeate basidiomes, smooth brown subicular hyphae, and brown basidiospores (Ginns 1976). Basidiospore sizes in the two species are similar. The average basidiospore size of the isotype of C. kinabalui at BPI, BPI 0290583, is 27 + 2.9 x 9.1 + 1.1 um, Q=3 + 0.3.
One isotype of C. murrillii (NY 00776553) is a large collection that is intermixed with small basidiome fragments of Rhizochaete radicata (Henn.) Gresl. et al. Hjortstam & Ryvarden (2001, 2008) erroneously reported C. kinabalui from Columbia and Ecuador; two of the voucher specimens were examined and identified as C. murrillii.
The affinities of Corneromyces Ginns are not known. Ginns (1976) considered creating a new family for Corneromyces but placed it instead in Coniophoraceae Ulbr. Citing the strongly amyloid, brown, thick-walled basidiospores, Julich (1979) proposed the family Corneromycetaceae Jiilich. Recent molecular phylogenetic studies have identified a new order, Amylocorticiales K.H. Larss. et al. This order is characterized by varied basidiome habit and hymenophore configuration, a monomitic nodose-septate hyphal system, and smooth thin- or thick-walled basidiospores that are amyloid in most species (Binder et al. 2010). Corneromyces may belong in the Amylocorticiales, which is sister to the Agaricales Underw., instead of with Coniophora DC. and allies in the Boletales E.-J. Gilbert.
Unusual crust and pileate basidiomycetes ... 377
Dendrophysellum amurense Parmasto, Consp. syst. cortic.: 206. 1968.
FIGs 8-9, 13-15
= Vararia amurensis (Parmasto) Nakasone, comb. nov. MycoBAnk MB812355
BASIDIOMES resupinate, effused, beginning as circular to irregular colonies, then coalescing, <5 x 20 mm, adnate, thin, <150 um thick, subceraceous, smooth to finely granulose over the irregular contours of the substrate, greyish yellow [4(B-C)(3-4)], greyish orange [5B(3-4)], to brownish orange (5C3), rimose, exposing white, felty to cottony context; cottony white mycelial tissue developed in substrate and occasionally beneath basidiome. MARGIN adnate, thinning out, white, farinaceous.
HYPHAL SYSTEM dimitic with clamped generative hyphae and aseptate dichohyphidia. MycrELiaL Tissue cottony, white, composed of skeletal-like hyphae; hyphae 2.2-3.5 um diam, occasionally clamped or appearing aseptate, sparsely branched, straight, non-staining, walls hyaline, thin to thick, smooth. SUBICULUM <100 um thick, basal layer next to substrate a compact, agglutinated tissue of short-celled hyphae arranged more or less parallel to substrate, then hyphae becoming upright, mostly indistinct and degraded although with a few, intact, phloxine-stained hyphae; subicular hyphae from basal layer 3-5 um diam, clamped, frequently branched, walls hyaline, thin, smooth; hyphae of upper subiculum 1.5-2.2 um diam, clamped, moderately branched, walls hyaline, thin, smooth. SUBHYMENIUM AND HYMENIUM indistinct, composed of dichohyphidia, gloeocystidia, and basidia intermixed with abundant, coarse, hyaline crystals. DICHOHYPHIDIA scattered in hymenium, not enclosed, dendriform, <20 x 20 um, with a main stalk, 2-2.5 um diam, and multiple, short branches, walls hyaline, slightly thickened, smooth, cyanophilous, dextrinoid. GLOEOCYSTIDIA scarce, inconspicuous, cylindrical to narrowly clavate, occasionally papillate, 23-27 x 3-4 um, clamped at base, protruding <15 um, containing oil-like material, negative in sulfovanillin, walls hyaline, thin, smooth. Basrip1a suburniform at first, then narrowly cylindrical, flexuous, about 26 x 5 um, clamped at base, protruding <10 um, 4-sterigmate, walls hyaline, thin, smooth. BasipiosporREs numerous, agglutinated, often collapsed, subfusiform to pip-shaped (in face view), 6-8(-9.3) x 3-3.7(-4) um, x = 7.1 + 0.7 x 3.3 + 0.3 um, Q = 2.1 + 0.3, walls hyaline, thin, smooth, acyanophilous, faintly amyloid.
HABITAT & DISTRIBUTION — On bark of dead Picea; known from the type locality in northeastern Asia.
TYPE SPECIMEN EXAMINED — RUSSIA. KHABAROVSK REGION: Selikhino, Kabansopka, on bark of fallen trunk of Picea jezoensis (Siebold & Zucc.) Carriere, 18 Aug 1961, E Parmasto (TAAM 015561, holotype).
378 ... Nakasone
COMMENTS — Vararia amurensis is characterized by clamped generative hyphae, dextrinoid and cyanophilous dendriform dichohyphidia, gloeocystidia, suburniform basidia, and subfusiform amyloid basidiospores. A typical catahymenium was not observed. Mature basidia are rare, and gloeocystidia are inconspicuous and easily overlooked. It is morphologically similar to V. mediospora var. makokouensis Boidin et al. from Gabon, which has basidiospores of the same size and shape but lacks clamp connections (Boidin et al. 1980). Parmasto (1968: 146) placed Dendrophysellum Parmasto in Corticiaceae subfam. Aleurodiscoideae Parmasto, but it is morphologically similar to taxa in Vararia sect. Fusamyspora Boidin & Lanq. that have dextrinoid and cyanophilous dendrohyphidia, smooth or slightly ornamented amyloid basidiospores, and sulfovanillin-negative gloeocystidia (Boidin & Lanquetin 1975). Thus, the monotypic genus Dendrophysellum is placed in synonymy under Vararia and D. amurense is transferred to Vararia.
Hyphoradulum conspicuum Pouzar, Ceska Mykol. 41: 26. 1987. FIGS 16-17, 25 = Pseudolagarobasidium conspicuum (Pouzar) Nakasone, comb. nov. MycoBank MB812356
BASIDIOME resupinate, widely effused, soft, fragile, cretaceous, spinose, yellowish white (4A2), greyish yellow (4B3), or greyish orange (5B3), no color change with KOH; cracks absent. HyMENIAL SURFACE composed of fragile, brittle, soft, cretaceous or chalky aculei, <2 aculei per mm, <3.5 mm long, terete to conical then gradually tapering to an acute or rounded apex, single at first then fused at base or along entire length, occasionally reticulate, smooth or studded with tiny, round tubercles, apices concolorous with base of aculei or pale yellow, often broken off. MARGIN cream white, byssoid, fibrillose. HyPHAL SYSTEM monomitic with clamped generative hyphae. ACULEI composed of non-agglutinated tramal hyphae arranged in a fascicle with embedded tramal cystidia curving into hymenium, at aculeal apex terminal hyphae undifferentiated, smooth; tramal hyphae (1.5-)2-3(-4) um diam, clamped, sparingly branched, even, walls distinct, hyaline, thin, smooth, sometimes weakly cyanophilous. SusrcuLum 300-700 um thick, a non-
Fics 14-24. Dendrophysellum amurense (holotype TAAM 015561). 14. Dichohyphidium with cyanophilous walls; 15. dichohyphidium with dextrinoid walls in Melzer’s reagent. Hyphoradulum conspicuum (isotype PRM 834887). 16. Aculei; 17. close-up of aculei. Polyporus polyacanthophorus (BPI 0261328). 18. Hyphal pegs from hymenial surface. Favolus curtipes (isotype HUH 00290582). 19. Close-up of poroid surface; 20. poroid surface of basidiome. Mycothele disciformis (isotype BPI 0261300). 21. Close-up of basidiomes. Polyporus epitheloides (BPI 0261319). 22. Close-up of apically encrusted hyphal pegs. Mycobonia winkleri (holotype F-15807). 23. Cross-section of “hyphal peg” or bulbil; 24. close-up of hymenial surface with “hyphal pegs” or bulbils. Scale bars: 14, 15 = 10 um; 16, 18, 19 = 2 mm; 17, 21, 24 = 1 mm; 20 = 20 mm; 22 = 0.5 mm; 23 = 50 um.
Unusual crust and pileate basidiomycetes ... 379
380 ... Nakasone
agglutinated tissue; subicular hyphae 1-5 um diam, clamped, walls hyaline, thin, smooth. SuBHYMENIUM <40 um thick, composed of short-celled hyphae in a dense non-agglutinated tissue; subhymenial hyphae 2-3 um diam, clamped, frequently branched, walls hyaline, thin, smooth. HYMENIUM <35 um thick, a dense palisade of hyphidia, cystidia, and basidia. HypHip1a scarce, inconspicuous, filamentous to subulate, 15-23 x 1.5-3 um, clamped at base, walls hyaline, thin, smooth. Cystip1a of two types: (a) arising from aculei trama, subiculum, and subhymenium, abundant, embedded, broadly cylindrical, clavate, or obclavate, often strangulated or moniliform, sometimes with a lateral lobe, stalked, apex obtuse, occasionally branched, (27—)40-80 x 5-10 um, tapering to 1.5-3 um diam at base, with a basal clamp, often with honey yellow, refractive contents, walls hyaline, thin, smooth, negative in sulfovanillin; (b) arising from hymenium, rare, clavate to cylindrical but sometimes with a small, lateral beak, 11-15 x 4-5 um, with a basal clamp, walls hyaline, thin, smooth. Basip1a clavate, often with a distinct stalk, (16-)20-35 x 5-6 um, clamped at base, walls hyaline, thin, smooth, 4-sterigmate. BASIDIOSPORES ellipsoid with a small apiculus, (4.8-)5-5.8(-6) x 3.5-4.4 um, average of isotype 5.4+0.4 x 4+0.2 um, Q=1.3 + 0.1, filled with numerous oil-like globules, walls hyaline, slightly thickened, smooth, cyanophilous, not reacting in Melzer’s reagent. HABITAT & DISTRIBUTION — Saprophytic on wood and bark of dead Cornus;
known from the type location, Bohemia.
TYPE SPECIMENS EXAMINED — CZECH REPUBLIC. Bouemica: “Velka hora” hill near
Karl&Stejn, on base of dead Cornus mas L., 2 Oct 1981, Z Pouzar (PRM 834886, holotype;
PRM 834887, isotype). ComMENTS — Pseudolagarobasidium conspicuum is characterized by soft, brittle, cretaceous or chalky aculei, strangulated or monilioid cystidia with honey-yellow contents, and ellipsoid basidiospores with slightly thickened, cyanophilous walls. Microbinding hyphae were not observed. Pouzar (1987) noted that old collapsed basidiospores have dextrinoid walls, which was not observed in this study. This is the first species of Pseudolagarobasidium J.C. Jang & T. Chen reported from Europe. The description above is based partially on Pouzar’s (1987) observations. Hyphoradulum conspicuum is transferred to Pseudolagarobasidium, and Hyphoradulum, a monotypic genus, becomes a synonym of Pseudolagarobasidium.
This species is most similar to P. pronum (Berk. & Broome) Nakasone
& D.L. Lindner with respect to color and texture of the basidiome.
Fics 25-26. Hyphoradulum conspicuum (isotype PRM 834887). 25. A. Basidiospores; B. basidia; C. hymenial cystidia; D. tramal cystidia; E. section through hymenium with tramal cystidia and immature basidia. Polyporus polyacanthophorus (TENN 57579). 26. A. Basidiospores; B. basidia.
382 ... Nakasone
Pseudolagarobasidium pronum develops microbinding hyphae and has slightly smaller basidiospores (4-5.5 x 3-3.7(-4.3) um) with weakly cyanophilous walls, whereas P. conspicuum lacks microbinding hyphae and has distinctly cyanophilous basidiospore walls. In addition, cystidia in P pronum do not contain refractive, honey-yellow contents. Pseudolagarobasidium conspicuum is known from Europe, whereas P. pronum is reported from Asia, Australia, Sierra Leone (Nakasone & Lindner 2012), and a first report from Jamaica (Runaway Bay, on Calliandra sp., 12 Feb 2006, F Dammrich 8139 at CFMR). See Nakasone & Lindner (2012) for a description of P pronum and other species of Pseudolagarobasidium.
The genus Mycobonia
Mycobonia Pat. was erected with the generic type Hydnum flavum Berk. (Patouillard 1894). Mycobonia was substituted for the illegitimate Bonia Pat., which had Bonia papyrina Pat. as its generic type (Patouillard 1892). These two generic names are not homotypic; thus, Mycobonia was published as a new genus rather than a replacement name. For a detailed nomenclatural history of Mycobonia, see Jiilich (1976) and Martin (1939). One important feature of Mycobonia is the dense stand of hyphal pegs penetrating the smooth hymenial surface. Hyphal pegs are not restricted to Mycobonia but are produced in a number of basidiomycete species. For a key to basidiomycete taxa with hyphal pegs see Nakasone (2013). Mycobonia is readily distinguished from other taxa with hyphal pegs and non-septate basidia by its substipitate and pileate basidiomes.
The classification of Mycobonia is controversial because of its unique combination of morphological features. Donk (1964: 294) placed Mycobonia in the Stereaceae Pilat even as he noted a close relationship to Pseudofavolus Pat. in the Polyporaceae Fr. ex Corda. Jiilich (1982: 186) created Mycoboniaceae Jiilich for Mycobonia, placing the family in the Polyporales. Later, Singer (1986: 171) reduced Mycoboniaceae to a synonym of Polyporaceae. Other researchers also noticed striking morphological similarities between Mycobonia and Polyporus (Corner 1984: 102; Kriiger 2002; Kriiger & Gargas 2010; Ryvarden 1991: 213, 2010: 119) and between Mycobonia and Pseudofavolus (Corner 1984: 36, Kruger 2002, Kriiger & Gargas 2010, Singer 1986: 171). Authors have generally agreed that Pseudofavolus is closely related to Polyporus s.s. (Nunez & Ryvarden 1995: 68, Ryvarden 1991: 213), while some consider Pseudofavolus a synonym of Polyporus (Corner 1984: 35, Kriiger 2002, Kriiger & Gargas 2010).
Molecular phylogenetic studies show that Mycobonia and Pseudofavolus are sister taxa embedded in the core polyporoid clade (Binder et al. 2013; Kruger 2002; Kriiger & Gargas 2004, 2010; Sotome et al. 2008), confirming
Unusual crust and pileate basidiomycetes ... 383
morphological observations. Based on phylogenetic studies and morphological similarities, Kriiger (2002) and Kriiger & Gargas (2010) reduced M. flava to a subspecies of Polyporus curtipes (Berk. & M.A. Curtis) Ryvarden. While agreeing with these authors that the molecular and morphological evidence overwhelmingly indicates that Mycobonia is a synonym of Polyporus, significant differences in ITS sequences and basidiospore shape and size argue for the recognition of M. flava and M. brunneoleuca as distinct species. Transfer of these taxa requires replacement names because both species epithets are preoccupied in Polyporus.
Mycobonia brunneoleuca (Berk. & M.A. Curtis) Pat., Bull. Soc. Mycol. France 16: 181. 1901 (1900). FIGs 18, 26-27 = Hydnum brunneoleucum Berk. & M.A. Curtis, Trans. Linn. Soc. London
22: 129. 1857, non Polyporus brunneoleucus Berk. 1846.
= Polyporus polyacanthophorus Nakasone, nom. nov. MycoBank MB812357
ErymMo.Loecy: from the Greek, poly- (many) + acantha (spine) + -phorus (bearing),
referring to the numerous hyphal pegs in the hymenial surface. BASIDIOME pileate, sessile or substipitate, coriaceous, flabellate, reniform. PitEus galeaeform or helmet-shaped, <10 x 8 cm; upper surface smooth, becoming radially rugulose, bright yellow when fresh, drying to dull greyish orange (5B6), brown (7D7), reddish brown (9E6), Vinaceous-Russet, Pecan Brown, or Kaiser Brown. Stipe reduced, usually black, 3.5-9 mm diam, attached to substrate by a thin, circular pad, <8-13 mm diam. CONTEXT <4 mm thick at base, thinning to 0.3 mm at pileus edge, pale orange (5A3), coriaceous. HYMENIAL SURFACE appearing smooth but densely covered with minute, conical hyphal pegs, <330 x 40-80 um, 9-12 pegs per mm, becoming fimbriate at apex, purplish tan when fresh, drying to greyish orange (5B3), brownish orange to light brown [6(C-D)5], or Cinnamon Buff.
HYPHAL SYSTEM dimitic with clamped generative and aseptate skeleto- binding hyphae. HypHaL PEGs a dense fascicle of non-agglutinated, sparsely branched skeleto-binding hyphae, <330 um long, originating 100-150 um below hymenium, projecting <180 tm, encrusted at apex with coarse, irregular, hyaline crystals; hyphae rigid, tapering to acute apex, <3 um, walls hyaline to light brown, slightly thick, smooth, cyanophilous, weakly dextrinoid. CONTEXT a densely interwoven, non-agglutinated tissue composed mostly of skeleto- binding hyphae and some generative subicular hyphae; subicular hyphae 3-6.5 um diam, clamped, strangulated, staining in phloxine, walls hyaline, thin, smooth; skeleto-binding hyphae (0.5—)1.5-5.5 um diam, occasionally inflated <12 um diam, then tapering to tips, with a narrow lumen, aseptate, rarely to
384 ... Nakasone
| Ke _
Oy ee. C (DD 0 10 20 um
Fic. 27. Polyporus polyacanthophorus (NY 00536422). A. Basidiospores; B. thin-walled hyphidia; C. cystidia; D. hyphidia from hyphal pegs; E. skeleto-binding hyphae from context of NY 00543220.
extensively branched, walls hyaline, <1.5 um thick, smooth, acyanophilous, not reacting in Melzer’s reagent. SUBHYMENIUM a dense, thickening, non- agglutinated tissue of upright, short-celled hyphae; subhymenial hyphae 3.5-4.5um, clamped, frequentlybranched, wallshyaline, thin,smooth. HyYMENIUM a dense palisade of hyphidia, cystidia, and basidia. HypHip1A or two types: (a) inconspicuous, filamentous, often with short branches at apex, (23-)32-60
Unusual crust and pileate basidiomycetes ... 385
(-80) x 2-4 um, clamped at base, walls hyaline, thin, smooth; (b) numerous, acicular or narrowly cylindrical then tapering to a subacute or acute tips, 1.5-4 um diam, with a narrow lumen, aseptate, sparsely branched, walls hyaline, slightly thick to thick, smooth, acyanophilous, not reacting in Melzer’s reagent (may be interpreted as terminal ends of skeleto-binding hyphae). Cystip1a of two types: (a) rare, inconspicuous, embedded, subfusiform to cylindrical, tapering slightly to apex, 45-55 x 6-7.5 um, clamped at base, walls hyaline, thin, smooth; (b) rare, obclavate, an enlarged bulbous base tapering to apex, 25-45 x (6-)10-15 um, aseptate, walls hyaline, <1 um thick, smooth, acyanophilous, not reacting in Melzer’s reagent. Basrp1A scarce, clavate with a stalk, <80 x 10-16 um, clamped at base, walls hyaline, thin, smooth, with 4-sterigmata. BASIDIOSPORES cylindrical to narrowly ellipsoid or broadly subfusiform with a small, distinct, hyaline apiculus, (13-)16-24(-26.5) x (7.2-)8-11(-12.3) um, average of three specimens 17.3-21 x 9.4-10.1 um, Q = 1.9-2.2, containing oil-like material, walls hyaline to light brown, thin to 0.7 um thick, smooth, acyanophilous or weakly cyanophilous, not reacting in Melzer’s reagent.
HABITAT & DISTRIBUTION — On logs, branches and twigs of dead hardwoods throughout Central and South America and the Caribbean region.
TYPE SPECIMEN EXAMINED — VENEZUELA. Fendler 129 (FH HUH-00290586, isotype of H. brunneoleucum).
REPRESENTATIVE SPECIMENS EXAMINED — BRAZIL. Prov. SAO PAuLo: S. Framisco dos Campos, Dec 1896, F Noark 333 (S F-177421); Caldas, pnu. Miras Gerais, Regnell (S F-249624). COLUMBIA. Dept. DEL Cauca: Cordillera Central, Hoya del Rio Palo, Ta Tolda, 1944, J Ceratrecas 19440 (BPI 0261323); Munchique, 2700 m elev., on dead wood, 5 May 1968, AL Welden 7318 (NY 00536422). COSTA RICA. PUNTARENAS PROVINCE: Monteverde, on recently cut small trees, 8 Jan 1973, AL Welden 3303 (NY 00543220); Sta. Elena, road to Elena Biological Reserve, on fence post, 17 Mar 1999, RH Petersen 10256 (TENN 57579). SAN JOSE PROVINCE: San Isidro de Coronado, on fallen log, 21 Jun 1972, AL Welden 8338 (NY 00536420); Monté Zurqui, alt. 2000-2500 m., on log, 13 Feb 1926, PC Standley & J Valerio no. 48084 (BPI 261325). Dota, Guadeloupe, Finca Jaular, km 66 on Interamerican Hwy, 1 Jul 1998, RH Petersen 9471 (TENN 56445). HEREDIA PROVINCE: Cerro Central de Zurqui, 1600 m., on dead log, 27 Dec 1929, CW Dodge no. 633 (BPI 261320). HONDURAS. Tegucigalpa, Escuela Agricola Panamericana, 31 Dec 1951, AS Mueller (BPI 261322). PANAMA. Cuiriquf PRovINCE: valley of upper Rio Chiriqui Viejo, 1600-1800 m., 10 Jul 1935, GW Martin 2714 (BPI 261324, as “M. flava”); 6 Jul 1953, GW Martin 2519 (BPI 261238, as “M. flava”); 7 Jul 1935, GW Martin 2631 (NY 00536417, as “M. flava”). PARAGUAY. Asuncion, San Antonii, July 1893, Malme (S F-249621). PUERTO RICO. Guajataca Community Forest, Verada, Nueva Trail, on deciduous wood, 26 Jun 1996, L Ryvarden (CFMR PR 5160).
DESCRIPTIONS & ILLUSTRATIONS — Gerlach & Loguercio-Leite (2011), Martin (1939: 248, figs 13-16, as M. flava), Jiilich (1976, figs 1-2 only), Reid (1976: 191, fig. 2).
ComMENTsS — Polyporus polyacanthophorus is a pileate species characterized by a dense stand of hyphal pegs in the hymenium and large, cylindrical to
386 ... Nakasone
ellipsoid basidiospores. Although considered a synonym of P. epitheloides (as M. flava) by some mycologists, it can be differentiated by its larger basidia and broader basidiospores. In addition, the thin-walled, fusiform to cylindrical cystidia observed in P polyacanthophorus are not present in P. epitheloides. Reid (1976) postulated that P. polyacanthophorus (as M. brunneoleuca) prefers higher elevation habitats than P epitheloides (as M. flava).
The description above is based on Martin (1939), Reid (1976), and personal observations. A replacement name is proposed for H. brunneoleucum because the species epithet is occupied in Polyporus. Although Martin (1939: 247) considered M. brunneoleuca a synonym of M. flava, the specimens he referenced from Panama are redetermined here as P. polyacanthophorus. Similarly, the drawings of M. flava in Julich (1976, Fig. 2) of an Argentinian specimen probably represent P. polyacanthophorus based on the basidiospore shape and size.
Mycobonia disciformis G. Cunn., Trans. Roy. Soc. New Zealand 83: 635. 1956. FIGS 21, 28
= Mycothele disciformis (G. Cunn.) Jiilich, Persoonia 8: 452. 1976.
BASIDIOMES resupinate, scattered to gregarious, disciform, centrally attached to substrate, orbicular to elongate, 1-10 mm diam, <0.5 mm thick, firm, membranous to ceraceous, spinose because of numerous hyphal pegs, exterior surface dark brown. HYMENIAL SURFACE smooth with numerous hyphal pegs penetrating surface, cream, dull sulfur-yellow, or light brown. HyPHAL PEGS cylindrical, tapering to a subacute apex, 20-30 per mm. Marcein distinct, abrupt, free, slightly involute.
HYPHAL SYSTEM monomitic with clamped generative hyphae. HyPHAL PEGS originating deep in subicular trama, a dense, agglutinated fascicle of hyphae with an central column of coarse, hyaline crystals, <1000 x 45 um, protruding <135 um. SuBicuLUM <800 um thick with a basal layer of compact hyphae arranged parallel to substrate, then hyphae becoming upright, forming a dense but non-agglutinated tissue; subicular hyphae 2-3 um diam, clamped, moderately branched, walls hyaline to brown, thin to slightly thick, smooth. HyMENIuM <70 um thick, a dense palisade of hyphidia and basidia. HypHip1a filamentous, occasionally branched at apex, 20-40 x 2-2.5 um, clamped at base, walls hyaline, thin, smooth. Basrp1a fragile, scarce, cylindrical to narrowly clavate, sometimes with a short, stalk-like base, (38-)45-60 x (7—)9-11(-14) um, clamped at base, walls hyaline, thin, smooth, 4-sterigmate, sterigmata 7-10 x 1.5-2 um. BASIDIOSPORES scarce, globose to subglobose or broadly ellipsoid, 8-13.5 x (6.5-)7-9(-10) um, filled with oil-like globules, walls hyaline, thin, smooth, acyanophilous or weakly cyanophilous, not reacting in Melzer’s reagent.
Unusual crust and pileate basidiomycetes ... 387
C B
2 any:
0 10 20 um
Aree eee ee eal
Fic. 28. Mycothele disciformis (isotype BPI 0261300). A. Basidiospores; B. basidia; c. hyphidia.
HABITAT AND DISTRIBUTION — On bark of living trunks of Rhopalostylis sapida H. Wendl. & Drude in New Zealand.
TYPE SPECIMEN EXAMINED — NEW ZEALAND. AucxkLanp: Henderson Valley, 130 m., Sharps Bush, on bark of R. sapida, 5 May 1952, SD Baker PDD 11491 (BPI 261300, isotype). In the original description, the month is mistakenly reported as April (Cunningham 1956: 636).
SPECIMENS EXAMINED — NEW ZEALAND. AUCKLAND: Titirangi, Atkinson Park, on R. sapida, 27 Jun 1953, JM Dingley (PDD 12643); Coromandel Peninsula, Camel’s Back, on R. sapida, Oct 1954, JM Dingley (PDD 14304); Waitakeres, Cascades, on R. sapida, 3 Apr 1954, SD Baker (PDD 14307).
DESCRIPTIONS AND ILLUSTRATIONS — Cunningham (1956, 1963: 173), Julich (1976).
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Comments — Mycothele disciformis is characterized by small fragile disciform basidiomes with prominent hyphal pegs. Epithele ryvardenii Nakasone is an effused species that is reminiscent of M. disciformis because it produces brown- pigmented hyphae and has similarly sized and shaped basidia and basidiospores (Nakasone 2013). However, M. disciformis is found on bark of living nikau, an endemic New Zealand palm, whereas E. ryvardenii is reported from Venezuela on wood. Mycothele disciformis is undoubtedly closely related to Epithele (Pat.) Pat. Whether it should be transferred to Epithele or retained as a monotypic genus may be resolved in the future with molecular evidence.
Mycobonia flava (Sw. : Fr.) Pat., Bull. Soc. Mycol. France 10: 77. 1894. Fics 22, 29-30
= Peziza flava Sw. : Fr., Prod.: 150. 1788, non Polyporus flavus Jungh. 1838.
= Hydnum flavum (Sw. : Fr.) Berk., Ann. Mag. Nat. Hist., 10: 380. 1843 [“1842”].
= Bonia flava (Sw. : Fr.) Henn., Hedwigia 36: 192. 1897, as “(Berk.) Pat.”
= Auricularia flava (Sw. : Fr.) Farl., Bibl. Index N. Amer. Fung.: 307. 1905.
= Grandinioides flava (Sw. : Fr.) Banker, Mem. Torrey Bot. Club 12: 179. 1906.
= Polyporus curtipes subsp. flavus (Sw. : Fr.) D. Kriiger, Cryptog. Mycol. 31: 399. 2010.
= Polyporus epitheloides Nakasone, nom. nov.
MycoBank MB812358 EryMo_oey: refers to similarity with the genus Epithele which is characterized by hyphal pegs.
BASIDIOME pileate, dimidiate, flabelliform or reniform, sessile or substipitate, coriaceous. PILEus applanate, subdiscoid, or shallowly cucullate, <8 x 4 cm; upper surface smooth then becoming radially rugulose, when dried dull, pallid ochraceous, greyish orange (5B5) to brown [6D(7-8)] or purplish chestnut, darkening at edges to brown (6E8). Stipe short, <5 mm long by 3 mm diam, often black, attached to substrate by a thin, circular, mycelial pad, <8 mm diam. ConTEXT 1-6 mm thick at base, thinning to 0.5 mm at pileus edge, light orange (5A4), thin, coriaceous. HYMENIAL SURFACE appearing smooth but densely covered with cylindrical hyphal pegs, <400 x 40-60 um, 5-11 pegs per mm, pale ochraceous when fresh, drying to greyish orange (5B4), Light Ochraceous Buff, purplish buff, or light rusty buff.
HYPHAL SYSTEM dimitic with clamped generative and aseptate skeleto- binding hyphae. HyPHAL psGs a dense fascicle of non-agglutinated, sparsely branched, skeleto-binding hyphae, originating in context, projecting <200 um, heavily encrusted with coarse, irregular, hyaline crystals; hyphae 1-3 um diam, aseptate, rarely branched, walls hyaline, slightly thick to thick, smooth, cyanophilous, weakly dextrinoid. CONTEXT a densely interwoven, Fic. 29. Polyporus epitheloides (NY 00536421). A. Basidiospores; B. papillate hyphidia;
C. thin-walled hyphidia; D. thick-walled hyphidia or skeleto-binding hyphae terminating in hymenium; E. cystidium.
390 ... Nakasone
non-agglutinated tissue composed primarily of skeleto-binding hyphae and indistinct generative hyphae, in pileus cuticle hyphae similar but compacted into a cutis; generative hyphae 2.2-4 um diam, clamped, walls hyaline, thin, smooth; skeleto-binding hyphae 2.2-5 um diam, occasionally inflated <20 um diam, aseptate, rarely to extensively branched, branches often elongate and whip-like, walls hyaline, thick, smooth, acyanophilous, not reacting in Melzer’s reagent. HyMENIUM a dense palisade of hyphidia, cystidia, and basidia. Hypuip1a of two types: (a) inconspicuous, scarce, filamentous, often with a few short branches at apex or papillate, 27-52 x 2-4 um, clamped at base, walls hyaline, thin, smooth; (b) numerous, acicular or narrowly cylindrical then tapering to a subacute to acute apex, 30-60 x 2-4 um, aseptate, walls hyaline, slightly thick, smooth, acyanophilous, not reacting in Melzer’s reagent (may be interpreted as terminal branches of skeleto-binding hyphae). CysTip1A rare, obclavate, an enlarged bulbous base then tapering to apex, 25-45 x (6-)10-15 um, aseptate, walls hyaline, <1 um thick, smooth, acyanophilous, not reacting in Melzer’s reagent. Bastp1a clavate, 26-40 x 6-12 um, clamped at base, walls hyaline, thin, smooth, with 4-sterigmata. BAsip1osPorEs cylindrical to subfusiform, 14-23(-26) x 5-8(-8.7) um, average of three specimens 15.6-22.5 x 5.2-7.4 um, Q = 3, filled with oil-like material, walls hyaline, thin, smooth, acyanophilous or weakly cyanophilous, not reacting in Melzer’s reagent. HABITAT & DISTRIBUTION — On logs, branches and twigs of deadangiosperms from southeastern United States (Florida, Louisiana) to southeastern Brazil. REPRESENTATIVE SPECIMENS EXAMINED — ARGENTINA. MIsIONES PROVINCE: Urugua-i Provincial Park, Wanda, 26 May 2001, D Kriiger 11279 (TENN 59088). BRAZIL. Rio GRANDE DU SOL PROVINCE: Sao Leopoldo, 1904, Rick (S F-249620); Rick- Fungi Austro-Americano Exsiccati nr. 141, 1906 (S F-177427, F-177429); Ijuhy, Exped. Imae Regnelliana no. 345B, 7/4 1893, GA Malme (S F-177430). COLOMBIA. Cauca District: ad pag. El Tambo, 1700 m, 13 Nov 1937, Kv Snidern (S F-177424). Valle del Cauca Dept., elev. 1980 m., 23 km from Cali, on Cali-Cisneros, on branch, 30 Aug 1976, KP Dumont & LA Molina, Dumont-Co 7669 (NY). COSTA RICA. PUNTRENAS PROVINCE: Coto Brus County, San Vito, Hacienda La Amistad, 3 Jul 1998, RH Petersen 9486 (TENN 56514); Tablazo, 1800 m, 8 Aug 1932, M Valerio (S F-177432). CARTAGO PROVINCE: mountains east of Tulis and north of Plantanillo, 2 Sep 1968, DE Stone 7141 (NY 00536421). CUBA. Fungi Cubensis Wrightiani no. 343, [Wright] (S F-15535); Monte Verde, Fungi Cubensis Wrightiani no. 237, 7-4 1915, C Wright (S F-15533). PINAR DEL RIO PROVINCE: San Diego de Los Bajos, on old log, 31 Aug to 3 Sep 1910, NL Britton, FS Earle & OS Gages, Britton 6823 (NY). JAMAICA. St. ANN ParIsH: near Reynolds bauxite mines, Lynford P.O., 1200-1400 ft., on dead branch of small tree, 23 Sep 1954, GR Proctor, F1288 (NY 00536418); Moneague to Hollymount via Union Hill, 700-750 m. elev., 6 Aug 1957, AL Welden 558 (NY 00536419). PARAGUAY. ALTro PARANA STATE: Ciudad del Este Hernandarias, Tati Jupi Reserve on Iaipu Lake, on
Fics 30-31. Polyporus epitheloides. 30. Basidiospores: A. from TENN 56514; B. from TENN 59088. Mycobonia winkleri (holotype F-15807). 31. A. Basidiospores; B. broom-like structures in pileus context; C. hyphal segments from pileus context.
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hardwood branch, 25 May 2001, RH Petersen (TENN 58933); SAN ANTONIO PROVINCE: Asuncion, Exped. Imae Regnellian Fungi, July 1893, GA Malme (S F-177425). UNITED STATES. Fioripa: Florida Caverns State Park, on dead wood, 5 May 1962, AL Welden 4428 (NY 00536416); Miami, on twigs, 15 Jan 1919, WH Long (CFMR FP 30869); Dade County, Matheson’s Hammock, on twigs, 23 Feb 1922, JA Stevenson 1630 (BPI 261319). Loutsi1ANa: Plaquemines Parish, Tulane University, F Edward Herbert Center, on dead wood, 25 Jul 1972, AL Welden 7764 (NY 00536411); St. Martinville, on fallen branches, 20 Aug 1898, AB Langlois 2817 (S F-249626).
DESCRIPTIONS & ILLUSTRATIONS — Burt (1919: 262), Corner (1984: 104), Julich (1976:
450, description only), Patouillard (1894, plate III, fig. 2), Reid (1976: 190, figs 1, 5),
Ryvarden (2010: 119). ComMENTsS — Polyporus epitheloides is a pileate species characterized by a dense layer of hyphal pegs in the hymenium and large cylindric to subfusiform basidiospores. The peculiar thick-walled cystidia seen in this species were also observed in P. polyacanthophorus and could be interpreted as terminally differentiated skeleto-binding hyphae. Jilich (1976: 451) refers to these structures as tramal cystidia and includes an illustration. The description above is based on Jiilich (1976), Reid (1976), and personal observations. Transfer of the species to Polyporus requires a new name because the epithet ‘flavus’ is preoccupied.
Polyporus epitheloides is closely related to P. polyacanthophorus [= M. brunneoleuca] but can be differentiated by its narrowly cylindrical to subfusiform basidiospores that are usually <8 um diam, Q = 3. Polyporus epitheloides has a wider distribution than P polyacanthophorus, extending into southeastern United States. ITS sequences of P. epitheloides (AY513571 as M. flava) and P. polyacanthophorus (AY513569, AY513570 also as M. flava) differ by 5%, confirming their status as distinct species. Another similar species is Pseudofavolus cucullatus (Mont.) Pat. [= Polyporus curtipes]. It can be distinguished from P. epitheloides by its angular to hexagonal pores, 1-3 per mm, and smaller, cylindrical basidiospores (11.5-)13-16 x 4-6 um (Ryvarden & Johansen 1980: 514). The isotype of Favolus curtipes Berk. & M.A. Curtis (South Carolina, Santee, Ravenel 378, HUH 00290582) was examined (see Fics. 19-20 for photographs of the poroid hymenophore).
Kriiger (2002) and Kriiger & Gargas (2010) considered M. flava to be a subspecies of P. curtipes despite a 4—7% difference in ITS sequences. They noted that M. flava, with spines, and Ps. cucullatus, with pores, were otherwise similar in basidiome coloration, hyphal construction, and shape and size of basidia and basidiospores. Because they are sister taxa embedded in Polyporus sensu stricto as demonstrated by phylogenetic analyses of the large subunit ribosomal RNA gene and ITS region, Kriiger (2002) and Kriiger & Gargas (2010) concluded that M. flava and Ps. cucullatus belong in Polyporus. Following Ryvarden (1991:
Unusual crust and pileate basidiomycetes ... 393
213), they accepted the name P. curtipes for Ps. cucullatus and then reduced M. flava to a subspecies of P. curtipes.
It should be noted that Kriiger (2002) and Kriiger & Gargas (2010) misidentified TENN 057579, FB 10256, as M. flava. Based on basidiospore size, this specimen represents P. polyacanthophorus [= M. brunneoleuca)]. Thus, ITS (AY513569, AY513570) and LSU sequences (AJ487934) correspond to P. polyacanthophorus whereas AY513571, ITS sequence of TENN 059088, FB 11279, is correctly identified as P. epitheloides [= M. flava].
The following descriptions and illustrations should be used with caution, for the authors did not differentiate between M. flava and M. brunneoleuca or did not provide basidiospore measurements, so it is not possible determine the species involved — Banker (1906), Berkeley (1843, plate 10, fig. 8), Gibertoni et al. (2006), Hennings (1900), Ibanez (1999), and Lloyd (1915). Although Martin (1939: 247) considered M. brunneoleuca a synonym of M. flava, the specimens he cited from Panama are redetermined here as P. polyacanthophorus.
Mycobonia winkleri Bres., Ann. Mycol. 9: 551. 1911. FiGs.23—24, 34
BASIDIOME dimidiate-sessile to subresupinate, 17 x 7 mm, 320-400 um thick, soft, fragile, greyish orange (5B4). PILEUS SURFACE warty to wrinkled; HYMENIAL SURFACE smooth with warts, 2-3 per mm, with a dark cavity or reddish brown, mucilaginous material at apex; context cottony, orange white (5A2).
HYPHAL SYSTEM dimitic with clamped generative and aseptate skeletal hyphae. Warts consisting of bulbils embedded in context and hymenium, composed of pseudoparenchymatous tissue enclosing a mass of dark reddish brown, amorphous material. PILEUS CORTEX a densely agglutinated tissue composed of irregularly thick-walled hyphae, thick-walled, broom-like structures, and scattered dichophyses; pileal hyphae 3-8 um diam, clamped, with numerous short branches, irregular, walls hyaline, thick, smooth; broom- like structures 17-30 x 3-8 um, with a thick, robust main trunk that gives rise to branches short and stubby or longer, <12 x 2 um, with knobby protuberances, walls hyaline, thick, smooth, cyanophilous, not reacting in Melzer’s reagent; dichophyses with short, slender branches radiating from a central point, walls hyaline, thick, smooth, cyanophilous, dextrinoid. Beneath cortex a narrow, moderately dense zone of generative subicular and skeletal hyphae, then a zone of open, loosely intertwined subicular hyphae and scattered dichophyses, about 140 um thick; subicular hyphae 3-5 um diam, clamped, moderately branched, walls hyaline, thin to 2 um thick, sometimes irregularly thickened, smooth, acyanophilous, not reacting in Melzer’s reagent; skeletal hyphae 1.3-2.2 um
394 ... Nakasone
diam, aseptate, sparsely branched, walls hyaline, thick, smooth, cyanophilous, dextrinoid in Melzer’s reagent. SUBHYMENIUM a dense, agglutinate tissue of hyphae similar to subicular hyphae. HYMENIUM a dense, agglutinated palisade of basidia. Basrp1a clavate, 25-28 x 12 um, bisterigmate (Bresadola, 1911). BASIDIOSPORES scarce, globose, subglobose or broadly lacrymoid, 6-7.5(-7.9) x (5.5-)6-7.2 um, walls hyaline, thin, smooth, cyanophilous, not reacting in Melzer’s reagent. TYPE SPECIMEN EXAMINED — S.O. BORNEO, ad Calamus rotang, 24 Jun 1902, H Winkler 2605 (S F-15807, holotype).
ComMENts — Mycobonia winkleri is a conundrum. The fungus displays a bewildering array of morphological features and the name cannot be placed in any extant genus. In the first place, the unusual structures that resemble hyphal pegs in M. winkleri (and probably the reason Bresadola placed the species in Mycobonia) are not composed of hyphae but consist of an amorphous reddish brown material. Second, the cyanophilous dextrinoid dichohyphidia developed in M. winkleri resemble those found in some species of Vararia P. Karst., such as V. minidichophysa Boidin & Lanq., V. microphysa Boidin & Langq., and V. tropica A.L. Welden. Boidin et al. (1980) described these structures as capillary dichophyses. Two species of Amyloflagellula Singer, A. verrucosa Agerer & Boidin and A. inflata Agerer & Boidin, also develop similar dichohyphidia (Agerer & Boidin 1981). Third, the broom-like structures found in the pileus cortex recall those produced in the pileipellis and gills of some Marasmius Fr. species. However, the rather large, dimidiate-sessile basidiome of M. winkleri is unlike that of Vararia (effused), Amyloflagellula (cup-shaped), or Marasmius (mushroom-shaped).
It is premature to erect a new genus for M. winkleri at this time. The type is fragile and somewhat deteriorated. Additional collections are needed to confirm that the disparate characters are produced by this taxon. The hyphal peg-like structures are particularly mysterious and in need of further study. DNA sequence data would be desirable to discover its closest relatives.
Acknowledgments
Curators and assistants of the following herbaria are thanked for arranging specimen loans: BPI, HUH, LY, NY, O, PDD, PRM, TAAM, TENN and S. Comments and corrections of Drs. Andrew M. Minnis, James H. Ginns, and Harold H. Burdsall, Jr., who critically reviewed this manuscript, are greatly appreciated.
Literature cited
Agerer R, Boidin J. 1981. The genus Amyloflagellula in West Africa (Basidiomycetes, ‘Cyphellaceae’). Sydowia 34: 1-12.
Banker HJ. 1906. A contribution to a revision of the North American Hydnaceae. Mem. Torrey Bot. Club 12: 99-194.
Unusual crust and pileate basidiomycetes ... 395
Berkeley MJ. 1843 [“1842”]. Notices of fungi in the herbarium of the British Museum. Ann. Mag. Nat. Hist. 10(suppl.): 369-385.
Binder M, Larsson K-H, Matheny PB, Hibbett DS. 2010. Amylocorticiales ord. nov. and Jaapiales ord. nov.: Early diverging clades of Agaricomycetidae dominated by corticioid forms. Mycologia 102: 865-880. http://dx.doi.org/10.3852/09-288
Binder M, Justo A, Riley R, Salamov A, Lopez-Giraldez F, Sjékvist E, Copeland A, Foster B, Sun H, Larsson E, Larsson K-H, Townsend J, Grigoriev IV, Hibbett DS. 2013. Phylogenetic and phylogenomic overview of the Polyporales. Mycologia 105: 1350-1373. http://dx.doi.org/10.3852/13-003
Boidin J, Gilles G. 1991. Basidiomycétes Aphyllophorales de ile de la Réunion. XVI. Les genres Hyphoderma, Hyphodermopsis, Chrysoderma nov. gen. et Crustoderma. Cryptog. Mycol. 12: 97-132
Boidin J, Lanquetin P. 1975. Vararia subgenus Vararia (Basidiomycetes Lachnocladiaceae): étude speciale des especes d'Afrique intertropicale. Bull. Trimestriel Soc. Mycol. France 91: 457-513.
Boidin J, Lanquetin P, Gilles G. 1980. Application du concept biologique de lespece aux Basidiomycetes: le genre Vararia (section Vararia) au Gabon. Cryptog. Mycol. 1: 265-384.
Bresadola J. 1911. Fungi Borneenses. Lecti a cl. Hubert Winkler anno 1908. Ann. Mycol. 9: 549-553.
Burt EA. 1919. The Thelephoraceae of North America. XI. Tulasnella, Veluticeps, Mycobonia, Epithele, and Lachnocladium. Ann. Missouri Bot. Gard. 6: 253-281. http://dx.doi.org/10.2307/2990131
Burt EA. 1926. The Thelephoraceae of North America. XV. Corticium. Ann. Missouri Bot. Gard. 13: 173-354. http://dx.doi.org/10.2307/2394065
Corner EJH. 1984. Ad Polyporaceas 11 — Polyporus, Mycobonia, and Echinochaete. Beih. Nova Hedwigia 78. 129 p.
Cunningham GH. 1956. Thelephoraceae of New Zealand— Parts VII and VIII. Part VIII — The genera Epithele and Mycobonia. Trans. Roy. Soc. New Zealand 83: 628-636.
Cunningham GH. 1963. The Thelephoraceae of Australia and New Zealand. Bull. New Zealand Dept. Sci. Industr. Res. 145. 359 p.
Donk MA. 1964. A conspectus of the families of Aphyllophorales. Persoonia 3: 199-324.
Gerlach ACL, Loguercio-Leite C. 2011. Hydnoid basidiomycetes new to Brazil. Mycotaxon 116: 183-189. http://dx.doi.org/10.5248/116.183
Gibertoni TB, Ryvarden L, de Queiroz Cavalcanti MA. 2006. Steroid fungi (Basidiomycota) of the Atlantic Rain Forest in northeast Brasil. Nova Hedwigia 82: 105-113. http://dx.doi.org/10.1127/0029-5035/2006/0082-0105
Ginns J. 1976. Corneromyces kinabalui, gen nov., sp. nov. (Aphyllophorales, Coniophoraceae). Mycologia 68: 970-975. http://dx.doi.org/10.2307/3758712
Hennings P. 1900. Hymenomycetineae. 104-276, in: A Engler, K Prantl (eds). Die natirlichen Pflanzenfamilien. Teil I, Abt. 1**. Leipzig, Wilhelm Engelmann.
(http://www. biodiversitylibrary.org/item/23586].
Hjortstam K. 1983. Studies in tropical Corticiaceae (Basidiomycetes) V. Specimens from East Africa collected by L. Ryvarden. Mycotaxon 17: 555-572.
Hjortstam K, Larsson K-H. 1995. Annotated check-list to genera and species of corticioid fungi (Aphyllophorales, Basidiomycotina) with special regards to tropical and subtropical areas. Windahlia 21. 75 p.
Hjortstam K, Ryvarden L. 2001. Corticioid species (Basidiomycotina, Aphyllophorales) from Colombia III. Mycotaxon 79: 189-200.
Hjortstam K, Ryvarden L. 2008. Some corticioid fungi (Basidiomycotina) from Ecuador. Syn. Fungorum 25: 14-27.
396 ... Nakasone
Hohnel F. 1910. Fragmente zur Mykologie. XII. Mitteilung, Nr. 574 bis 641. Sitzungsber. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 119: 877-958.
Hohnel F, Litschauer V. 1907. Beitrage zur Kenntnis der Corticieen (II. Mitteilung). Sitzungsber. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Cl, Abt. 1, 116: 739-852.
Ibanez CG. 1999. Cultural studies of non-poroid Aphyllophorales. Mycotaxon 71: 457-471.
Julich W. 1975. On Cerocorticium P. Henn., a genus described from Java. Persoonia 8: 217-220.
Julich W. 1976. Studies in hydnoid fungi X I. On some genera with hyphal pegs. Persoonia 8: 447-458.
Jiilich W. 1979. Studies in resupinate Basidiomycetes — VI. On some new taxa. Persoonia 10: 325-336.
Julich W. 1982 [“1981”]. Higher taxa of Basidiomycetes. Biblioth. Mycol. 85. 485 p.
Kirk PM, Cannon PE, Minter DW, Stalpers JA (eds.). 2008. Dictionary of the fungi, 10" edition. Wallingford, CABI.
Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. Eyre Methuen, London.
Kotlaba F, Pouzar Z. 1964. Preliminary results on the staining of spores and other structures of Homobasidiomycetes in cotton blue and its importance for taxonomy. Trans. Brit. Mycol. Soc. 47: 653-654.
Kriger D. 2002. Monographic studies in the genus Polyporus (Basidiomycotina). PhD dissertation, Department of Botany, University of Tennessee. Knoxville.
Kriiger D, Gargas A. 2004. The basidiomycete genus Polyporus — an emendation based on phylogeny and putative secondary structure of ribosomal RNA molecules. Feddes Repert. 115: 530-546. http://dx.doi.org/10.1002/fedr.200311052
Kriiger D, Gargas A. 2010. Unusual polypore fungi — a taxonomic emendation of Polyporus (Basidiomycotina) after ribosomal spacer characters. Cryptog. Mycol. 31: 389-401.
Liberta AE. 1969. Descriptions of the nomenclatural types of corticiums described by Burt. Nova Hedwigia 18: 215-233.
Lloyd GC. 1915. Letter no. 56, pp. 1-12. Mycological Writings 4.
Maekawa N, Suhara H, Kinjo K, Kondo R. 2003. Corticioid fungi (Basidiomycota) in mangrove forests of the islands of Iriomote and Okinawa, Japan. Mycoscience 44: 403-409. http://dx.doi.org/10.1007/S10267-003-0133-0
Martin GW. 1939. New or noteworthy fungi from Panama and Colombia. III. Mycologia 31: 239-249. http://dx.doi.org/10.2307/3754518
Nakasone KK. 2008. Type studies of corticioid Hymenomycetes described by Bresadola. Cryptog.
Mycol. 29: 231-257.
Nakasone KK. 2013. Taxonomy of Epithele (Polyporales, Basidiomycota). Sydowia 65: 59-112.
Nakasone KK, Lindner DL. 2012. Taxonomy of Pseudolagarobasidium (Polyporales, Basidiomycota).
Fungal Diversity 55: 155-169. http://dx.doi.org/10.1007/s13225-012-0161-1
Nufez M, Ryvarden L. 1995. Polyporus (Basidiomycotina) and related genera. Syn. Fungorum 10. 85 p.
Parmasto E. 1968. Conspectus systematis corticiacearum. Tartu, Academiae Scientiarum R.PS.S.
Estonicae. Patouillard MN. 1892. Champignons nouveaux Extra-Européens. Bull. Soc. Mycol. France 8: 46-56. Patouillard MN. 1894. Espéces critiques d Hyménomycetes. Bull. Soc. Mycol. France 10: 75-81. Pouzar Z. 1987. Taxonomic studies in resupinate fungi III. Ceska Mykol. 41: 26-28. Reid DA. 1976. Notes on polypores. 2. Mem. New York Bot. Gard. 28: 179-198. Ridgway, R. 1912. Color standards and color nomenclature. Washington, DC, publ. by author.
Unusual crust and pileate basidiomycetes ... 397
Ryvarden L. 1991. Genera of polypores — nomenclature and taxonomy. Syn. Fungorum 5. 363 p.
Ryvarden L. 2010. Stereoid fungi of America. Syn. Fungorum 28. 206 p.
Ryvarden L, Johansen I. 1980. A preliminary polypore flora of East Africa. Oslo, Fungiflora.
Singer R. 1986. The Agaricales in modern taxonomy. Koenigstein, Koeltz Scientific Books.
Sotome K, Hattori T, Ota Y, To-anun C, Salleh B, Kakishima M. 2008. Phylogenetic relationships of Polyporus and morphologically allied genera. Mycologia 100: 603-615. http://dx.doi.org/10.3852/07-191R
Talbot PHB. 1951. Studies of some South African resupinate Hymenomycetes. Bothalia 6: 1-116.
Thiers B. 2014 [continuously updated]. Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg. org/ih/ [accessed January 2014].
Trierveiler-Pereira L, Baltazar JM, Loguercio-Leite C. 2009. Santa Catarina Island mangroves 4 — xylophilous basidiomycetes. http://www.mycotaxon.com/resources/checklists/trierveiler- v109-checklist.pdf [Summary: Mycotaxon 109: 107-110. http://dx.doi.org/10.5248/109.107].
Wu S-H, Chen Z-C. 1990. New records of Corticiaceae (Basidiomycetes) collected from the National Taiwan University campus. Taiwania 35: 64-67.
ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MY COTAXON
http://dx.doi.org/10.5248/130.399 Volume 130, pp. 399-419 April-June 2015
A new Graphilbum species from western hemlock (Tsuga heterophylla) in Canada
JAMES REID & GEORG HAUSNER*
Department of Microbiology, University of Manitoba, Winnipeg, MB, R3T 2N2
* CORRESPONDENCE TO: hausnerg@cc.umanitoba.ca
ABSTRACT — A new species of Graphilbum, G. tsugae isolated from Tsuga heterophylla, is described from British Columbia, Canada. Although rDNA ITS data analysis demonstrates its close relationship to Graphilbum rectangulisporium, morphologically the unique apical projections arising from the perithecial necks of G. tsugae easily differentiate it from the former and all other described Graphilbum and Ophiostoma species. We also correct the epithet orthography and the parenthetical basionym author citation of the name G. rectangulisporium.
KEY worps — ophiostomatoid fungi, bark beetles, morphology, taxonomy, ITS rDNA region.
Introduction
During a reassessment of isolates of ophiostomatoid fungi in our culture collection whose taxonomic placements had never been resolved, we examined three strains from a single location in southern British Columbia isolated from two felled logs of Tsuga heterophylla. These logs had shown clear evidence of having been attacked by bark beetles — many bore holes were evident and peeling back the bark around the holes revealed well-developed galleries — but neither beetle adults nor larvae were present and the surrounding sapwood was only slightly stained. However, hand lens examination did show both remnant hyphae and possible reproductive structures scattered throughout gallery interiors.
Slabs of bark-covered wood about 12 inches long, 6-8 inches wide, and 1-3 inches thick were harvested from such areas using a hatchet and a chisel. The bark was then removed to expose the outer sapwood surfaces and the inner bark surfaces of these slabs. After the wood slabs and bark pieces were examined carefully to ensure no adult beetles or larvae were present in any of the exposed galleries, these materials were placed and sealed in collection bags for return to the laboratory.
400 ... Reid & Hausner
When we began our detailed study of this species, the Ophiostomatales accommodated three genera: Ceratocystiopsis H.P. Upadhyay & W.B. Kendr.; Grosmannia Goid.; and Ophiostoma Syd. & P. Syd., among which species economically important tree pathogens and blue-stain fungi are found (Olchowecki & Reid 1974, Wingfield et al. 1993, Hausner et al. 2005).
Based on partial LSU rDNA and 6-tubulin sequences, Zipfel et al. (2006) provided some support for the circumscription of the above three genera. But Zipfel et al. (2006, p. 95) and other authors (Hafez et al. 2012, 2013) also noted that the use of molecular criteria might require the erection of new genera to accommodate either previously misplaced or subsequently newly discovered species of Ophiostoma sensu lato.
Recently de Beer & Wingfield (2013), using partial LSU and SSU rDNA sequences, have begun redefining the Ophiostomatales. They include only one family, the Ophiostomataceae, in which they recognize six distinct lineages represented by the genera Ceratocystiopsis H.P. Upadhyay & W.B. Kendr. emend. Zipfel et al. (2006), Fragosphaeria Shear, Leptographium Lagerb. & Melin sensu lato (includes Grosmannia), Ophiostoma sensu lato (includes Pesotum and Sporothrix), Raffaelea Arx & Hennebert emend. T.C. Harr. in Harrington et al. (2008), and Graphilbum H.P. Upadhyay & W.B. Kendr. emend. Z.W. de Beer et al. (2013a). They also clearly implied the possible need for further generic inclusions. Relevant comments are also found in related accompanying papers (de Beer et al. 2013a & b).
The genera of the Ophiostomataceae include economically important tree pathogens and blue-stain fungi (e.g., Lagerberg et al. 1927, Olchowecki & Reid 1974, Wingfield et al. 1993, Hausner et al. 2005) whose members lack forcible ascospore discharge, have deliquescent asci, develop sticky ascospore droplets at the apex of their perithecial necks, and typically produce slimy/sticky conidia on various simple or complex conidiophore structures. Species assigned to Ceratocystiopsis resemble those of Ophiostoma except in Ceratocystiopsis the small dark perithecia only develop short perithecial necks and produce falcate, sheathed ascospores (Upadhyay 1981, Zipfel et al. 2006, Plattner et al. 2009). These species also have a lower tolerance to cycloheximide than do Ophiostoma species (Harrington 1981, Hausner et al. 1993). Species formerly assigned to Grosmannia (above) were distinguished by the fact they have Leptographium states.
As we could not definitively establish the identity of our three isolates using only morphological criteria, we compared nuclear small subunit rDNA (rSSU) and rDNA internal transcribed spacer (ITS) region (ITS1, 5.8S gene, and ITS2) sequences from our isolates with those of other Ophiostoma species. We also assessed the growth rates and cultural characteristics of our isolates at different
Graphilbum tsugae sp. nov. (Canada) ... 401
temperatures and compared our results with data published for other species. We did not assess mating type features, as we had earlier when describing another ophiostomatoid species (Reid et al. 2010), as mating type studies had not been conducted for any of the species with which we compared our current isolates.
Based on the revisions discussed above, we believe our results show that our isolates represent a new taxon within the genus Graphilbum that we describe here as Graphilbum tsugae.
Materials & methods
Isolation of strains studied, culturing procedures, and growth-temperature experiment
Laboratory isolations were made from field collected wood and bark samples and grown on agar plates in a 20°C incubator; these were examined periodically for evidence of fungi of interest.
We obtained three isolates that, based solely on morphological criteria, appeared to represent an undescribed Ophiostoma species. Strains examined and/or sequenced are detailed in TaBLeE 1. Living cultures and dried specimens of our new species have been deposited in the University of Alberta Microfungus Collection and Herbarium (UAMH), Edmonton, Canada; the additional DNA sequences retrieved from GenBank employed in the analyses are also listed in TABLE 1.
Based on the intent of the experiment, strains were grown and characterized employing either malt extract agar supplemented solely with yeast extract (Reid et al. 2010) or such agar that also contained wood chips (Reid & Hausner 2010); fertile perithecia were regularly produced on the latter plates. These were then dried, and one was selected to be the type specimen. The parent culture was isolate WIN(M) 1391, obtained from a perithecial spore drop produced at the apex of a perithecium found in the gallery of an unidentified bark beetle. Preparations for microscopy were mounted in 85% lactic acid and at least 50 measurements were made for each significant morphological feature. Colour designations are based on Rayner (1970).
In a previous study (Reid et al. 2010) where we had found a group of very similar isolates growing on pieces of substrate in very close proximity to each other — as was the case here — we had processed three of them in a temperature growth-rate response experiment, and they produced very similar growth-temperature files. They and the other isolates in the group were described as Ceratocystis norvegica J. Reid & Hausner [sic; = Georg Hausner].
Because of that experience, the precise identity of the gene regions sequenced in our current three isolates, and the lack of any significant morphological variation noted between them, we used only the designated holotype isolate, VAMH 11701 [WIN(M) 1391] for determining the temperature growth-rate response of G. tsugae.
We employed the protocols of van Wyk et al. (2006), as modified by Reid et al. (2010). However, because of the cultures’ relatively slow growth, the first measurements were not made until the second day after plate inoculation. We used eight replicate plates at each test temperature. Although initially, the total temperature-growth test
402 ... Reid & Hausner
TABLE 1. The rDNA ITS region sequences evaluated
TAXON STRAINS/ISOLATES/SOURCE! GENBANK Ceratocystiopsis minuta °CBS 145.59 DQ128173 C. cf. minuta SL-K 70 DQ128175 Grosmannia penicillata RJ-T 0125 AM943882 Ophiostoma abietinum CMW 397 DQ396788 O. dentifundus CMW 13016? AY495434 CMW 13017? AY495435 O. ips CMW 22843 DQ539549 O. grandicarpum Strain D AJ293884 O. microcarpum YCC 459 AB506676 YCC 612 GU134170 O. piceae Ci-37 EF506934 O. pulvinisporum CMW 9026 AY546715 O. rectangulisporium TEM:FPH 7756 AB242825 YCC 617 GU134171 O. cf. rectangulisporium CMW 26261 > EU785451 CMW 26259° EU785450 CMW 26258 > EU785449 C2477 GU129987 C2300 GU393357 253GRJ © JX444645 1313RJP¢ JX444594 1039RJP * JX444590 1037RJP * JX444589 1284RJP ¢ JX444591 1308RJP * JX444593 1306RJP ¢ JX444592 246bGRJ JX444644 CMW 22830 4 DQ539536 CMW 22829 4 DQ539535 CMW 22832 4 DQ539538 CMW 22831 4 DQ539537 1132RJ ¢ (=voucher 1138RJ) JQ289021 O. rostrocoronatum CBS 434.77 AY194509 Graphilbum tsugae WIN(M) 1391 *(=UAMH 11701) KJ661745 WIN(M) 1384 ¢ (=UAMH 11700) = WIN(M) 1387 *(=UAMH 11699) — Ophiostoma sp.1 WIN(M) 1602 HM363177 WIN(M) 1603 HM363164 Ophiostoma sp. 2 RJ-0771 ! AM943895 RJ-0704 § AM943894
Ophiostoma sp. 3 PR-2006c (=CMW 27315) DQ674367
Graphilbum tsugae sp. nov. (Canada) ... 403
Pesotum fragrans CMW 22853 DQ539561 C01-014a § AY194513 NLC 348 8 AY194518 CBS 279.54 §(=C1224) AF198248 WIN(M) 1388 8 _— C990 DQ062976 P cupulatum C1194 AF198230 Sporothrix inflata CMW 12529 AY495428
‘Strain numbers followed by identical superscript letters (* to 8) have identical ITS regions and only one representative was used in the ITS phylogenetic analysis.
*CBS = CBS-KNAW Fungal Biodiversity Center, Utrecht, Netherlands; WIN(M) = culture collection of J. Reid, Microbiology, University of Manitoba, Winnipeg, Manitoba, Canada; UAMH = University of Alberta Microfungus Collection and Herbarium, Devonian Botanic Garden, Edmonton, Alberta, Canada; CMW, Tree Pathology Co-operative Program, Forestry and Agricultural Biotechnology Institute, University of Pretoria, South Africa; STE-U, Stellenbosch University, South Africa.
period was to have lasted seven days, we incubated all plates further at 20°C to record morphological development.
DNA extraction, amplification protocols and DNA sequencing
DNA extraction, purification, and agarose electrophoresis protocols followed Hausner et al. (1992). Genomic DNA served as the amplification template using the Invitrogen-Life Technologies PCR System (Buffer and Taq polymerase, Invitrogen, Fredrick, MD). Primers SSUZ and LSU4 (Hausner et al. 2005) were used to amplify the ITS regions. The PCR primer sequences, amplification conditions, sizes of the expected PCR products, and preparation of sequencing templates for fragments have been described previously (Hausner & Wang 2005, Hausner et al. 2005). DNA sequencing templates were prepared with the aid of the Promega Wizard SV Gel and PCR clean- up system (Promega, Madison, WI). Purified double-stranded PCR products were sequenced in both directions using the cycle-sequencing protocols performed according to the manufacturer's recommendations (Perkin Elmer Applied Biosystems, Foster City, CA), and automated Fluorescent DNA sequence analysis was performed using an ABI Prism 310 Genetic Analyzer system (PEAB at the University of Calgary, DNA sequencing facility, Calgary, AB).
Analyses of DNA sequence data
To add to sequences generated herein, we used the UAMH 11701 ITS sequence in a BLAST search to recover additional sequences from NCBI databases for comparison with other relevant taxa. Fifty ITS rDNA sequences were aligned with CLUSTAL-X (Thomson et al. 1997) and, when appropriate, modified with the alignment editor program GeneDoc v2.5.010 (Nicholas et al. 1997, http://www.psc.edu/biomed/genedoc). Phylogenetic trees were generated using the Molecular Evolutionary Genetic Analysis program package (MEGA 5.1; Tamura et al. 2011). Trees were generated with the Maximum parsimony (MP), Neighbor joining (NJ), and Maximum likelihood (ML) methods. MEGA 5.1 was also used for determining the best-fit DNA models and the best-fit substitution model for ML analysis. In order to evaluate node support within the
404 ... Reid & Hausner
tree topologies, the bootstrap option (1000 replicates) was selected for all tree-building programs.
Finally, MrBayes (version 3.1; Ronquist & Huelsenbeck 2003) was used for Bayesian analysis. The NEXUS file format necessary for the alignment (input) file was generated with the file converting option available within DAMBE (Xia 2001). The DNA substitution model setting for Bayesian analysis was chosen based on evaluating the ITS region alignment with the Modeltest 3.7 program (Posada & Crandall 1998). The Bayesian inference of phylogenies was initiated from a random starting tree, and four chains were run simultaneously for 3 million generations; trees were sampled every 100 generations. The first 25% of trees generated were discarded (burn-in); the remaining trees were used to construct a 50 % majority rule consensus tree and compute the posterior probability values.
Phylogenetic trees were drawn with the TreeView program (Page 1996) and figure annotations added using Corel Draw (Corel Corporation and Corel Corporation Limited).
Results
Cultural and morphological characteristics
The temperature-related growth response of isolate VAMH 11701 (Fie. 1) depicts a relatively broad growth temperature optimum, i.e. 20-25°C, dropping off sharply towards both 15 and 30°C. The mycelium amount produced differed between test temperatures but was always appressed, with only limited sparse, central aerial mycelium. During the first 7 days, the mycelium remained hyaline (translucent) and without coloration or reproductive structure development.
All eight plate sets from the various test temperatures were then placed in a single 20°C incubator.
70 60 Mean 90 colony 40 diam
20 10
0 0
10 15 20 25 30 35
Temperature ( C)
Fic. 1: Temperature-growth curve for Graphilbum tsugae holotype strain UAMH 11701 incubated at various temperatures for 7 days.
Graphilbum tsugae sp. nov. (Canada) ... 405
By dayl0, mycelium in all original 20°C and 25°C cultures had reached Petri dish margins. It was still mostly hyaline and appressed, but with a ring of slightly more floccose white-colored mycelium around the paler center (Fic. 2A); these cultures were virtually identical in growth rates and appearance.
The 15°C and 30°C cultures had also grown further; their diameters were still quite small, but the 10°C and 35°C cultures showed only numerous short hyphae extending from their inoculation discs; apparently the fungus could survive a short exposure to the relatively high 35°C temperature.
All plate sets were then returned to the 20°C incubator until day 17.
In all original 20°C cultures, the mycelium had completely covered the dish surfaces. It was still quite peripherally appressed, more floccose centrally, and while still primarily white, there were definite scattered amber tints and a central darkening (Fic. 2B). Very occasionally scattered immature differentiating perithecia were observed, but only a few showed what might have been developing neck initials.
On day 17 in the original 25°C cultures, the mycelium had also reached dish margins. Generally it was still white, but now there were scattered slight amber tints. Less aerial mycelium was present peripherally than centrally, but that difference was now less pronounced. Overall there was less aerial mycelium than in the 20°C plates, but the central dark (Fuscous Black) pigmentation (Fic. 2C) was more pronounced and perithecial development more advanced. In six dishes the perithecia were quite numerous but fewer in the remaining two, suggesting that 25°C may be slightly better for perithecial production.
On day 17 in the original 10°C cultures, all isolates were growing, but no mycelium had reached a dish margin. Colony diameters averaged 70 mm, the mycelium was white and peripherally appressed to slightly floccose centrally, but no perithecial initials were present. In the original 15°C cultures, mycelium, which reached the dish margins randomly, was white, appressed peripherally, and slightly floccose centrally.
On day 17 in the original 30°C cultures, mycelium had reached the dish margins uniformly, but again the amounts of central and peripheral mycelium differed markedly. The peripheral mycelium was white and appressed while the central mycelium was more floccose and showing numerous presumptive perithecium initials. And while all original 35°C cultures were growing, no mycelium had reached a dish margin. Colony diameters averaged 53 mm, and in every case the mycelium was white, appressed peripherally and more floccose centrally, and there was no evidence of perithecia.
After inspection, we retained only the original 20°C and 25°C cultures, which were thereafter both maintained at 20°C.
By day 24, these retained cultures were similar in appearance. New mycelium was less dense, older mycelium had collapsed centrally, and scattered amber
406 ... Reid & Hausner
Fic. 2. Graphilbum tsugae (holotype, VAMH 11701 [WIN(M) 1391]). Morphology after growth at various temperatures. A. Original 20 °C plate at day 10. B. Original 20 °C plate at day 17. C. Original 25 °C plate at day 17. D. Original 25 °C plate at day 24. E. Original 25 °C plate after c. 7 weeks; F. Original 20 °C plate after c. 7 weeks.
Graphilbum tsugae sp. nov. (Canada) ... 407
colored patches, which were more common, often had turned pale yellowish- brown. More dark pigment was present centrally in the agar, and mounds of floccose white hyphae had formed at the colony margins. Although many perithecial initials were seen, no mature perithecia were found (Fic. 2D, an original 25°C culture).
Fic. 2E illustrates changes that occurred in an original 25°C culture over a further period of approximately 6-8 weeks. Note the increased amount of marginal mounded mycelium, increased area of central dark pigmentation, slight further overgrowth of superficial aerial mycelium, and finally, what appear to be abundant immature perithecia. Fic. 2F shows an even older original 20°C culture with extensive development of — now collapsed — marginal mounded mycelium but no apparent superficial mycelial overgrowth; there was also a restricted central dark pigmentation. Numerous perithecia were produced, but we only rarely saw any evidence of sporulation. Fics. 2E-F both show an extensive development of cinnamon-colored pigment throughout.
Because we rarely observed fully mature perithecia exuding spore droplets on agar dishes, we grew the strains on agar plus wood chips where mature perithecia were readily produced and their nature could be fully characterized (see TABLE 2 and Taxonomy section).
TABLE 2. Morphological comparisons between G. rectangulisporium and G. tsugae
CHARACTER PERITHECIAL BASE Shape Width (um) Height (um) PERITHECIAL NECKS Length (um)
Base width (um) Tip width (um) OSTIOLAR HYPHAE Length (um)
Number Shape (um)
ASCOSPORES (uum)
ANAMORPH
COLONY COLOR
Graphilbum rectangulisporium ¢
Globose 70-110 70-110
190-500
including ostiolar hyphae 20-35 10-19
13-43
5-15 Cylindrical with apical taper
2.0-3.5 x 1.0-1.7 with sheath; rectangular in side and face view
None found
White; remaining so on 2% MEA
*Ohtaka et al. 2006; Kirk et al. 2008; ‘Reid et al. 2010.
Graphilbum tsugae
Obpyriform, ampulliform or globose ?; 75-120(-167.5) 75 -113(-140)
(87-)138-200(-280) including ostiolar hyphae 27.5-50
1225225
(20-)25-40 Various Crooked to jagged in outline; at times irregularly cellular; tapering to a point. 3-5 x 1 with sheath; oblong to rectangular; sheath flaring at corners
Conidiophores simple to branched; not identifiable to genus; Conidia hyaline, 1-celled, oblong, 2-5 x 1-1.5 um.
White initially, coloring (pale yellowish brown) on aging on 2% MEA + 1% YES
408 ... Reid & Hausner
DNA sequence analysis
The rDNA SSU sequence (GenBank accession: HQ634824) for isolate UAMH 11701 — then as WIN(M) 1391 — was previously included in a study examining rDNA SSU intron distributions among ophiostomatoid fungi and related taxa (Hafez et al. 2012; p. 102). It showed that although UAMH 11701 was part of the Ophiostomataceae clade, it did not group with representatives of Grosmannia, Ceratocystiopsis or Ophiostoma sensu stricto. Instead, within the phylogenetic tree it assumed the most basal position among the sampled Ophiostomataceae taxa.
Our rDNA sequence analysis of the internal transcribed spacer (ITS) region (ITS1, 5.88, and ITS2; KJ661745) yielded even more surprising results. Using this sequence set from isolate UAMH 11701 — again as WIN(M) 1391— in a BLASTN analysis, revealed that its sequences are similar to those from this
Pesotum fragrans AF1982481° Ophiostoma microcarpum GU134170 19-9 Pesotum fragrans DQ062976 8% | + Ophiostoma sp. PR-2006c DQ674367
109 Ophiostoma microcarpum GU134170 ig Pesotum fragrans DQ539561 ; Ophiostoma cf. rectangulosporium Ja289021° Grap hilbum S) ® ! Ophiostoma cf. rectangulosporium JX444593 ©
Ophiostoma cf. rectangulosporium JX444644 Ophiostoma cf. rectangulosporium EU785451 B Ophiostoma sp. RJ-0771 AM943895 f Ophiostoma rectangulosporium GU134171 * Ophiostoma cf. rectangulosporium GU129987 ; Ophiostoma rectangulosporium AB242825 Ophiostoma sp. WIN(M)1602 HM363177 Ophiostoma sp. WIN(M)1603 HM363164 az Ophiostoma cf. rectangulosporium GU393357 a Graphilbum Sugee WIN(M)1391° Ophiostoma grandicarpum AJ2938 @ , Ceratocystiopsis minuta DQ1281 es Ceratocystiopsis cf. minuta DQ128175 @ Ophiostoma pulvinisporum AY546715
100
100 Ophiostoma ips DQ539549 es @) Ophiostoma piceae EF506934 ie Pesotum cupulatum AF198230 a7 Ophiostoma rostrocoronatum AY 194509
Ophiostoma abietinum DQ396788
55 Sporothrix inflata AY495428 Ophiostoma dentifundum AY495434"
Grosmannia penicillata AM943882
-——_———__ Node support @ = node support >99 for 0.05 values: MB, NJ, MP, ML
Fic. 3: Phylogenetic tree based on ITS sequence data. Levels of confidence for the major nodes in the phylogenetic tree are based on posterior probability (PP) values (MrBayes; first value), and bootstrap support (BS) in NEIGHBOR (second value), DNAPARS (third value), and Maximum likelihood analysis (fourth value). Nodes that received less than 50% support (BS or PP) were collapsed. Tree topology and branch lengths are based on Bayesian analysis and are proportional to the number of substitutions per site. GenBank accession numbers are listed next to species names.
Graphilbum tsugae sp. nov. (Canada) ... 409
region in strains identified as Pesotum fragrans, Ophiostoma rectangulisporium, Ophiostoma cf. rectangulisporium, and O. microcarpum (Fic. 3, TABLE 1). Comparisons within this ITS data set also strongly suggest that there are several yet to be described cryptic species within this clade. In particular the circumscription of O. rectangulisporium and isolates designated as O. cf. rectangulisporium need to be reevaluated, as the molecular data obtained failed to group these taxa into an appropriate clade relative to their assigned names. All these species are currently expected to be treated as members of the now holomorph genus Graphilbum (see below), whose species form a monophyletic group apart from species of Ophiostoma sensu stricto.
Taxonomy
Graphilbum tsugae J. Reid & Georg Hausner sp. nov. FIGS. 4-7 MycoBank MB809811 Differs from all previously described species of Graphilbum by its apical ostiolar projections being not true ostiolar hyphae but elongated tapering neck cells with jagged outlines, its rDNA sequences, and its variably solitary or sporodochial anamorphic conidiophores.
Type: Canada, British Columbia, Highway 1, Annis Mountain, near Salmon Arm, from a felled log of Tsuga heterophylla (Raf.) Sarg., isolated from spore drop at apex of a perithecium in unidentified bark beetle gallery, 21 September 1987, collected and isolated by J. Reid. (Holotype, UAMH 11701 [dried culture on agar plus wood chips], derived from herein designated isotype culture WIN(M) 1391.
EryMo_oey: tsugae, referring to the host genus Tsuga.
Maximum growth occurring on 2% malt-extract agar amended with yeast extract over a temperature range from 20°C to 25°C; maximum diameter 60 mm after seven days, i.e., relatively slow growing. No measurable growth at 10°C and below or 35°C and above. Colonies remaining only white over this period.
Anamorph apparently pleomorphic on wood chip agar plates. Some conidia developing terminally on short, simple, non-branched to slightly branched conidiophores. Others on variously branched conidiophore clusters that do not develop in any consistent branching pattern. Some are initially hyaline and verticillate, but later secondary cymose branches develop (see Seifert et al. 2011, fig. 100D) arising from beneath a septum in the parent branch. In others, branches that have arisen initially from a common point continue to branch along their length in a cymose fashion. Often these variously branched forms aggregate in clusters on the substrate, and then appear superficially as sporodochia until mounted. Conidiogenous cells proliferating in an imprecisely determined manner, as at times either sympodial or percurrent, or both presumptive forms can be observed in single mounts. Conidia one- celled, hyaline, oblong and parallel-sided in plan view (see Olchowecki &
410 ... Reid & Hausner
Fic. 4: Graphilbum tsugae (wood chip agar cultures). A. Simple conidiophores bearing conidia. B. Conidia in plan view (see Olchowecki & Reid 1974, p.1678 for definition). Scale bars: A = 35 um; B = 8 um.
Reid 1974, p. 1678 for definition); generally with slightly rounded ends, but in some cases one end flattened when fully abscised; very uniform in size, cylindrical in cross section (arrow); 3-5(-6) (mean + s.d. = 4.0 + 1.05) um long; frequently produced in slimy masses that occasionally form around the bases of perithecia developing on the wood chips. In such masses the more complicated conidiophores appear.
Mature perithecia black; initials produced abundantly on malt extract supplemented with yeast extract rarely maturing. Abundant mature perithecia developing in two months on plates containing wood chips embedded in the agar. Bases obpyriform, ampulliform, or globose, 75-120 (-168) um wide (mean + s.d. = 102 +16 um), 75-113 (-140) um high (mean + s.d = 94 + 19 um). Surface smooth to slightly irregular, often adorned with short dark hair-like hyphae. Neck black, (87-) 138-200 (-280) um long (mean + s.d. = 166 + 49 um), smooth to occasionally hairy; base 27.5-50 um wide (mean + s.d. = 39.1 + 6.24 um), tapering to 12.5-25 um (mean + s.d. = 18 + 4.0 um) just below the point of diverging apical projections; these projections are not ostiolar hyphae, but elongated terminal neck cells, (20-)25-40 um long (mean + s.d. = 33.5 + 7 um), crooked to jagged cellular extensions that taper to a point, apparently developing by differentiation from normal terminal neck cells; base 2-5 um
Graphilbum tsugae sp. nov. (Canada) ... 411
Fic. 5: Graphilbum tsugae (wood chip agar cultures). Variously branched conidiophores. A. Single verticillate conidiophore bearing conidia; arising from beneath septum in a parent branch (arrow). B. Portion of a compound aggregation of conidiophores showing successive cymose branches (see Seifert et al. 2011, fig.100D) in a single element (arrows). C. Aggregation of verticillate conidiophores formed in culture on and around the wood chips. Scale bars: A = 12 um; B = 30 um; C = 65 Um.
412 ... Reid & Hausner
Graphilbum tsugae sp. nov. (Canada) ... 413
wide (mean + s.d = 3.5 + 0.7 um). Ascospores hyaline, one-celled, oblong to rectangular with slightly rounded ends, rarely slightly constricted in the middle; sheath present but slightly flared and projecting out at the corners in plan view; 3-5 (mean + s.d. = 3.8 +. 0.7 x 1.0 um). ADDITIONAL MATERIAL EXAMINED: CANADA, British CoLumBiaA, Highway 1, Annis Mountain, near Salmon Arm, from a second felled log of Tsuga heterophylla at the same location from which the holotype material was obtained, 21 September, 1987. (1) A dried culture grown from isolate WIN(M)1387 (= UAMH 11699] originally isolated aseptically from a slab of stained wood found adjacent to the beetle galleries in the log; (2) A dried culture grown from isolate WIN(M)1384 [= UAMH 11700] that had been isolated from perithecial ooze extruded from a broken perithecium in the beetle galleries in a second small slab of wood taken from this second log. Both samples collected and isolated by J. Reid.
Finally, we correct errors in the citation of:
Graphium rectangulisporium (Ohtaka, Masuya & Yamaoka) Z.W.de Beer & M.J. Wingf. CBS Biodiversity Series 12: 269. 2013 [as “rectangulosporium (R.W. Davidson)” ]. = Ophiostoma rectangulisporium Ohtaka, Masuya & Yamaoka, Can. J. Bot. 84: 290. 2006
[as “rectangulosporium”; orthographic correction to comply with McNeill et al. 2012: Art. 60.8].
The miscitation of “R.W. Davidson” as the basionym author appears to have
been a transcriptional error.
Discussion
Kim (2010) and Kim et al. (2011) drew attention to a sterile, white, fast growing isolate (GU393357) obtained from Orthotomicus erosus (Wollaston) from Pinus spp. growing in California that groups closely with our new species (Fic. 3). However, both its reported growth rate and colour range differ markedly from those of G. tsugae. We believe both our morphological data and molecular analysis support our isolates as distinct from all other species formerly assigned to Ophiostoma. They are, however, linked by molecular criteria to a small group of other former Ophiostoma species whose anamorphs had been considered members of the then anamorphic genus Graphilbum. This observation is significant due to recent changes proposed by de Beer et al. (2013a, figs. 2 & 3, pp. 4 & 6) and de Beer & Wingfield (2013, figs. 4b & 5b, pp. 29 & 31). In both these papers Graphilbum is used as a holomorph- generic name in trees, and in de Beer & Wingfield (2013), the species group to be included in Graphilbum is discussed on page 39.
Fic. 6 (left): Graphilbum tsugae (wood chip agar cultures). A-C. Mature perithecia. A. Single mature perithecium and a detached perithecial neck. B. Cluster of five mature perithecia. C. Mature perithecia growing on wood chips in agar medium. Note the droplets in which aggregated conidiophores are often found. D- F. Apical neck projections. Scale bars: A, B = 110 um; C = 650 um; D = 15 um; E= 17 um; F = 10 um.
414 ... Reid & Hausner
Fic. 7 Graphilbum tsugae (wood chip agar cultures). Ascospores. Scale bars: A = 8 um; B = 7 um.
Graphilbum tsugae sp. nov. (Canada) ... 415
De Beer et al. (2013b, p. 268) formally redefined the generic name as: Graphilbum H.P. Upadhyay & W.B. Kendr., Mycologia 67: 800; 1975 emend. Z.W. de Beer, Seifert & M.J. Wingf. [type species Gra. sparsum] = Ceratocystis Ellis & Halst. section Ips H.P. Upadhyay pro parte, Monogr. Ceratocystis and Ceratocystiopsis, p. 70. 1981).
On page 269 discussing Graphilbum sparsum H.P. Upadhyay & W.B. Kendr. (1975), de Beer et al. (2013b), also state: ... “Graphilbum sparsum is the type species of the genus... re-introduced here to accommodate species previously treated in the P fragrans complex ... ”
Although this resulted in eight species being formally assigned to the genus Graphilbum — for only six of which phylogenetic data exists — there is now a ninth: an anamorphic species placed in Graphilbum by phylogenetic data (Romon et al. 2014). However, it is clear that de Beer and Wingfield (2013, p. 39) felt strongly that additional isolates studied by them might well deserve similar placement when further supportive data becomes available. Some such potential species are listed with former Ophiostoma spp. discussed in their species review under the name of Graphilbum (de Beer & Wingfield 2013, p. 39; figs. 4 b, p. 29, 5b, p. 31) and also seen in de Beer et al. (2013a, fig. 3, p. 6). Using phylogenetic data, three other species represented by new combinations in Hyalorhinocladiella proposed by Harrington et al. (2010) — H. ips, H. macrospora, H. tingens — were shown to represent anamorphs of three different Ophiostoma species (de Beer et al. 2013b).
The foregoing changes were made in conformance with the new rule of fungal nomenclature (McNeill et al. 2012, Article 59, Note 2), “... all legitimate fungal names are treated equally... regardless of the life history stage of the type... .” For a history of the origin of this approach see Hawksworth (2011). Following this practice, we describe our holomorphic isolates as representing a new species of the genus Graphilbum, originally an anamorphic name as emended above, even though there are some significant morphological inconsistencies among the species now included therein. Our new species introduction will undoubtedly add to these inconsistencies and to the impetus to erect further new genera.
Our ITS sequence analysis, which showed Graphilbum tsugae only distantly related to species accommodated in Ceratocystiopsis, Grosmannia, and Ophiostoma, grouped it in a clade previously referred to as the “P. fragrans & O. rectangulosporium clade” (Jankowiak 2012). Earlier studies based on ITS region analyses also showed that P. fragrans and O. rectangulisporium form a distinct lineage among the Ophiostomatales (Lu et al. 2009, Paciura et al. 2010, Jankowiak & Kolarik 2010, Jankowiak & Bilanski 2013ab, Romon et al. 2014), hinting at possible generic status for these taxa. Based on this history, we feel the ITS data support placing our new species, at least for now, in Graphilbum.
416 ... Reid & Hausner
Because of our ITS data, we critically compared (TABLE 2) our new species morphologically only to O. rectangulisporium, as described and named in Ohtaka et al. (2006). This comparison clearly separates G. tsugae from the only other described Graphilbum species to which it could