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MYCOTAXON

THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY & NOMENCLATURE

VOLUME 119

JANUARY-MARCH, 2012

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CONSISTING OF I-XII + 511 PAGES INCLUDING FIGURES

ISSN 0093-4666 (PRINT) http://dx.doi.org/10.5248/119.cvr ISSN 2154-8889 (ONLINE)

Iv ... MYCOTAXON 119

MY COTAXON

VOLUME ONE HUNDRED NINETEEN — TABLE OF CONTENTS

COVER SECTION

digi 11s DAR aL AN gan Ui ACN El! PPR ALE CGO NL MER MAD Bo eR AUE ge LO OLR eRe viii

VIC WETS Es, bch eee beac dn MME eee: PR n ohh et RA SET PAIRS Sra ta chk lt ix

SUOTESSHOT: PD FOCCHLUTES SB ors pil trices seo ead gatos ae ice eget eee a ee pag + x

POPPI ENTE Oem Mee en SMe SS) ee ce at Nate eh Me vd tery de cteey Se ate te pe ees xi

RESEARCH ARTICLES

Four new species of Diorygma from India

Bharati Sharma & Pradnya Khadilkar

Gyalidea minuta in Central Europe — new data on its distribution,

ecology, and morphological variation Dariusz Kubiak & Jifi Malicek

New species and record of Sporidesmium from southern China

Jian Ma, Li-Guo Ma, Yi-Dong Zhang, Ji-Wen Xia & Xiu-Guo Zhang

Phlebopus spongiosus sp. nov. (Boletales, Boletinellaceae) with a

sponge-like tissue Nguyen Duc Hoang Pham, Haruki Takahashi,

Toshimitsu Fukiharu, Kiminori Shimizu, Ba Dung Le & Akira Suzuki

Type studies in Helvella (Pezizales) 1

Fidel Landeros, Teresa Iturriaga & Laura Guzman-Davalos

Psilocybe s.s. in Thailand: four new species and a review of previously

recorded species Gaston Guzman, Florencia Ramirez Guillén,

Kevin D. Hyde & Samantha C. Karunarathna

A new Corynesporella species and two first records from China

Li-Guo Ma, Jian Ma, Yi-Dong Zhang & Xiu-Guo Zhang

Queletia mirabilis, a rare gasteroid species in Gran Canaria, Spain

G. Moreno, M.A. Ribes, V. Escobio & A.N. Khalid

New species and a new record of Solicorynespora from southern China

Jian Ma, Li-Guo Ma, Yi-Dong Zhang, Ji-Wen Xia & Xiu-Guo Zhang

Two new species of Endophragmiella from southern China

Jian Ma, Yi-Dong Zhang, Li-Guo Ma,

Rafael F. Castafieda-Ruiz & Xiu-Guo Zhang 103

Alveariospora, a new anamorphic genus from trichomes of

Dimorphandra mollis in Brazil Meiriele da Silva,

Rafael FE. Castafieda-Ruiz, Olinto Liparini Pereira

& Robert Weingart Barreto 109

JANUARY-MARCH 2012... V

Intraornatosporaceae (Gigasporales), a new family with two new genera

and two new species Bruno T. Goto, Gladstone A. Silva,

Daniele M.A. de Assis, Danielle K.A. Silva, Renata G. Souza,

Araeska C.A. Ferreira, Khadija Jobim, Catarina M.A. Mello,

Helder E.E. Vieira, Leonor C. Maia & Fritz Oehl 117

Radulomycetopsis (Agaricomycetes), a new corticioid genus from India

G.S. Dhingra, Priyanka & Jaspreet Kaur 133

First record of Dendrothele mangiferae (Agaricales, Basidiomycota)

from the Neotropics Sergio P. Gorjén 137

Two new fungi from Mexico: Anaseptoidium gen. nov. and

Cylindrosympodium sosae sp. nov. Rafael E Castanteda-Ruiz,

Gabriela Heredia, Rosa M. Arias-Mota,

Marc Stadler, Masatoshi Saikawa & Eric H.C. McKenzie 141

Candelariella, Ochrolechia, Physcia, and Xanthoria species

new to Turkey Kenan Yazici, André Aptroot & Ali Aslan 149

A new species and new combinations and records of Hypotrachyna

and Remototrachyna from Bolivia Adam Flakus,

Pamela Rodriguez Saavedra & Martin Kukwa 157

Mycena moconensis, a new species in section Polyadelphia

from Argentina N. Niveiro, O. Popoff, D. Desjardin & E. Alberté 167

A new species of Fulvifomes (Hymenochaetaceae) from Cambodia

Li-Wei Zhou & Wei-Min Zhang 175

Fungal pathogens of ‘cat's claws’ from Brazil for biocontrol of

Macfadyena unguis-cati Meiriele da Silva,

Robert W. Barreto & Olinto Liparini Pereira 181

Typification of Lecanora expallens and L. expallens var. conizaea

(Lecanorales, Ascomycota) Martin Kukwa 197

The morphology of Ganoderma species with a laccate surface

Mabel Gisela Torres-Torres & Laura Guzman-Davalos 201

Four new polypore species from the western United States

Josef Vlasak, Josef Vlasak Jr. & Leif Ryvarden 217

Studies on Croatian Basidiomycota 2: Marasmiellus milicae sp. nov.

Armin MeSi¢, Zdenko Tkaléec & Vladimir Antonin 233

Zelodactylaria, an interesting new genus from semi-arid

northeast Brazil Alisson Cardoso Rodrigues Da Cruz,

Luis Fernando Pascholati Gusmao, Rafael F. Castafieda-Ruiz,

Marc Stadler & David W. Minter 241

A new species of Ijuhya, I. oenanthicola

Christian Lechat & Michel Hairaud 249

vI ... MYCOTAXON 119

Two new species of Hyphoderma (Agaricomycetes) from India

Priyanka & G.S. Dhingra 255

New records of noteworthy gasteroid fungi from Pakistan

N. Yousaf, A.R. Niazi & A.N. Khalid 261

Pseudocercosporella stellariicola, a new hyphomycete occurring on

Stellaria aquatica Mi-Jeong Park, Ji- Hyun Park & Hyeon-Dong Shin 269

A key to the peltigericolous fungi in Turkey

Mehmet Gokhan Halici, Mehmet Candan & Aysen Tiirk 277

Taxonomy of the Gymnopus inusitatus group and the new G. inusitatus

var. cystidiatus from Hungary Vladimir Antonin,

Péter Finy & Michal TomSovsky 291

New records of Pezizales from Pakistan T. Ashraf & A.N. Khalid 301

A new Hysterostomella species from the Cerrado in Brasilia

National Park C.A. Inacio, R.C. Pereira-Carvalho,

E.S.C. Souza, H.B. Sales & J.C. Dianese 307

Penicillium mallochii and P. guanacastense, two new species

isolated from Costa Rican caterpillars Karol G. Rivera, Joel Diaz,

Felipe Chavarria-Diaz, Maria Garcia, Mirjam Urb,

R. Greg Thorn, Gerry Louis-Seize, Daniel H. Janzen & Keith A. Seifert 315

Two new species of Terriera from Yunnan Province, China

Juan-fang Song, Ling Liu, Yuan-yuan Li & Cheng-Lin Hou 329

Gloeocystidiellum kenyense in Azores and Madeira M. Teresa Telleria,

Margarita Duejfias, Ireneia Melo, Esperanza Beltran-Tejera,

J. Laura Rodriguez-Armas, Isabel Salcedo & Maria P. Martin 337

A new species of Skeletocutis on bamboo (Polyporaceae)

in tropical China Li-Wei Zhou & Wen-Min Qin 345

Paxillus albidulus, P. ammoniavirescens, and P. validus revisited

Else C. Vellinga, Erin P. Blanchard, Stephen Kelly & Marco Contu 351

Two new Heteroconium species and two other forest microfungi

newly recorded from China = Shou-Cai Ren, Jian Ma & Xiu-Guo Zhang 361

Second record of Ramularia hypericicola — collected in Turkey

on a new host Faruk Selcuk, Elsad Hiiseyin & Ali S. Bulbul 369

Lichenological notes 4: A Revision of Acarospora gallica

(Acarosporaceae) Kerry Knudsen & Jana Kocourkova 373

The genus Xylaria in the south of China - 3.

Xylaria atroglobosa sp. nov. Hai-Xia Ma, Larissa Vasilyeva & Yu Li 381

A new name for a Coltricia (Basidiomycota) from India

Juliano M. Baltazar & Rosa Mara B. da Silveira 385

Tuber in China: Tuber microspermum and T. microspiculatum spp. nov.

Li Fan, Jin-Zhong Cao, Zhao-Hui Zheng, & Yu Li 391

JANUARY-MARCH 2012... VII

The first report of Ceriporia lacerata (Phanerochaetaceae, Basidiomycota)

in Korea Yeongseon Jang, Ha Eun Choi,

Young Woon Lim, Jin Sung Lee & Jae-Jin Kim 397

Marasmioid and gymnopoid fungi of the Republic of Korea.

5. Marasmius sect. Hygrometrici Vladimir Antonin, Rhim Ryoo,

Kang-Hyeon Ka & Hyeon-Dong Shin 405

A new hairy species of Leptogium (Collemataceae) from China

Hua-Jie Liu & Shuai Guan 413

Abrachium, a new genus in the Clathraceae, and Itajahya reassessed

Tiara Sousa Cabral, Paulo Marinho,

Bruno Tomio Goto & Juri Goulart Baseia 419

Nomenclatural notes 13. An incorrect neotype designation and

provision for a lectotype and an epitype for Helvella fusca

Fidel Landeros & Richard P. Korf 431

Herpothallon weii, a new lichen from China _ Yu-Liang Cheng, Jing Ning,

Hou-Ping Xu, Lu-Lu Zhang, Hai-Ying Wang & Zun-Tian Zhao 439

Four new records of lecideoid lichens from China Lu-Lu Zhang,

Li-Song Wang, Hai-Ying Wang & Zun-Tian Zhao 445

Puccinia, Uromyces, and Xenodochus species new to Turkey

Sanli Kabaktepe & Zeliha Bahcecioglu 453

Pseudobaeospora deckeri sp. nov., a new agaric from

central California Christian Schwarz 459

A new species of Volvariella from India Gunasekaran Senthilarasu,

Rahul Sharma & Sanjay K. Singh 467

A new species of Pleurophragmium from India

Maria A. D’Souza & D.J. Bhat 477

New combinations in Lactifluus. 2. L. subgenus Gerardii

D. Stubbe, X.-H. Wang & A. Verbeken 483

BOOK REVIEWS AND NOTICES Else C. Vellinga (EDITOR) 487

MyYCOBIOTAS ONLINE 493

ABSTRACTS: Niveiro & Albert6—Checklist of the Argentine Agaricales. 1; Costa

& al.—Checklist of endophytic fungi from tropical regions; Novakova et al.—List

of cultivable microfungi...of the Czech and Slovak Republics; Ghobad-Nejhad

& Hallenberg—Checklist of Iranian non-gilled/non-gasteroid hymenomycetes

(Agaricomycotina); Bahcecioglu & Kabaktepe—Checklist of rust fungi in Turkey

NOMENCLATURE

Minimizing the chaos following the loss of Article 59:

Suggestions for a discussion Walter Gams, Richard A. Humber,

Walter Jaklitsch, Roland Kirschner & Marc Stadler 495

Nomenclatural novelties proposed in volume 119 509

vill ... MYCOTAXON 119

ERRATA FROM PREVIOUS VOLUMES

VOLUME 118

p. 248, Figure caption, line 2

for: 1-2 in LM (holotype), 3 in SEM (K(M) 166454), 4 in SEM (holotype).

read: 5-6 in LM (holotype), 7 in SEM (K(M) 166454), 8 in SEM (holotype).

PUBLICATION DATE FOR VOLUME ONE HUNDRED EIGHTEEN

MYCOTAXON for OCTOBER-DECEMBER, VOLUME 118 (I-1x + 1-465)

was issued on January 5, 2012

JANUARY-MARCH 2012... IX

REVIEWERS — VOLUME ONE HUNDRED NINETEEN

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 volume.

Tetiana V. Andrianova

Andre Aptroot

Ahmet Asan

Juliano Marcon Baltazar

Timothy J. Baroni

Scott T. Bates

Reinhard Berndt

José Luiz Bezerra

Wolfgang von Brackel

Uwe Braun

Mehmet Candan

Paul EF Cannon

Lori M. Carris

Rafael F. Castaneda Ruiz

Ovidiu Constantinescu

Vagner G. Cortez

Pedro Crous

Bao-Kai Cui

Yu-Cheng Dai

Dennis Desjardin

Maria Martha Dios

John A. Elix

Martin Esqueda

Harry C. Evans

Adam Flakus

Jacques Fournier

Francisco C.O. Freire

Walter Gams

Zai-Wei Ge

Tatiana Baptista Gibertoni

Sergio P. Gorjon

A.M. Gugliotta

Shou- Yu Guo

Nils Hallenberg

Roy E. Halling

Richard T. Hanlin

Adriana Hladki

Jan Holec

Kentaro Hosaka

Fabien Charles Cossi

Hountondji

Jae-Seoun Hur

Per M. Jorgensen

Alfredo Justo

Bryce Kendrick

Sevda Kirbag

Roland Kirschner

Kerry Knudsen

Deepika Kumari

Roman Labuda

Karl-Henrik Larsson

Bernardo E. Lechner

James C. Lendemer

De-Wei Li

Cristiano Losi

Guozhong Li

Robert Liicking

Elia C. Martini

Eric H.C. McKenzie

John McNeill

Pierre-Arthur Moreau

Abdul Rehman Niazi

Machiel E. Noordeloos

Lorelei L. Norvell

Clark L. Ovrebo

Zdenék Palice

Shaun R. Pennycook

Jadergudson Pereira

Ronald H. Petersen

Stephen W. Peterson

Olga Vinnere Pettersson

Donald H. Pfister

Michele D. Piercey-

Normore

Christian Printzen

Scott A. Redhead

Peter Roberts

Gerardo Robledo

Andrea Irene Romero

Amy Y. Rossman

Robert A. Samson

Ivan Sanchez-Castro

André Luiz C.M. de A.

Santiago

Trond Schumacher

Keith Seifert

B.M. Sharma

Ewald Sieverding

Gladstone Alves da Silva

Rosa Mara Borges da

Silveira

Harrie J.M. Sipman

Lucyna Sliwa

Dana Tancinova

M. Teresa Telleria

Michal TomSovsky

Larissa Trierveiler-Pereira

Alexander Urban

Joe Vaughan

Else Vellinga

Orvo Vitikainen

Alfredo Vizzini

Roy Watling

A.J.S. Whalley

Alan Wood

Xiu-Guo Zhang

X ... MYCOTAXON 119

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MY COTAXON

http://dx.doi.org/10.5248/119.1

Volume 119, pp. 1-10 January-March 2012

Four new species of Diorygma from India

BHARATI SHARMA & PRADNYA KHADILKAR

Agharkar Research Institute, G.G. Agarkar Road, Pune - 411004, India

CORRESPONDENCE TO *: bharatiomsharma@gmail.com

ABSTRACT—Four new species of the lichen genus Diorygma from India are described — D.

agumbense, D. dandeliense, and D. karnatakense from the Western Ghats and D. rupicola

from Northeast India.

KEYWwoRDS—ascomycetes, taxonomy, Ostropales, Graphidaceae

Introduction

The lichen genus Diorygma Eschw. is widespread in tropical and subtropical

regions and c. 45 species are known from around the world (Kalb et al. 2004,

Archer 2006, 2007, Archer & Elix 2008, Caceres 2007, Sharma & Makhija

2009a,b, Makhija et al. 2009, Tripp et al. 2010), 21 of which are known from

India. The main characters that define Diorygma are lirellate ascocarps with a

heavily pruinose disc, parallel paraphyses with thin lumina, thick gelatinous

walls, and tips that anastomose to form an epithecium, Graphis-type asci, and

hyaline to (rarely) brownish trans-septate or muriform ascospores. Kalb et

al. (2004), who monographed the genus, and confirmed its placement within

Graphidaceae through molecular phylogenetic analysis.

The combination of norstictic and salazinic acids is very rare in Diorygma.

In our previous studies on Graphidaceae from India, we proposed four new

Diorygma species with norstictic and salazinic acids as major compounds:

D. dealbatum B.O. Sharma & Makhija, D. inaequale B.O. Sharma & Makhija,

D. manipurense B.O. Sharma & Makhija, and D. verrucirimosum B.O. Sharma

& Makhija (Sharma & Makhija 2009a). Since then, we have encountered

four additional new species of Diorygma, which are described here. Two —

D. dandeliense and D. karnatakense — contain only norstictic and salazinic

acids, while D. agumbense (with protocetraric acid) and the saxicolous

D. rupicola are also new to science. A key to the identification of Diorygma

species in India is provided.

2 ... Sharma & Khadilkar

Materials & methods

Approximately 80 specimens representing four Diorygma species were collected from

different regions of India Western Ghats —Karnataka, Northeast India —Meghalaya.

Their external morphology was studied using a Zeiss dissecting microscope; sections

for anatomical studies were stained with Lugol's iodine, mounted in lactophenol, and

examined with a Zeiss Axioskop microscope. Images were captured using a Nikon

Coolpix 8400 digital camera.

The specimens were identified by comparison with protologues or descriptions and

photographs of known species in the most recent treatments (Kalb et al. 2004, Archer

2006, 2007, Archer & Elix 2008, Makhija et al. 2009; Sharma & Makhija 2009a,b).

Chemical constituents were identified by thin-layer chromatography using methods

standardized for lichen products (Culberson & Kristinsson 1970, Culberson 1972, White

& James 1985) with the solvent systems benzene-dioxane-acetic acid (180:45:5), hexane-

ethyl ether-formic acid (130:80:20), and toluene-ethyl acetate-formic acid (139:83:8).

All examined specimens are deposited in Ajrekar Mycological Herbarium (AMH).

Taxonomy

Diorygma agumbense B.O. Sharma & Khadilkar, sp. nov. FIG. 1

MycoBank MB 561545

Species nova distincte inter Diorygma specibus ob continentem acidum protocetraricum

sed ascosporis 8-nae differt.

Type: India. Karnataka: Hebri, Agumbe-Udipi road, 16.12.1974, Patwardhan & A.V.

Prabhu (Holotype, AMH 74.3041).

ErymMo_oey: Referring to Agumbe, the collecting locality

Thallus corticolous, pale greenish-grey, rough, cracked, delimited by thin,

black hypothallus. Ascocarps lirelline, concolorous with the thallus, 1-1.5 mm

long, mostly simple, rarely branched, slightly raised to mostly immersed. Disc

narrow, slit-like, rarely + open, covered with white pruina. Exciple convergent,

non-carbonized, non-striate. Hymenium hyaline, laterally KI+ blue, clear, not

inspersed. Asci 8-spored, hyaline, muriform, I+ blue violet, peripheral and

central locules of more or less equal size, regularly arranged in rows, 87-125

x 20-37 um.

CHEMISTRY—Cryptostictic, protocetraric, and stictic acids present (K+

yellow, P+ red).

ECOLOGY & DISTRIBUTION— in semi-evergreen forest in open places on the

roadsides. Known only from the type locality.

REMARKS—Diorygma africanum Kalb et al., D. hololeucum (Mont.) Kalb et

al, and D. pruinosum (Eschw.) Kalb et al., the only other Diorygma species

known to have protocetraric acid as a major compound, differ in having a

distinctly divergent exciple, 1-2-spored asci, and an absence of stictic acid. The

combination of protocetraric and stictic acid complex is rather unique in the

genus and no species previously has been reported with this chemistry.

Diorygma spp. nov. (India) ... 3

Figures 1-4. Habit: 1. Diorygma agumbense (Holotype AMH 74.3041); 2. D. dandeliense (Holotype

AMH 04.276); 3. D. karnatakense (Holotype AMH 04.280); 4. D. rupicola (Holotype AMH 77.779).

Scale bars = 1 mm.

Diorygma erythrellum (Mont.) Kalb et al. and D. minisporum Kalb et al.

both have 8-spored asci but differ from D. agumbense in small ascospores

and chemistry. Diorygma erythrellum has ascospores of 30-65 x 12-20 um

and produces norstictic acid, whereas D. minisporum has transversely septate

ascospores of 17-20 x 5-6.5 um and lacks lichen substances.

Diorygma dandeliense B.O. Sharma & Khadilkar, sp. nov. Fic. 2

MycoBank MB 561546

Similis Diorygma dealbatum sed ascospois majoribus et excipulum convergentum differt.

Type: India, Karnataka, Dandeli forest, 5.10.2004, U.V. Makhija (Holotype, AMH

04.276).

Erymo oey: Referring to Dandeli, the collecting locality

4 ... Sharma & Khadilkar

Thallus corticolous, pale greenish-grey, smooth. Ascocarps_lirelline,

concolorous with the thallus, 0.5-2 mm long, mostly simple, rarely branched,

curved, slightly raised to immersed. Disc narrow, slit-like, rarely + open,

covered with white pruina. Exciple convergent, non-carbonized, non-striate.

Hymenium hyaline, laterally KI+ blue, clear, not inspersed. Asci 1-spored,

hyaline, muriform, I+ blue violet, peripheral and central locules of more or

less equal size, regularly arranged in rows, 125-175 x 27-35 um.

CHEMISTRY—Norstictic and salazinic acids present (K+ yellow forming red

crystals).

ADDITIONAL SPECIMENS EXAMINED—INDIA. KARNATAKA: Dandeli forest, 5.10.2004,

ULV. Makhija, AMH 04.277; B.C. Behera, AMH 04.269.

REMARKS—Diorygma dandeliense shares a similar morphology and chemistry

with D. dealbatum, which can be differentiated by its shorter (105-147 um)

ascospores and a divergent exciple.

The new species is very similar to D. megasporum Kalb et al. in morphology

and ascospore range. However, the asci of D. megasporum contain 2-6

ascospores and the thallus lacks salazinic acid.

Both D. excipuloconvergentum Makhija et al. and D. albocinerascens Makhija

et al., which are morphologically and chemically very close to D. dandeliense,

have long (< 6 mm) lirellae, distinctly striate exciple, and 1-2-spored asci.

Diorygma salvadoriense Kalb et al. also contains norstictic and salazinic

acids but differs from the new species in having distinctly divergent exciple and

a carbonaceous basal layer.

Diorygma karnatakense B.O. Sharma & Khadilkar, sp. nov. FIG. 3

MycoBank MB 561547

Similis Diorygma salvadoriense sed peripheralis et centralis sporae loculis equalis differt.

Type: India, Karnataka, Dandeli forest, 5.10.2004, U.V. Makhija (Holotype, AMH

04.280).

ETYMOLOGy: From the state name, Karnataka.

Thallus corticolous, epiphloeodal, greenish-grey, dull, cracked, smooth.

Ascocarps lirelline, concolorous with the thallus, 0.5-2 mm long, simple,

curved, immersed, ends acute or rounded. Disc narrow, slit-like, covered with

white pruina. Exciple convergent, non-carbonized, non striate. Hymenium

hyaline, laterally KI+ blue, clear, not inspersed. Asci 1-4-spored, hyaline,

muriform, I+ blue violet, peripheral and central locules of more or less equal

size, 130-200 x 32-42 um.

CHEMISTRY— Norstictic and salazinic acids present (K+ yellow, forming red

crystals).

ADDITIONAL SPECIMENS EXAMINED—INDIA. KARNATAKA: Dandeli forest, 5.10.2004,

UV. Makhija, AMH 04.266, 04.441, 04.291.

Diorygma spp. nov. (India) ... 5

REMARKS—Diorygma karnatakense is close to D. salvadoriense, D. reniforme

(Fée) Kalb et al., D. rufopruinosum (A.W. Archer) Kalb et al., D. albocinerascens,

and D. excipuloconvergentum in having large ascospores and in containing

norstictic and salazinic acids as major substances. However, D. salvadoriense,

D. rufopruinosum, and D. reniforme do not produce ascospores with the same-

sized peripheral and central cells. Moreover, D. salvadoriense and D. reniforme

have a distinctly divergent, orange brown to basally carbonized, entire exciple.

Diorygma albocinerascens and D. excipuloconvergentum from Maharashtra,

India, also containing norstictic and salazinic acids and morphologically

similar, differ from D. karnatakense in a distinctly striate exciple.

Diorygma rupicola B.O. Sharma & Makhija, sp. nov. Fic. 4

MycoBank MB 561548

Similis Diorygma hieroglyphicum sed habitus saxicola et excipulum convergentum

differt.

Type: India, Meghalaya, Mawrong, on Cherrapunjee Road, 27.10.1977, P.G. Patwardhan

& M.B. Nagarkar (Holotype, AMH 77.779).

ErymMo.oey: From the Latin rupes, rock, + -cola, dweller, a reference to the species

habit on rock.

Thallus saxicolous, grey-brown colored. Ascocarps concolorous with the

thallus, short, 0.2-0.5 mm long, simple, or irregularly branched, curved,

immersed. Disc narrow, rarely + open, pruinose. Exciple convergent, apically

1-2 striate, carbonized at the tips. Hymenium hyaline, laterally KI+ blue, clear,

not inspersed. Asci 1-2-spored, hyaline, muriform, I+ blue violet, peripheral

and central locules of more or less equal size, regularly arranged in rows, 88-

147 x 25-42 um.

CHEMISTRY—Constictic, cryptostictic, and stictic (all major) and norstictic

(minor) present (K+ yellow).

ECOLOGY & DISTRIBUTION— collected from the tropical and sub-tropical

regions of Meghalaya, Nagaland and Sikkim at ~1500-2000 m.

ADDITIONAL SPECIMENS EXAMINED—INDIA. MeGuHataya: Wiloe, 30.10.77, P.G.

Patwardhan & M.B. Nagarkar, AMH 77.1092; Cherrapunjee Road, 27.10.77, P.G.

Patwardhan & M.B. Nagarkar, AMH 77.816, 77.817; Mohmtheid-Cherrapunjee road,

28.10.77, P.G. Patwardhan & M.B. Nagarkar, AMH 77.1027, 77.1031, 77.1040; Mawrong,

on Cherrapunjee Road, 27.10.77, P.G. Patwardhan & M.B. Nagarkar, AMH 77.2048;

Mawsynram, 30.10.77, P.G. Patwardhan & M.B. Nagarkar, AMH 77.1135; Nagaland,

12 km to Zabuza, 7.11.77, P.G. Patwardhan & M.B. Nagarkar, AMH 77.1468; Sikkim,

near Tangshi view point, Gangtok, 17.10.77, P.G. Patwardhan & M.B. Nagarkar, AMH

77.1947.

REMARKS—Diorygma rupicola is morphologically very close to D. hiero-

glyphicum (Pers.) Staiger & Kalb, which differs in having a corticolous

habit, divergent exciple, and ascospores with a gelatinous sheath. Diorygma

albocinerascens, a corticolous species with similar exciple morphology and

6 ... Sharma & Khadilkar

spore size, can easily be distinguished from the new species in its much narrower

(18-27 um) ascospores and cryptostictic, methylstictic, norstictic and salazinic

acids (all major).

Diorygma monophorum (Nyl.) Kalb et al., resembles D. rupicola in exciple

characters, 1-2-spored asci, large and very broad ascospores (105-165

x 35-60 um), but it differs in having hypostictic, hypoconstictic, and a-

acetytylhypoconstictic acids (major or submajor).

Key to the species of Diorygma from India

LatAscosporesrexcee ding 200 Uris ELE OUAP wwacce! Mane. cienn Se shawn PL risen Be sewn Pee ora Baw ¢ 2

Ib; ‘Ascosporesnotexceeding:-200 punnain lemethie a sa 5 ese ele ldse Fate ee bce te ban ide bee 5

2a. Exciple distinctly divergent; disc exposed; thallus grayish white, powdery, smooth;

ascocarps concolorous with the thallus, 1-3 mm long, emergent; disc yellowish

to brownish; exciple divergent, non-carbonized; asci 1-spored; ascospores

muriform, (135—)168-205 x 54-96 um with 5-7.5 um thick sheath; constictic

(major) and stictic (major) acids present

1 neh be eaal Stee ene D. rufosporum (Patw. & C.R. Kulk.) B.O. Sharma & Makhija

2beExciple distinctly convergentsdise hidden: 3.53 22.2 ap Page Page wae oad ane fea dae 3

3a. Norstictic acid absent; thallus corticolous, greenish white; ascocarps 0.5-9 mm long

and 0.1-0.25 mm broad, simple, rarely branched, disc narrow, pruinose; exciple

convergent, 2—6-striate, blackish brown at apices and dark brown laterally; asci

1-spored; ascospores muriform, 231-244 x 59-76 um, consalazinic, constictic,

cryptostictic, and stictic acids (major) present.... D. megaspermum Makhija et al.

Sbr NOrstichic- ACicupresenl irs Lai cide RE edd dat He cats cents ede Rua +

4a. Thallus corticolous, glaucous, whitish to greenish gray; ascocarps 1-4 mm long

simple to rarely branched, whitish, or concolorous with the thallus; disc slit like

or broader, blackish brown to black, white pruinose when open; exciple poorly

developed, 3-4 indistinct striation, blackish brown at apices, pale yellow to

almost hyaline, convergent; asci 1-4-spored; ascospores muriform, 151-294

x 38-63(-84) um, cryptostictic, norstictic, and stictic acids present

oe ercatly Bane N MatgsNy Bathe, tale he Bue Aad ey D. megistosporum Makhija et al.

4b. Thallus corticolous, smooth to cracked, pale glaucous green to olivaceous buff

or whitish; ascocarps 1-6 mm long simple to rarely branched, immersed to

semi-emergent, concolorous with the thallus; disc white, slit-like, later

moderately open, white pruinose; exciple poorly developed, pale yellow,

indistinctly 2—4-striate, blackish brown at apices; ascospores 1-2-4/ascus,

147-273 x 34-67 um; constictic, cryptostictic, methylstictic, norstictic, and

SalaZiIiG ACIDS PTESEME epson Sesion eras wR chara dees D. excipuloconvergentum

Dd -rAScUspores exceéedine POON AN TENSth 22 #t.0.) 1th! tla w8 et Luma! UR oe Te BM on I 2 6

5b. Ascospores not exceeding 100 um in length ............... 0c eee eee 20

Gah Hallie Saxe OlGUsa to vance Bens, Veracths hecnchs Vevache Mara ¥a tered Maavah Means Paar ey 3 7

Diorygma spp. nov. (India) ... 7

7a. Thallus saxicolous, grey-brown colored; ascocarps concolorous with the thallus,

short, 0.2-0.5 mm long, simple, or irregularly branched; exciple convergent,

apically 1-2 striate, carbonized at the tips; asci 1-2-spored, muriform, 88-147

x 25-42 um; constictic, cryptostictic, stictic (major), and norstictic (minor)

PIERS TI let as ante! Peeve elt con tel Neva slh era WeeN cre Leh cn Piers tL Micra Mita Midas D. rupicola

7b. Thallus saxicolous, greenish to pale gray, rough; ascocarps concolorous, curved,

short, round to elongate, simple to branched, very closely arranged, immersed

to slightly emergent; disc narrow to broad; exciple divergent, non-carbonized,

poorly developed, yellowish brown laterally and basally, 2-3 indistinct striate

at the apical region; asci 1-6-spored; ascospores muriform, 143-172 x 29-34

um; constictic, norstictic (trace) and stictic acids present

RTT SA SOR gn en eat Pe veer ae Ear) D. saxicola B.O. Sharma & Makhija

Sas Protocetratic acid: present: Fics iii eae hee eare ogre ls as Ce Pe ces. poe Leen 9

SHAProtocewraricacicl AD Serit tweety Bema t Mine t ly Benen ey Uatuctyls, Bitnadls Bain Ale A we teatabe de best 10

9a. Thallus corticolous, whitish green; ascocarps concolorous with the thallus,

circular, oval, short curved, 1-3 mm long; disc grayish, broad; exciple

divergent, non-carbonized to carbonization sometimes restricted to the

basal corners; asci 1-spored; ascospores muriform, 147-160 x 34-42 um;

PEOLOCE ATIC ACI mes rs ret, men Ts eee ee D. pruinosum

9b. Thallus corticolous, pale greenish-grey; ascocarps lirelline, concolorous with the

thallus, 1-1.5 mm long, mostly simple, rarely branched; disc narrow, covered

with white pruina; exciple convergent, non striate; asci 8-spored, muriform,

87.5-125 x 20-37.5 um; cryptostictic, protocetraric and stictic acids present

willy seach vey led oe Pe eh dF bey eee Fh See elu PG lege Pe ee ee D. agumbense

LOA pARSEL SSS POLE St inet tot tebad es Saba hep aba dcr Sebel tebe geben teh Aabag tee poked pekabsdhe pdakgy he pet 11

LOb-ssciimoke Tiare SPOLet He. iste. Milde, ABE es At eh deat as ntl ita ab gel e 12

11a. Thallus corticolous; exciple convergent; ascospores 1-8/ascus, 85-197 x 29-71

lm; constictic, stictic, and cryptostictic acids present.......... D. megasporum

11b. Thallus corticolous, ashy white; ascocarps white, straight to curved, branched,

0.5-2.5 mm long, immersed; disc 0.3-0.6 mm broad; exciple convergent to

divergent, indistinctly striate; asci 1-8-spored; ascospores 75-145 x 24-33.6 um;

norstictic and stictic acids present

fy 2 Pedy ee ies 2 oe D. subalbatum (Patw. & Makhija) B.O. Sharma & Makhija

l2asExciple distinctly convergent -dischidden: +2. -...45 20 tb need hh niet nec mee dee 13

12bExcipledistinctly-diversetitzdise exposed mas ccns..caecie cael toed seco + ole 16

1B Fer oC) Tel Leics 6101 8 {0 UF AR LOLS Poy AO Td tS ee ee Ee Tt eo RS nh 14

13h. sxsch more thanslespOretiae se, ork, sort Atee ea ies hacer Ad sGrocth sy «ices aro 15

14a. Thallus corticolous, pale greenish-grey; ascocarps lirelline, concolorous with

the thallus, 0.5-2 mm long; disc narrow, covered with white pruina; exciple

convergent, non striate; asci 1-spored, muriform, 125-175 x 27.5-35 um;

norstictic and salazimcacids present. hs u keh ley ht A x D. dandeliense

14b. Thallus corticolous, greenish grey with red tinge; ascocarps long flexuous,

branched, curved, up to 9 mm long; disc slightly open, pale brown; exciple

8 ... Sharma & Khadilkar

convergent to divergent, non-carbonized; asci 1-spored; ascospores muriform,

105-113 x 33-42 um; norstictic acid present

peg Hheed chard Qthreg diner a dibierg D. longilirellatum B.O. Sharma & Makhija

15a.Thallus corticolous, whitish gray, finely cracked; ascocarps 0.5-6 mm long simple,

dendroidly branched, immersed, straight or curved, concolorous with the

thallus; disc narrow, slit-like, light brown, slightly pruinose; exciple convergent,

2-3-striate, blackish brown at apices, asci 1-2-spored; ascospores 135-150 x

18-27 um; cryptostictic, methylstictic, norstictic, and salazinic acids present

Pia Wt cadandl patted, sara dac Marmara Masmatese maaan seman Maan Marat eae D. albocinerascens

15b. Thallus corticolous, greenish-grey, dull; ascocarps concolorous with the thallus,

0.5-2 mm long, simple, curved, immersed; disc narrow, covered with white

pruina; exciple convergent, non striate; asci 1-4-spored, muriform, 130-200 x

32.5-42.5 «um; norstictic and salazinic acids present .......... D. karnatakense

loa Norsticticand salamimic-acids;preserits..x.... 8) ke 0s, att Re, ath Ray ate ay ena ene 17

LG by-Sallaz tite aid AS OIMA Etec ote eben rhe suhag ath fe bwa bs Sebald che Abad tabs Akal tc ube ch fo tou-dch foal 18

17a. Thallus corticolous, glaucous grey, deeply cracked, distinctly verrucose; ascocarps

crowded, concolorous with the thallus, irregular, short, 1-2 mm long; disc

narrow to wide, sunken, 0.2-0.4 mm wide, reddish brown covered with

white pruina; exciple, divergent, uncarbonized, base distinctly orange brown;

asci 1-spored, muriform, 97-126 x 25-33 um; norstictic and salazinic acids

JORCSCI Epa. cr ead torts tree ten Bee Me EON ha Sea ee D. verrucirimosum

17b. Thallus corticolous, white with brown tinge, smooth, thick; ascocarps rounded,

lirellate, concolorous with the thallus, immersed, irregular, simple to branched,

curved, 0.5-2.5 mm long; disc narrow to broad, 0.2-0.7 mm; exciple divergent,

uncarbonized, laterally poorly developed, base distinctly orange brown; asci

1-spored, ascospores muriform, 105-147 x 33-37 um; norstictic and salazinic

AGIA e.g fo ad dita. od Aihie ob dais oo dase dace dchareG achece Facsre hd the-o 4 D. dealbatum

18a. Peripheral spore locules distinctly smaller than the central ones. Thallus

corticolous, rough; ascocarps 1-5 mm long, raised from the thallus; exciple

non-carbonized, yellowish orange; asci 1-spored; ascospores muriform,

100-130 x 44-48 um; norstictic and con-norstictic acids present

wri: Lope ake Sy Eat OR ely ae ica ts oar pelate oe pe ae 5 D. tuberculosa (Stirt.) Kalb et al.

18b. Peripheral and central spore locules of equal size ......... 2... cece ee eee ee 19

19a. Thallus corticolous, greenish with whitish tinge, smooth; ascocarps off white

to concolorous with the thallus, immersed 1-4 mm long; disc broad, brown,

0.2-0.4 broad, pruinose; exciple non-carbonized, poorly developed, yellowish

brown laterally and basally not striate; asci 1-spored; ascospores muriform,

80-131 x 42-52 um, with a 5-15 um thick sheath; stictic, constictic and

norstictic acids present? wm. «£5 tss canoe saws Pe Fae Fa D. hieroglyphicum

19b. Thallus corticolous, creamy white, grayish; ascocarps 0.4-4 mm long, round to

irregular, branched, immersed to slightly raised; disc brownish grey covered by

yellowish cream pruina; exciple divergent, non-carbonized, orange yellow at the

Diorygma spp. nov. (India) ... 9

base; asci 1(-2)-spored; ascospores 84-134.4 x 29.4-42 um, with a 2.3-5 um

sheath; constictic acid (trace), norstictic, and stictic acid (major) present.

NE Pe MID. LNG aces orcas gee gia D. junghuhnii (Mont. & Bosch) Kalb et al.

2a. SEIU S OTE og lh. ng shang g dihareg dibareg Aineerg dined y deneerg & Deen een theres neers 21

20D A XSCIMOne, Una Wes POLE Ce Sete ai le tad ena ne Te ee Ds Res RM Se oa Se 22,

21a. Thallus corticolous, glaucous green with a yellow and or red tinge, uneven;

ascocarps 1-6 mm long, simple to irregularly branched; disc brown, 0.2-0.3

mm broad; exciple non-striate, divergent, non carbonized, distinctly orange at

the base; asci 1-spored; ascospores muriform, 79-96 x 29-33 um; norstictic and

salavinieacidsspresenits. yes uhh ele ne eae Pee BaP eae Le D. inaequale

21b. Thallus corticolous, grayish white, smooth to rough, deeply cracked and

appearing areolate; ascocarps concolorous with the thallus, 1-1.5 mm long,

simple to branched, grouped, immersed; exciple hyaline, poorly developed, 2-3

striate, pale brown laterally, blackish brown at apices, convergent; asci 1-spored;

ascospores muriform; 57-96 x 24-36 um; consalazinic, constictic, cryptostictic,

and stictic acids (major); Maharashtra state, [AMH 04.13]... D. “patwardhanii”

274, Norsticticiand salazinicacid presenti, 4... 4.1 kl oR ec he gas cit 23

22 SalaZziiii G-ACiLADSENly gles sre sctg dye -sclon discon den pcb tigen dr pemadie ected ype gaia 24

23a. Thallus corticolous, grayish, smooth to rough; ascocarps 1-2 mm long, simple,

semi-emergent, concolorous with the thallus; exciple 2-3 striate, pale brown

laterally, blackish brown at apices, convergent; asci 1-2-spored; ascospores

muriform, 75-99 x 24-30 um; norstictic, salazinic, and methylstictic acids

PIES CIN a wy Ho tog traracteg drprseien srpesatag ding von drngetea dh et D. panchganiense Makhija et al.

23b.Thallus corticolous, brownish white, + smooth to rough; ascocarps concolorous

with the thallus, 0.2-0.3 mm long mostly simple to rarely branched; disc brown,

0.4-0.5 mm broad; exciple 2-5 striate, apically dark brown, divergent; asci 2-4

spored; ascospores muriform, 76-92 x 21-25 um; Norstictic and salazinic acids

POLSON 0% cathy, welelinths, Bettina, Bocwenth, Bocwe thy Wee hey Belt Meee Bs ah dma D. manipurense

24a. Thallus corticolous, glaucous green, smooth to cracked; ascocarps 3-7 mm long

simple to branched, immersed to emergent; disc reddish brown to blackish

brown; exciple 2-5 striate, orange brown, carbonized at apices, convergent;

asci 8-spored; ascospores muriform, 46-63 x 17-25 um; stictic acid (major);

Maharashtra states, [AMIE F224) xls cceel gush, een BBs ows os ss D. “microsporum”

24b. Thallus corticolous, greenish white, smooth to cracked; ascocarps 0.5-3 mm

long simple to branched, concolorous with the thallus, fissure-like; exciple 2-3-

striate, pale brown to blackish-brown at apices, convergent, asci 4—8-spored;

ascospores muriform, 66-99 x 12-36 um; constictic, cryptostictic, and stictic

AcIdS Present. dias Pitae esate #utuas Ata eee Cle D. albovirescens Makhija et al.

(Note: species in quotations marks — D. “patwardhanii” and D. “microsporum” — are

recorded by Makhija et al. (2009) and are probably new to science, but are not formally

described due to scanty material.)

10 ... Sharma & Khadilkar

Acknowledgments

We are grateful to the Department of Science and Technology, Government of India,

New Delhi for the financial support. The authors are grateful to A. Aptroot, J. Lendemer,

and R. Liicking for reviewing the manuscript, critical revision, and suggestions to

improve the quality of the manuscript.

Literature cited

Archer AW. 2006. The lichen family Graphidaceae in Australia. Bibliotheca Lichenologica 94:

1-19}.

Archer AW. 2007. Key and checklist for the lichen family Graphidaceae (lichenized Ascomycota) in

the Solomon Islands. Systematics & Biodiversity 5(1): 9-22.

http://dx.doi.org/110.1017/S1477200006002040

Archer AW, Elix JA. 2008. Three new species in the Australian Graphidaceae (lichenized

Ascomycota). Australasian Lichenology 63: 26-29.

Caceres MES. 2007. Corticolous crustose and microfoliose lichens of northeastern Brazil. Libri

Botanici 22: 1-168.

Culberson CE 1972. Improved conditions and new data for the identification of lichen products by

a standardized thin-layer chromatographic method. Journal of Chromatography 72: 113-125.

http://dx.doi.org/1/10.1016/0021-9673(72)80013-X

Culberson CF, Kristinsson H. 1970. A standardized method for the identification of lichen products.

Journal of Chromatography 46: 85-93. doi:10.1016/S0021-9673(00)83967-9

Kalb K, Staiger B, Elix JA. 2004. A monograph of the lichen genus Diorygma — a first attempt. Symb.

Bot. Ups. 34(1): 133-181.

Makhija U, Chitale G, Sharma BO. 2009. New species and new records of Diorygma (Graphidaceae)

from India: species with convergent exciples. Mycotaxon 109: 379-392.

http://dx.doi.org/110.5248/109.379

Sharma BO, Makhija U. 2009a. Four new species in the lichen genus Diorygma. Mycotaxon 107:

87-94. http://dx.doi.org/10.5248/107.87

Sharma BO, Makhija U. 2009b. New species and new reports of Diorygma (lichenized Ascomycotina,

Graphidaceae) from India. Mycotaxon 109: 209-217. http://dx.doi.org/10.5248/109.209

Tripp EA, Lendemer JC, Harris RC. 2010. Resolving the genus Graphina Mill. Arg. in North

America: new species, new combinations, and treatments for Acanthothecis, Carbacanthograhis,

and Diorygma. Lichenologist 42(1): 55-71. http://dx.doi.org/110.1017/S0024282909990296

White FJ, James PW. 1985. A new guide to microchemical techniques for the identification of lichen

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MY COTAXON

http://dx.doi.org/10.5248/119.11

Volume 119, pp. 11-16 January-March 2012

Gyalidea minuta in Central Europe - new data on its distribution,

ecology, and morphological variation

DARIUSZ KUBIAK? & JIRi MALICEK?

‘Department of Mycology, Warmia and Mazury University in Olsztyn

Oczapowskiego 1A, PL-10-719 Olsztyn, Poland

*Department of Botany, Faculty of Science, Charles University

Bendtska 2, CZ-128 01 Praha 2, Czech Republic

* CORRESPONDENCE TO: darkub@uwm.edu.pl

ABSTRACT — New localities of Gyalidea minuta in Central Europe are reported. The

distribution, ecology, and morphological variation of the species are discussed and

differentiating characters presented.

Key worps — lichenized fungi, lichens, Solorinellaceae, Poland, Czech Republic

Introduction

The genus Gyalidea was resurrected by Vézda (1966) and by Vézda & Poelt

(1991). Currently included in the Solorinellaceae (Baloch et al. 2010). Gyalidea

was earlier referred to the Asterothyriaceae and Gomphillaceae (see Henssen

& Liicking 2002, Lumbsch & Huhndorf 2010). The genus is characterized by

a crustose thallus, zeorine apothecia with a mostly well-developed thalline

margin, an Ostropales-type ascus, a non-amyloid hymenium with simple

to sparsely branched and septate paraphyses, and hyaline, muriform (or

transversally septate) ascospores (Vézda 1966; Vézda & Poelt 1991; Henssen &

Liicking 2002; Aptroot & Licking 2002). The thallus is usually inconspicuous

and the apothecia small. Gyalidea is a cosmopolitan genus with over 40 species

(Vézda & Poelt 1991; Kirk & Cooper 2009), many of which have been described

in the last two decades (see Lumbsch et al. 2009). They are rather rare lichens,

some known only from the type material or very few collections. Most species

grow on soil, rocks, mosses, or plant debris, and only a few occur on tree bark. In

Europe there are only two epiphytic species, G. minuta (van den Boom & Vézda

1995) and the recently described G. fruticola (Svensson & Thor 2007). Until

recently, G. minuta had been found only in three localities in southwestern and

12 ... Kubiak & Maliéek

western Europe. Here we report the first record of G. minuta in Central Europe

and discuss its distribution, ecology, and morphological variation.

Materials & methods

The specimens were morphologically examined by standard microscopic techniques.

Hand-cut sections and squash preparations were examined in water, a 10% aqueous

solution of KOH, and Lugol's solution. Measurements of well-developed free ascospores

lying outside the asci were measured in water at x1000 magnification. The description

below is based on the isotype and newly collected specimens.

The taxon

Gyalidea minuta Van den Boom & Vézda, Mycotaxon 54: 423. 1995. Fic. 1

Tye: Portugal, Algarve (holotype, herb. van den Boom 14875 [not seen]; isotype, herb.

A. Vézda (PRA-V-05556!)).

THALLUuS epiphloedal or partly endophloedal, corticolous, scattered among

substrate wrinkles, grayish green, without visible prothallus. APOTHECIA sessile,

hyaline with a brownish tinge, translucent when wet, 0.15-0.2 mm diam., 0.1

mm tall (0.2-0.4 x 0.1-0.15 mm in water preparation). HyMENIuM colorless,

45-75 um tall. Paraphyses simple, not broadening towards the tips, 1.5-2.0 um

in diam., indistinctly septate. Asci cylindrical-clavate, wall slightly thickened

at apex, 8-spored, 28-40 x 7-9 um. Ascospores ellipsoid, with rounded to

attenuate ends, muriform, with 3-5 transverse septa, and 1-3(-4) longitudinal

septa, (9.5-)10-17(-20) x 4.5-7.5(-9) um. PYcnipiA not observed. CHEMISTRY

not tested by TLC.

Eco.Loey -— In western Europe the species has been reported from the bark

of Alnus glutinosa and Fraxinus sp. trees in shady, humid forests and among

shrubs (van den Boom & Vézda 1995, Sparrius et al. 2002).

In Poland, it has been found only in forest conditions. One locality is a

narrow strip of land (ca. 50 m wide and about 2 km long) adjacent to a lake

with an anthropogenic forest community of an unusual structure. The tree

layer consists of middle-aged pine and the shrub layer of Berberis sp., Crataegus

sp., Euonymus europaeus, Rhamnus cathartica, Sambucus nigra in which the

common buckthorn (R. cathartica) dominates. The dense shrub-layer and the

proximity to the lake give rise to highly humid sheltered conditions. Gyalidea

minuta grows here on the bark of the buckthorns only and is intermixed with

lichens common to this area (Anisomeridium polypori (Ellis & Everh.) M.E.

Barr, Bacidina sulphurella (Samp.) M. Hauck & V. Wirth, Coenogonium pineti

(Schrad. ex Ach.) Licking & Lumbsch, Lepraria incana (L.) Ach., L. lobificans

auct., Melanelixia fuliginosa (Fr. ex Duby) O. Blanco et al., Micarea micrococca

(K6rb.) Gams ex Coppins, Psoroglaena abscondita (Coppins & Vézda) Hafellner

& Turk, Parmelia sulcata Taylor).

Gyalidea minuta in Central Europe... 13

FIGURE 1. Habit of Gyalidea minuta. Scale bar = 0.5 mm.

The second locality is in a pine forest planted in soils that naturally favor

oak-linden-hornbeam forest. In this habitat, G. minuta grows at the bottom of

Acer pseudoplatanus trunks mixed with abundant Bacidina sulphurella. Other

lichen species include Lepraria incana, L. elobata Tonsberg, and Parmelia

sulcata.

In the Czech Republic G. minuta has been collected in a shady, damp forest

of mostly Alnus glutinosa and Fraxinus excelsior within part of an old spruce-

beech forest reserve. Here the species grows abundantly around bark rifts of

Fraxinus trunks accompanied by Lepraria sp., Lecanora pulicaris (Pers.) Ach.,

and Phlyctis argena (Ach.) Flot.

GEOGRAPHICAL DISTRIBUTION (FIG. 2) -Known only from Europe,

G. minuta had been previously reported from Portugal (van den Boom & Vézda

1995), northern France, and Belgium (Sparrius et al. 2002).

SPECIMENS EXAMINED: POLAND. WaARMIA-MasuRIA: OLSZTYN LAKE DISTRICT,

Olsztyn, Lupsztych settlement, near lake Ukiel (53°46'50"N 20°24'20"E), 10.VII. &

10.X1.2001, leg. D. Kubiak (OLTC L-1425, PRA-V-03137, PRA-V-03138); 15.[X.2002,

leg D. Kubiak (OLTC L-1976); 26.IV.2005, leg D. Kubiak (OLTC L-2879); 7.1I.2009,

leg. D. Kubiak (OLTC L-3113); Olsztyn, Dajtki settlement, near lake Ukiel (53°46'53"N

20°25'20"E): 6.V1.2009, leg. D. Kubiak (OLTC L-3190). CZECH REPUBLIC. EASTERN

14 ... Kubiak & Maligek

FIGURE 2. Distribution of Gyalidea minuta.

@ - previously published localities, A - new localities.

Bouemia: Zelezné hory Mts, Nasavrky—Horni Bradlo, Polom Nature Reserve (ca.

49°47'34"N 15°45'16"E), 24.1X.2009, leg. J. Malicek, J. Halda & A. Miiller (herb. J.

Maliéek 2018).

ADDITIONAL SPECIMENS EXAMINED: PORTUGAL: ALGARVE: Serra de Monchique,

road 267 to S. Marcos da Serra (Alferce), 1.8 km E of crossing to Monchique (37°19.0'N

8°32.3'W), 28.VI1I.1993, leg. Pv.d. Boom (PRA-V-05556, isotype).

Comments - This constitutes the first report of Gyalidea minuta from Central

Europe. The first specimen from Poland, found in 2001, differs from the type

in apothecial shape and in the number and arrangement of septa. A. Vézda

(pers. comm.) suggested that these differences were great enough to support

a new species, but a final taxonomic decision was postponed in view of the

small specimen size. No new material was found until 2009 when the first

author found a larger population within 2 km of the first locality. In the same

year the species was also found in the Czech Republic. Analysis of the rather

rich material from both countries showed that all specimens represent a single

Gyalidea minuta in Central Europe... 15

species, Gyalidea minuta. Detailed measurements have shown that the Central

European specimens deviate slightly: the ascospores are slightly smaller with

3-4 transverse septa and the apothecia are generally larger and more flattened

(TABLE 1). Furthermore, the exciples of dead apothecia are not persistent, as

in the isotype specimen. These differences are probably due to the fact that

the previously collected specimens are small and poorly developed and do not

reflect the full variability of the species.

TABLE 1. Gyalidea minuta measurements: comparison between western and

central European specimens.

WESTERN EUROPEAN CENTRAL EUROPEAN

APOTHECIA:

Diam. (mm) in preparation 0.20-0.30 0.25-0.40

HYMENIUM:

Height (ttm) 50-60(-65) 45-75

ASCOSPORES:

Length (um) (9.5-)12-17(-20) 10-16

Width (um) 5-9 4.5-7.5

Transverse septa 3-5 3-4

Longitudinal septa 1-3 0-3(-4)

It is worth noting that despite the relatively detailed and long-term

observations in Poland, there are very few records of G. minuta. This may

indicate that the species has an ephemeral nature, although it is also easily

overlooked in the field.

Despite its variability, Gyalidea minuta is quite distinctive due to its

occurrence on the bark of trees and shrubs, its very small, pale and almost

translucent apothecia, and the barely visible thallus. The only other epiphytic

Gyalidea species — the recently described G. fruticola (Svensson & Thor 2007),

which also occurs on the bark of shrubs — differs in its larger, whitish grey

to light yellowish brown apothecia and considerably longer, more septate

ascospores. Additionally, G. fruticola has a well-developed true exciple that

almost encloses the disc. In wet conditions, G. minuta resembles Coenogonium

pineti in the field, but the microscopic features differ markedly. Rich photo

documentation of the isotype and Central European collections is available on

http://www. jjh.cz/foto/.

Acknowledgments

We are grateful to the late Antonin Vézda for his valuable taxonomic comments,

Pieter P.G. van den Boom for the examination of our specimens, and Zdenék Palice

and Ludmila Kirschnerova for making the isotype specimen from PRA herbarium

available. We are also grateful to Zdenék Palice (Praha) and Harrie Sipman (Berlin) for

very helpful comments and peer-reviewing the paper, and to Mark Leonard for critical

reading and English correction.

16 ... Kubiak & Maliéek

Literature cited

Aptroot A, Liicking R. 2002. Proposal to conserve Gyalidea (lichenized fungi: Asterothyriaceae,

Ostropales) against an additional name, Solorinella. Taxon 51: 565.

http://dx.doi.org/10.2307/1554877

Baloch E, Liicking R, Lumbsch HT, Wedin M. 2010. Major clades and phylogenetic relationships

between lichenized and non-lichenized lineages in Ostropales (Ascomycota: Lecanoromycetes).

Taxon 59: 1483-1494.

Henssen A, Liicking R. 2002. Morphology, anatomy, and ontogeny in the Asterothyriaceae

(Ascomycota: Ostropales), a misunderstood group of lichenized fungi. Ann. Bot. Fenn. 39:

273-299.

Kirk PM, Cooper J. 2009. CABI Bioscience database: Index fungorum.

[http://www.indexfungorum.org/Names/Names.asp (viewed online on 12 April 2011)]

Lumbsch HT, Papong A, Naikatini A. 2009. A new terricolous species of Gyalidea (Gomphillaceae,

Ascomycota) from Fiji. Nova Hedwigia 88: 111-116.

http://dx.doi.org/10.1127/0029-5035/2009/0088-0111

Lumbsch HT, Huhndorf SH. 2010. Myconet Volume 14. Part One. Outline of Ascomycota—2009.

Fieldiana, Life and Earth Sciences 1: 1-42. http://dx.doi.org/10.3158/1557.1

Sparrius LB, Diederich P, Signoret J, Sérusiaux E. 2002. The lichen flora of the Boulonnais (France,

Pas-de-Calais). Belg. J. Bot. 135: 50-75.

Svensson M, Thor G. 2007. Gyalidea fruticola, a new corticolous lichen from Europe. Lichenologist

39: 335-338. http://dx.doi.org/10.1017/S0024282907006743

Van den Boom PPG, Vézda A. 1995. A new species and a new variety of the lichen genus Gyalidea

from Western Europe. Mycotaxon 54: 421-426.

Vézda A, Poelt J. 1991. Die Flechtengattung Gyalidea Lett. ex Vézda (Solorinellaceae). Eine Ubersicht

mit Bestimmungsschliissel. Nova Hedwigia 53: 99-113.

Vézda A. 1966. Flechtensystematische Studien IV. Die Gattung Gyalidea Lett. Folia Geobot.

Phytotax. 1: 311-340.

MYCOTAXON

http://dx.doi.org/10.5248/119.17

Volume 119, pp. 17-25 January-March 2012

New species and record of Sporidesmium from southern China

JIAN Ma, Li-Guo Ma, Y1-DONG ZHANG, JI-WEN XIA & XIU-GUO ZHANG

Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China

*CORRESPONDENCE TO: zhxg@sdau.edu.cn, sdau613@163.com

ABSTRACT — Four species of Sporidesmium were collected from decaying twigs in tropical

and subtropical forests in southern China. Sporidesmium liquidambaris sp. nov. on

Liquidambar formosana, S. antidesmatis sp. nov. on Antidesma ghaesembilla, and S. machili

sp. nov. on Machilus chinensis, are described, illustrated, and compared with closely related

taxa. Sporidesmium takashii is recorded as new for the Chinese mycota.

Key worps — conidial fungi, taxonomy

Introduction

The genus Sporidesmium (anamorphic Pleosporales) was established by

Link (1809) with S. atrum Link as type species. Ellis (1958, 1971) defined the

genus as having integrated, terminal, monoblastic, determinate or percurrent

conidiogenous cells on distinctive, unbranched conidiophores, and acrogenous,

solitary, transversely septate or distoseptate conidia. Under this generic concept,

about 250 species are included in this genus. However, Sutton & Hodges

(1979) and Hughes (1979) felt that Sporidesmium is heterogeneous, an opinion

confirmed by Shenoy et al. (2006) who showed the polyphyletic nature of

Sporidesmium and its allies. Kirk (1982) proposed the new genus Sporidesmiella

P.M. Kirk for those species with cylindrical to cuneiform or obovoid, distoseptate

conidia. Subramanian (1992) refined the generic concept of Sporidesmium in

a stricter sense as ‘conidiophores simple, determinate or irregular extending

proliferations, conidia solitary, gangliar, acrogenous and euseptate, and proposed

several novel anamorphic genera, including Ellisembia Subram., Penzigomyces

Subram., Repetophragma Subram., and Stanjehughesia Subram., based on

the absence/presence of conidiophores, type of conidiophore extension, and

conidial septation (euseptate/distoseptate). Hernandez-Guitiérrez & Sutton

(1997) and Shoemaker & Hambleton (2001) further modified this approach

by proposing Imimyces A. Hern. Gut. & B. Sutton, Linkosia A. Hern. Gut. &

18 ... Ma & al.

B. Sutton, and Imicles Shoemaker & Hambl. However, Wu & Zhuang (2005)

merged Penzigomyces into Sporidesmium and Imicles into Ellisembia, thereby

expanding the generic concepts of Sporidesmium (euseptate) and Ellisembia

(distoseptate) to include fungi with typically lageniform, doliiform, or nodose,

percurrently extending conidiophores.

Sporidesmium is worldwide in distribution, usually found as a saprobe on

rotten wood, dead branches, and decaying leaves of various plant species. (Ellis

1958, 1971, 1976, Wu & Zhuang 2005). During ongoing surveys of conidial

fungi associated with woody debris in tropical and subtropical forests of

southern China, four species clearly related to Sporidesmium sensu stricto were

collected from decaying twigs. Three represent species new to science while the

fourth is a new record from China.

Sporidesmium liquidambaris Jian Ma & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 563262

Conidiophora macronemata, nonramosa, 11-72 x 5.5-9.5 um. Cellulae conidiogenae

monoblasticae, integratae, terminales, per usque ad 7 extensiones lageniformes vel

ampulliformes percurrentes. Conidia solitaria, obclavata, verruculosa, 11-26-euseptata,

110-165 x 20-24 um, apicem versus ad 3-4.5 um attenuata, basi truncata 4-5 am lata.

Type: China, Fujian Province: Mount Wuyi, on decaying twigs of Liquidambar

formosana Hance (Hamamelidaceae), 16 Aug 2009, J. Ma, (Holotype HSAUP H5059;

isotype HMAS 146156).

EryMo_oey: in reference to the host genus, Liquidambar.

Anamorphic fungi. COLONIES on natural substrate effuse, brown to dark

brown, hairy. Mycelium partly superficial, partly immersed in the substratum,

composed of branched, septate, pale brown, smooth-walled hyphae, 1.5-3 um

thick. CONIDIOPHORES distinct, single or in groups, erect, unbranched, straight

or flexuous, cylindrical, dark brown to black, smooth, septate, 11-72 x 5.5-9.5

lum. CONIDIOGENOUS CELLS monoblastic, integrated, terminal, brown to dark

brown, smooth, 8.5-11.5 x 5-7 um, with up to 7 lageniform or ampulliform

percurrent extensions. Conidial secession schizolytic. Conrp14 holoblastic,

acrogenous, solitary, straight or curved, obclavate, verruculose, brown to dark

brown, apical cells pale brown or subhyaline, 11-26-euseptate, 110-165 um

long, 20-24 um thick in the broadest part, tapering to 3-4.5 um near the apex,

4-5 um wide at the truncate base.

ComMENTs —-Among the known Sporidesmium species, S. liquidambaris

resembles S. ghanaense M.B. Ellis (Ellis 1958), S. matsutakashii Subram.

(Subramanian 1992), and S. tengii W.P. Wu (Wu & Zhuang 2005) in conidial

shape. However, S. liquidambaris is distinguished from S. ghanaense (conidia

31-53 x 10-14 um, 5-9-euseptate) and S. tengii (conidia 45-50 x 7-8.5 um,

8-euseptate) by its larger conidia with more septa. It differs from S. matsutakashii

Sporidesmium spp. nov. (China) ... 19

A B

Fic. 1. Sporidesmium liquidambaris. A. Conidiophores with conidia. B. Conidia.

(conidia 9.5-14 um wide) by its wider conidia that lack an apical mucilaginous

appendage. In addition, the conidia of S. liquidambaris are verruculose while

those of S. ghanaense, S. matsutakashii, and S. tengii are smooth.

Sporidesmium antidesmatis Jian Ma & X.G. Zhang, sp. nov. FIG. 2

MycoBank MB 563263

Conidiophora macronemata, nonramosa, 17-70 x 6-7.5 um. Cellulae conidiogenae

monoblasticae, integratae, terminales, determinatae, cylindrica. Conidia solitaria,

obclavata, laevia, brunnea, 8-11-euseptata, 108-150 x 9-11 um, apice rotundata, ad 3-4

um lata, et muco subgloboso usque 12-24 um diam. tecta, ad basim 4.5-6 um lata.

Type: China, Hainan Province: Bawangling, on decaying twigs of Antidesma ghaesembilla

Gaertn. (Euphorbiaceae), 11 Dec 2010, J. Ma, (Holotype HSAUP H5254; isotype HMAS

146157).

ETryMoLoey: in reference to the host genus, Antidesma.

Anamorphic fungi. COLONIES on natural substrate effuse, brown to dark

brown, hairy. Mycelium partly superficial, partly immersed in the substratum,

composed of branched, septate, pale brown, smooth-walled hyphae, 1-2.5 um

20 ... Ma & al.

Fic. 2. Sporidesmium antidesmatis. A. Conidiophores with conidia. B—C. Conidia.

thick. CONIDIOPHORES macronematous, mononematous, single or in groups,

erect, unbranched, straight or flexuous, cylindrical, brown to dark brown,

smooth, septate, 17-70 x 6-7.5 um. CONIDIOGENOUS CELLS monoblastic,

integrated, terminal, determinate, cylindrical, brown, smooth, 4.5-7.5 x 5-6

um. Conidial secession schizolytic. Conrp1A holoblastic, acrogenous, solitary,

Sporidesmium spp. nov. (China) ... 21

straight or curved, obclavate, smooth-walled, brown, 8-11-euseptate, 108-150

um long, 9-11 um thick in the broadest part, tapering gradually towards the

apex, base truncate, apex rounded, 3-4 um wide, and invested in a subglobose

drop of mucilage ca 12-24 um diam, base 4.5-6 um wide.

COMMENTS - Sporidesmium antidesmatis superficially resembles S. fragilissimum

(Berk. & M.A. Curtis) M.B. Ellis (Ellis 1958) and S. eupatoriicola M.B. Ellis

(Ellis 1958). However, S. antidesmatis differs from S. fragilissimum (conidia

32-92 x 8-9 um, verruculose) by its larger, smooth-walled conidia, and from

S. eupatoriicola (conidia 60-195 um long, apex 4-6 um wide, 14-31-euseptate)

by its slightly shorter conidia with narrower apices and fewer septa. In addition,

the conidia of S. antidesmatis have an apical mucilaginous appendage while

those of S. fragilissimum and S. eupatoriicola do not.

Sporidesmium machili Jian Ma & X.G. Zhang, sp. nov. FIG. 3

MycoBank MB 563264

Conidiophora macronemata, nonramosa, 15-60 x 3-5.5 um. Cellulae conidiogenae

monoblasticae, integratae, terminales, determinatae. Conidia solitaria, obclavata, ad

longa rostrata, laevia, 5-7-euseptata, 70-160 x 5-7.5 ym, basi truncate, ad 1-1.5 ym lata,

cellula apicali versus attenuate, pallide brunnea vel subhyalina, aseptata, rostro, ad usque

125 Xx 0.5-1.5 um.

Type: China, Guangdong Province: Chebaling National Nature Reserve, on decaying

twigs of Machilus chinensis (Benth.) Hemsl. (Lauraceae), 19 Oct 2010, J. Ma, (Holotype

HSAUP H5407; isotype HMAS 146158).

EryMo_oey: in reference to the host genus, Machilus.

Anamorphic fungi. COLONIES on natural substrate effuse, brown, hairy.

Mycelium partly superficial, but mostly immersed in the substratum,

composed of branched, septate, pale brown, smooth-walled hyphae, 1.5-3 um

thick. CONIDIOPHORES macronematous, mononematous, single or in groups,

erect, unbranched, straight or flexuous, cylindrical, brown, smooth, septate,

15-60 x 3-5.5 um. CONIDIOGENOUS CELLS monoblastic, integrated, terminal,

determinate, lageniform, brown, smooth, 13-23 x 3-4.5 um. Conidial secession

schizolytic. Conrp14_ holoblastic, solitary, acrogenous, straight or curved,

obclavate to long-rostrate, smooth-walled, brown to pale brown, 5-7-euseptate,

70-160 um long (rostrum included), 5—7.5 um thick in the broadest part, 1-1.5

um wide at the truncate base, apex extended into a pale brown to subhyaline,

aseptate, smooth rostrum, up to 125 um long, 0.5-1.5 um wide.

Comments - In terms of conidial morphology, Sporidesmium machili is similar

to S. circinophorum Matsush. (Matsushima 1975), S. takashii (Subramanian

1992), S. longirostratum M.B. Ellis, and S. tropicale M.B. Ellis (Ellis 1958).

However, S. machili differs from S. circinophorum (conidia 120-220 x

10-12 um, 8-14-euseptate) and S. tropicale (conidia 80-250 x 12-15 um,

22 ... Ma & al.

i (

20pm

20m

| \ D H

| | 20um

Fic. 3. Sporidesmium machili. A. Conidiophores with developing conidia.

B. Conidiophores with mature conidia. C-D. Conidiophores. E-H. Conidia.

wigz

wilgz

wilQZ

7-19-euseptate) by its smaller conidia with fewer septa, and from S. longi-

rostratum (conidia 42-72 x 6-9 um, 3-5-euseptate) by its longer and slightly

narrower conidia with more septa. The conidia of S. machili are smooth-

walled and are 5-7-eusepta while those of S. takashii are verruculose and

5-10-eusepta. In addition, the conidial rostrum in S. machili is filiform, pale

brown to subhyaline and aseptate, which is obviously different from those of

S. circinophorum, S. takashii, and S. tropicale.

Sporidesmium takashii Subram., Proc. Indian natn. Sci. Acad. B 58: 183, 1992. Fic. 4

Anamorphic fungus. COLONIES on natural substrate effuse, brown, hairy.

Mycelium partly superficial, partly immersed in the substratum, composed

of branched, septate, pale brown, smooth-walled hyphae, 1-3 um thick.

CONIDIOPHORES macronematous, mononematous, single or in groups,

erect, unbranched, straight or flexuous, cylindrical, brown, smooth, septate,

Sporidesmium spp. nov. (China) ... 23

—_20nm 20um

Fic. 4. Sporidesmium takashii. A. Conidiophores with conidia. B—C. Conidia.

13-60 x 3-5.5 um. CONIDIOGENOUS CELLS monoblastic, integrated, terminal,

determinate, cylindrical, brown, smooth, 10-16 x 3-4 um. Conidial secession

schizolytic. Conrp14_ holoblastic, solitary, acrogenous, straight or curved,

obclavate, rostrate, smooth-walled, brown to pale brown, 8-13(-16)-euseptate,

24 ... Ma & al.

68-150 um long, 5.5-7.5 um thick in the broadest part, tapering to 1.5-2.2 um

near the apex, 2.5-4 um wide at the truncate base.

SPECIMEN EXAMINED: CHINA, HAINAN PROVINCE: Xinglong Tropical Botanical

Garden, on decaying twigs of unidentified broad-leaved tree, 6 Dec 2009, J. Ma, HSAUP

H5212, HMAS 146159.

ComMENTS - Matsushima (1975) described this fungus as Sporidesmium sp.

Subsequently, Subramanian (1992) assigned it as a new species, S. takashii,

based on the earlier description. Sporidesmium takashii is morphologically

most similar to S. circinophorum and S. tropicale. However, S. takashii conidia

are distinctly smaller than those of S. circinophorum (conidia 120-220 x 10-

12 um) and S. tropicale (conidia 80-250 x 12-15 um). In addition, S. takashii

differs from S. circinophorum (conidia versicolored, apex 2.5-3.5 um wide)

and S. tropicale (conidia verruculose, apex 2-4 um wide) in its concolorous,

smooth-walled conidia with a narrower apex. Compared with the morphology

of S. takashii described by Matsushima (1975) and Subramanian (1992), our

collection fits well with the original description except for having more septa

(8-16 vs 5-10) and smooth-walled conidia. This is the first report of this species

in China.

Acknowledgments

The authors express gratitude to Dr Bryce Kendrick and Dr Eric H.C. McKenzie for

serving as pre-submission reviewers and for their valuable comments and suggestions.

This project was supported by the National Natural Science Foundation of China (Nos.

31093440, 30499340, 30770015) and the Ministry of Science and Technology of the

People’s Republic of China (Nos. 2006FY120100, 2006FY110500-5).

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Sporidesmium spp. nov. (China) ... 25

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Sutton BC, Hodges Jr CS. 1979. Eucalyptus microfungi. Chaetophragmiopsis gen. nov. and other

hyphomycetes. Nova Hedwigia 29: 593-607.

Wu WP, Zhuang WY. 2005. Sporidesmium, Endophragmiella and related genera from China. Fungal

Divers. Res. Ser. 15: 1-351.

MY COTAXON

http://dx.doi.org/10.5248/119.27

Volume 119, pp. 27-34 January-March 2012

Phlebopus spongiosus sp. nov. (Boletales, Boletinellaceae)

with a sponge-like tissue

NGUYEN Duc HOANG PHAM?*?’, HARUKI TAKAHASHI,

TOSHIMITSU FUKIHARU4, KIMINORI SHIMIZU,

BA DuNG LE° & AKIRA SUZUKI”7

' Graduate School of Horticulture, Chiba University, 648 Matsudo, Matsudo 271-8510, Japan

? Division of Microbiology, Biotechnology Center of Hochiminh City,

km 1900, Highway 1A, Dist. 12, Hochiminh City, Vietnam

> 284-1, Ouhama, Ishigaki, Okinawa 907-0001, Japan

* Natural History Museum and Institute, Chiba,

955-2 Aoba-cho, Chuo-ku, Chiba 260-8682, Japan

° Medical Mycology Research Center, Chiba University,

1-8-1 Inohana, Chuo-ku, Chiba 260-8673, Japan

° Dalat University, 01 PhuDongThien Vuong St., Dalat City, LamDong Province, Vietnam

’ Faculty of Education, Chiba University, 1-33 Yayoi-cho, Inage-ku, Chiba 263-8522, Japan

* CORRESPONDENCE TO: hoangphamedu@yahoo.com

ABSTRACT - A new species, Phlebopus spongiosus, is described with a peculiar sponge-like

tissue in which the hollow spaces in the tubes are filled in by pleurocystidia-like elements, and

a thin membrane of dissepiments encloses the immature pores. It occurs in citrus orchards

(Citrus maxima) in southern Vietnam.

Key worps - tropics, black bolete, edible mushroom, crypta

Introduction

Heim (1936) proposed Phlebopus as a subgenus of Boletus to accommodate

Boletus colossus R. Heim. Singer (1936) elevated Phlebopus to genus with

P. colossus (R. Heim) Singer as type species. Subsequently, Heinemann &

Rammeloo (1982), Singer et al. (1983), and Singer (1986) emended the generic

limits to encompass all taxa of Phaeogyroporus (Singer 1944). With 12 species

(Kirk et al. 2008: 522), Phlebopus was classified in the Boletinellaceae (Boletales)

by Binder & Bresinsky (2002).

Phlebopus is a tropical or subtropical genus with boletoid basidiomata

producing an olivaceous brown spore print, ellipsoidal smooth basidiospores

28 ... Pham & al.

(light microscope), and hyphae with clamp connections. In Southeast Asia, the

two dominant edible taxa — P portentosus (Berk. & Broome) Boedijn (Saccardo

1888, Boedijn 1951) and P marginatus Watling & N.M. Greg. (Saccardo 1888,

Watling & Gregory 1988) — are regarded as conspecific by Watling & Gregory

(1988) and Watling & Li (1999).

During surveys of mycobiota in southern Vietnam, we collected in several

orchards a taxon with a habit reminiscent of P portentosus. In the present paper,

this taxon is described and illustrated as new to science.

Materials & methods

Geographical coordinates of localities were determined by GPS (Garmin - eTrex

Legend”) and Google Earth 5.0.11733.9347 software. Color notation in parentheses was

determined using the Methuen Handbook of Colour (Kornerup & Wanscher 1981).

The microscopic characteristics were observed using a Nikon Labophot-2 equipped

with DIC (Differential Interference Contrast) optics after hand-sectioning and

mounting with 10% ammonium hydroxide aqueous solution. Abbreviations used:

ccd

Q = mean length/width ratio measured from “n” number of spores; m = mean spore

length and width; P. = Phlebopus.

A strain isolated from basidioma hyphae is maintained on modified Melin and

Norkrans medium (Brundrett et al. 1996: 224), 20°C, in darkness.

All specimens are deposited in the herbarium of the Natural History Museum and

Institute, Chiba, Japan (CBM).

Taxonomy

Phlebopus spongiosus Pham & Har. Takah., sp. nov. PL. 1-6

MycosBank MB 518549

Differs from other Phlebopus species by sponge-like tissues with cystidioid elements

filling hollow spaces in immature tubes, a thin membrane of subcylindrical dissepiments

enclosing the immature pores, and a dark brown spore print.

TYPE: Vietnam. TienGiang: MyTho City, Highway 1A (UTM, 0646165, 1151001), 20 Jan

2006, N.D.H Pham (Holotype, CBM FB-38014)

ErymMo_oey: From the Latin, referring to the sponge-like tissue

MACROCHARACTERS ~ Pileus 50-120(-500) mm broad, at first hemispherical

with incurved margin, then becoming plano-convex to convex with decurved

margin; surface dry, at first subtomentose, then glabrescent; brown to yellowish

brown (5D7-8 to 5E7-8) overall, sometimes tinged with olive (3F4-5) on old

specimens. Flesh 10-20 mm thick at the center of pileus, soft, yellow (3A7

to 3B7-8) to vivid yellow (3A8), in places gradually bluing when exposed;

odor indistinct when fresh; taste after cooking slightly like persimmon rind.

Hymenophore decurrent to subdecurrent at first, then more or less depressed

around the stipe; tubes < 3 mm long in dried material, vertically arranged;

hymenium usually with two thirds of the tubes filled in with yellowish orange

Phlebopus spongiosus sp. nov. (Viet Nam) ... 29

Figures 1-3. Phlebopus spongiosus. 1: Basidiomata in its habitat. 2: Vertical section of the immature

basidioma; bar = 2 cm. 3: Vertical section of the hymenophore in dried specimens showing the

sponge-like tissue arrow); bar = 1 mm.

(5A7-8 to 5B7-8) to yellowish brown (5E5-6 to 5F5-6), sponge-like tissue

projecting from the sides of the tube walls, but this occluding tissue disappearing

in old specimens; pores small (1-3 per mm), angular to subcircular, more

elongated around the stipe, covered by a thin membrane in young specimens,

brown (7D8-7E8) when young, then brown (7E5-6) to dark brown (7F5-6).

Stipe 80-150 x 15-50 mm, central, subequal or somewhat enlarged at the base,

sometimes tapering downward, solid; surface dry, subtomentose to tomentose,

longitudinally striate toward the base; entirely yellowish brown (5E4-8 to 5F4-

8), becoming darker where handled; basal mycelium brownish yellow. Spore

print dark brown to reddish brown (8F5-6).

MICROCHARACTERS - Basidiospores m = 8.3 x 6.4 um [7.9-8.8(-9.1) x

5.9-7.1(-7.4) um, n = 47, Q = 1.30], ovoid to shortly ellipsoid, smooth under

the light microscope, brown (5D6-7), inamyloid, thick-walled. Basidia 21-27

x 8-10 um (without sterigmata), clavate, 4-spored; sterigmata 3-4 um long.

Hymenophoral trama divergent-bilateral of the Boletus subtype, hyaline,

with gelatinized hyphae 4.5-10 um diam.; mediostratum 80-110 um diam.

when young, 40-60 um diam. at maturity. Sponge-like tissue composed of

heteromorphous terminal cells occasionally protruding into the tube lacunae

from the sides of the tube walls; constituent elements numerous, (29-)45-53 x

4.5-7.4 um, cystidioid but not truly pleurocystidia, broadly clavate or fusoid-

ventricose to ventricose-rostrate, thin-walled, with yellow (4B-5 to 4A-6)

30 ... Pham & al.

Figures 4-5. Phlebopus spongiosus. 4: Vertical section through part of the hymenophore showing

the sponge-like tissue (arrow) made up of heteromorphous terminal cells (*) unusually protruding

into the lacunae of the tubes from the sides of the tube walls; bar = 50 um ina, = 30 um in b. 5: SEM

figures of basidiospores; bar = 2.5 um.

intracellular pigment, converging toward the center of the tube chamber at

least in young specimens but gradually disrupted. Dissepiments resembling

Phlebopus spongiosus sp. nov. (Viet Nam) ... 31

gp Gy

2 b c d

FiGurE 6. Phlebopus spongiosus. A: Basidiospores and basidia. B: Heteromorphous terminal

cells protruding into the lacunae of the tubes from the sides of the tube walls. C: Dissepiments.

D: Terminal cells of the pileipellis. Bar = 10 um.

cheilocystidia, remarkably projecting beyond the hymenium, 25-33(-42) x

4-5(-6.5) um, subcylindrical to cylindrical-clavate, thin-walled, distinctly

clamped at base, with yellow (4B-5 to 4A-6) intracellular pigment, forming the

membranous thin layer on the immature pore surface but gradually collapsing.

Pileipellis consisting of repent, appressed, interwoven hyphal elements;

terminal cells 22-29 x 3.3-4.7 um, cylindrical, thin-walled, with yellow (4B7-

8) intracellular pigment, golden yellow to brownish (5B7 or 5C7) in the wall

mass. Pileitrama composed of cylindrical, loosely interwoven hyphae 4-6 um

diam., colorless or greyish yellow, smooth, inamyloid, thin-walled. Stipitipellis

densely constructed, brown (6D8 or 6E7); terminal cells similar to those of the

pileipellis. Stipe trama composed of longitudinally arranged, cylindrical hyphae

2-4.5 um diam., thin-walled, colorless or brownish (6C4-6C5), inamyloid,

rather compactly arranged compared to the pileitrama. Clamp connections

constantly present in all tissues and in vitro.

Hasitat - Almost year-round, solitary to scattered in citrus farms,

(dominated by Citrus maxima (Burm.) Merr).

ADDITIONAL SPECIMENS EXAMINED: VIETNAM. TiENGIANG: MyTuo Ciry, Highway

1A (UTM, 0646165, 1151001), 25 Jun 2007, N.D.H Pham (CBM FB-38670; culture

Phle01, Laboratory of Microbiology, Biotechnology Center of HoChiMinh City).

HoCuiMinu City: TanPuu Dist., Cautre Company, 16 May 2007, N.D.H Pham

(CBM FB-38017); Dist. 12, Biotechnology Center of HoChiMinh City (10°50'58.95"N,

106°37'03.89"E), 30 Mar 2008, N.D.H Pham (CBM FB-38671).

Discussion

The most distinctive features of Phlebopus spongiosus are the sponge-

like tissue composed of numerous fusoid-ventricose to ventricose-rostrate

pleurocystidioid elements, the subcylindrical to subclavate dissepiments

32 ... Pham & al.

forming a thin membrane that covers the immature pores, and the dark-brown

spore print not found in any other known Phlebopus species.

Although its dark brown spore print appears foreign to Phlebopus sensu

Singer (1986), the shortly ellipsoid smooth basidiospores and clamp connections

merit placing P spongiosus within Phlebopus (Singer 1986).

Its medium to large dark yellowish-brown boletoid basidioma bears a

superficial resemblance to the Southeast Asian P. portentosus (Boedijn 1951,

Heinemann & Rammeloo 1982, Singer et al. 1983) and P. marginatus originally

described from Australia (Watling & Gregory 1988, Grgurinovic 1997). Both

taxa differ from P. spongiosus in their olivaceous brown spore print and lack of

distinct hymenial cystidia (Boedijn 1951; Pegler 1986; Watling & Li 1999).

It should be noted that P spongiosus basidiomata are sometimes accompanied

by a crust-like mycelial structure around nearby citrus tree roots reminiscent

of “crypta” as defined by Singer (“sleeve-like formations around tree roots in

tropical and subtropical conditions;” 1986: 10). Further critical investigations,

including cultural and molecular studies, are needed to confirm its relationship

with P. spongiosus basidiomata.

The crypta-like mycelium of P spongiosus is comparable to that of three

neotropical taxa: P. tropicus (Rick) Heinem. & Rammeloo (Singer 1944, Singer

& Digilio 1957, Heinemann & Rammeloo 1982, Singer et al. 1983), P. beniensis

(Singer & Digilio) Heinem. & Rammeloo (Heinemann & Rammeloo 1982,

Singer et al. 1983), and P brasiliensis Singer (Singer et al. 1983). Those taxa

differ from P. spongiosus mainly by lacking distinct hymenial cystidia and having

olivaceous-brown spore print. Moreover, according to the Phlebopus key by

Singer et al. (1983), the wide pores (21 mm diam.) or boletinoid/subgyrose

hymenophore separate P. tropicus from P. spongiosus. Miller et al. (2000) noted

that Puerto Rican P. beniensis specimens had well-developed cheilocystidia but

provided no information regarding the presence or absence of pleurocystidia.

Phlebopus spongiosus also shares well developed fusoid-ventricose to

ventricose-rostrate pleurocystidia-like elements with P cystidiosus Heinem &

Rammeloo from Ethiopia (Heinemann & Rammeloo 1982). However, the lack

of clamp connections causes P. cystidiosus to seem out of place in Phlebopus.

Its glabrous non-reticulate stipe, ellipsoid basidiospores, and absence of clamp

connections make P cystidiosus seem more closely related to Rubinoboletus

Pilat & Dermek (Heinemann & Rammeloo 1983, Watling & Gregory 1988, Li

& Watling 1999, Watling & Li 1999).

The thin membrane enclosing the immature pores might also suggest a

relationship between P. spongiosus and the secotioid boletes (Thiers 1984).

However, the secotioid boletes are generally associated with an often misshapen

pileus, irregularly oriented tubes, and a typically less-developed stipe, characters

not seen in P. spongiosus.

Phlebopus spongiosus sp. nov. (Viet Nam) ... 33

Acknowledgments

We are grateful to Ms. Dao Kieu Dung, the owner of citrus farms in MyTho City,

TienGiang Province and ChoLach District, BenTre Province, Vietnam who supported

us for studying in field trip. We are grateful to Dr. Roy Watling (Caledonian Mycological

Enterprises, UK) and Dr. Timothy J Baroni (State University of New York at Cortland,

USA) for reviewing of the manuscript. We are grateful to Dr. Roy E Halling (New York

Botanical Garden, USA) for suggestions on an early draft of the manuscript. We also

thank Mr. Le Duy Thang (University of Science in Hochiminh City, Vietnam) and Dr.

Bryn TM Dentinger (Royal Botanic Gardens, Kew, UK) for help in our research.

Literature cited

Binder M, Bresinsky A. 2002. Derivation of polymorphic lineage of gasteromycetes from boletoid

ancestors. Mycologia 94: 85-98. http://dx.doi.org/10.2307/3761848

Boedijn KB. 1951. Some mycological notes. Sydowia 5: 211-229.

Brundrett M, Bougher N, Dell B, Grove T, Malajczuk N. 1996. Working with mycorrhizas in

forestry and agriculture. Australian Centre for International Agricultural Research (ACIAR),

Canberra, Australia.

Grgurinovic CA. 1997. Larger fungi of South Australia. The Botanic Garden of Adelaide & State

Herbarium and The Flora & Fauna of South Australia Handbooks Committee, Adelaide,

Australia.

Heim R. 1936. Observations sur la flore mycologique malgache, HI. Trois bolets gigantesques

d'Afrique et de Madagascar. Revue Mycol 1: 3-18.

Heinemann P, Rammeloo J. 1982. Observations sur le genre Phlebopus (Boletineae). Mycotaxon

15: 384-404.

Heinemann P, Rammeloo J. 1983. Gyrodontaceae p.p. Boletineae. Flore illustrée des champignons

d'Afrique centrale 10: 173-198.

Kirk PM, Cannon PE, Minter DW, Stalpers JA. 2008. Dictionary of the fungi, 10™ edition. CABI

Europe, Wallingford, UK.

Kornerup A, Wanscher JH. 1981. Methuen handbook of colour, 3" edition. Eyre Methuen, London,

UK.

Li T-H, Watling R. 1999. New taxa and combinations of Australian boletes. Edinb Jour Bot 56:

143-148. http://dx.doi.org/10.1017/S0960428600002419

Miller OK, Lodge DJ, Baroni TJ. 2000. New and interesting ectomycorrhizal fungi from Puerto

Rico, Mona, and Guana Islands. Mycologia 92: 558-570. http://dx.doi.org/10.2307/3761516

Pegler DN. 1986. Agaric flora of Sri Lanka. Kew Bulletin Additional Series 12. Royal Botanic

Garden Kew, UK.

Saccardo PA. 1888. Sylloge fungorum VI. Edwards Bother Inc., Ann Arbor, Michigan, USA reprint

in 1944.

Singer R. 1936. Das system der Agaricales. Annls Mycol 34: 286-378.

Singer R. 1944. New genera of fungi. Mycologia 36: 358-368.

Singer R. 1986. The Agaricales in modern taxonomy. Koeltz Scientific Books, Koenigstein,

Germany.

Singer R, Digilio APL. 1957. Las boletaceas austrosudamericanas. Lilloa 28: 247-268.

Singer R, Araujo I, Ivory MH. 1983. The ectotrophically mycorrhizal fungi of the Neotropical

lowlands; especially central Amazonia. Beihefte zur Nova Hedwigia, Hefl 77. J Cramer, Vaduz,

Liechtenstein.

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Thiers HD. 1984. The secotioid syndrome. Mycologia 76: 1-8. http://dx.doi.org/10.2307/3792830

Watling R, Gregory NM. 1988. Observations of the Boletales of the Cooloola Sandmass, Queensland

and notes on their distribution in Australia. Proceedings of the Royal Society of Queensland

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UK.

MY COTAXON

http://dx.doi.org/10.5248/119.35

Volume 119, pp. 35-63 January-March 2012

Type studies in Helvella (Pezizales) 1

FIDEL LANDEROS*?3*, TERESA ITURRIAGA” & LAURA GUZMAN-DAVALOS?

‘Facultad de Ciencias Naturales, Universidad Aut6noma de Querétaro,

Avenida de las Ciencias s/n, Santiago de Querétaro, Qro., 76270, Mexico

*Departamento Biologia de Organismos, Universidad Simon Bolivar, Caracas, Venezuela

’Departamento de Botanica y Zoologia, Universidad de Guadalajara,

Apartado postal 1-139, Zapopan, Jal., 45101, Mexico

CORRESPONDENCE TO *: *landeros@uaq.mx, *titurri@usb.ve, *lguzman@cucba.udg.mx

ABSTRACT — Sixteen type and some additional specimens representing fifteen species

assigned to Helvella were studied with the aim of providing new data for their improved

delimitation and recognition. Specimens were received on loan from DAOM, FH, H, K, MA,

MICH, S and UPS. A synoptic key to the studied species is provided to contribute to a better

understanding of the genus and the taxonomy of the morphospecies. Original descriptions

are emended to include data on the anatomy of the apothecial excipulum and stipe layers,

apothecium and stipe reactivity to Melzer’s reagent, and ascospore ornamentation. Reasons

are given for maintaining H. subglabra, H. atra, H. griseoalba, and H. costifera as independent

taxa and for considering Helvella hyperborea a synonym of H. costifera. Previously imprecise

descriptive terms are clarified.

Key worps — Ascomycota, holotype, isotype, lectotype, revision

Introduction

The genus Helvella L. includes approximately 52 species (Kirk et al. 2008).

Its distribution is basically north temperate, with only one tropical species

recognized (Dissing 1979). Traditional taxonomy of the genus has been

based mainly on ascoma shape and color and the presence or absence of

projecting hyphae on the sterile surface (ectal excipulum) of the apothecium

(Dissing 1966b, Weber 1972, Abbott & Currah 1997). Characters such as

ascus development (aporhynchous or pleurorhynchous), ascospore shape and

size, paraphysis color, and the presence of pigment in the apothecium ectal

excipulum cells have proven to be of taxonomic value in some species (Weber

1972; Harmaja 1977b, 1979; Abbott & Currah 1997).

Although Nannfeldt (1937) proposed using the term “apothecium”

specifically for Helvella to describe the entire ascoma, Weber (1972, 1975) and

36 ... Landeros, Iturriaga & Guzman-Davalos

Abbott & Currah (1988, 1997) used “apothecium” exclusively for the upper

ascoma — i.e., the hymenium and its immediate supporting tissues — without

the stipe. Eckblad (1968) commented that some authors carefully avoid the

term apothecium for pileate forms like Helvella, whereas Dissing (1966b) and

Calonge & Arroyo (1990) employed this term loosely, either for the complete

ascoma or just for the upper part. Here, we follow Weber's (1972) concept,

applying the term apothecium only for the upper ascoma, except that where she

differentiated stipe tissues with “ectal excipulum” and “medullary excipulum,”

we use “outer layer” and “inner layer,’ because in discomycetes “excipulum”

currently is interpreted as the “tissue or tissues containing the hymenium in an

apothecium” (Kirk et al. 2008).

In all Helvella species the medullary excipulum and stipe inner layer are

composed of “textura intricata” (interwoven hyphae) while the ectal excipulum

and stipe outer layer are a “textura angularis” (almost isodiametric cells without

intracellular spaces) (Korf 1952, Dissing 1966, Eckblad 1968, Weber 1972).

These cells are arranged in rows perpendicular to the surface. In glabrous

ascomata, the ectal excipulum external layer is a palisade of clavate hyphal

end-cells, ending more or less at the same level (Fic. 1) (Weber 1972). Hairy

ascomata have an ectal excipulum outer layer that Eckblad (1968) described as

“consisting of more or less swollen chains of cells often converging into distinct

clusters.’ These clusters are called “hyphal fascicles” (Weber 1972) or “chains of

cells and/or fascicled hyphal chains” (Abbott & Currah 1997). Macroscopically,

they give the appearance of pubescence or hairs of different lengths (Frcs 2-3)

and are important in the taxonomy of the genus (Dissing 1966). Descriptive

terms used in the literature — “finely pubescent,’ “furfuraceous,” “granulose,”

“hairy, “pruinose,’ “pubescent, “roughened,” “velvety,” “villose” (Dissing 1966;

Weber 1972, 1975; Dennis 1981; Abbott & Currah 1997) — have generated

confusion through their different meanings, incorrect application, and/or the

lack of consistent usage. Kaushal (1991) alone consistently uses only “pubescent”

to describe the hairs; furthermore he included the hyphal fascicle length for

each species.

Only three papers have described the stipe tissue: GOmez & Herrera (1965)

and Weber (1972, 1975). Gomez & Herrera concluded that because stipe tissue

is continuous with apothecium tissue, it is not a useful character. Although

Weber (1972) described both tissues as similar in color and structure, she did

indicate color differences between tissues in some species descriptions.

Pigment distribution, mainly in ectal excipulum cells (see Figs 19-21) and

paraphyses — ie., intracellular (cytoplasmic), in the cell wall, or encrusting

(deposited pigment forming an irregular crust on the wall) — is controversial.

Eckblad (1968), Weber (1972), and Harmaja (1977b, 1979) considered

pigmentation to be diagnostic. Eckblad (1968) wrote it is a characteristic mostly

Helvella type studies — 1... 37

Hh \ EH ON

teen aie

may

WOOOAMdDDO

GHSOOSSO

Fics 1-9. 1: Ectal excipulum of a glabrous apothecium (Helvella leucopus var. populina). 2: Hyphal

fascicle of the outer layer of a subpubescent stipe (H. griseoalba). 3: Hyphal fascicle of the ectal

excipulum of a pubescent apothecium (H. costifera)—a: short, b: large. 4: Pleurorhynchous asci

(H. pocillum). 5: Aporhynchous asci (H. crassitunicata); 6: Common paraphyses (H. subglabra).

7: Paraphyses with a thick-walled brown cap or collar at the apex (H. paraphysitorquata). 8: Thick-

walled paraphyses (H. crassitunicata). 9: Ascospores (H. solitaria) —a: ornamented, b: smooth.

Scale bars: 1-8 = 20 um, 9 = 8 um.

used on species level in Pezizales, but “[u]nfortunately too many descriptions

simply state whether the paraphyses are coloured or not, not where the pigments

are located” With respect to Helvella, Eckblad (1968) noted, “[c]hemically

and genetically it is probably a very short step from pigmented paraphyses to

38 ... Landeros, Iturriaga & Guzman-Davalos

hyaline ones, e.g. from the greyish-black paraphyses of Helvella lacunosa to

the colourless paraphyses of H. crispa.’” Weber (1972) recognized two patterns

of pigment distribution: 1) intracellular pigments, occurring in species with

buff, tan or brown ascomata, and 2) pigments associated with hyphal walls,

characteristic of gray to black species. On the other hand, Dissing (1966b)

and Abbott & Currah (1997) considered pigmentation as highly variable and

of limited use in circumscribing Helvella species. Dissing (1966b) indicated it

is difficult to assess, especially in species where hymenium color variation is

very large (as in H. lacunosa Afzel.) and hymenium color ranges from pale

gray to gray brown to black due to pigments in the paraphyses. Only Dissing

(1964, 1966b) went beyond just describing the colors of the structures; he also

evaluated the color before and after treatment with cotton blue. Unfortunately,

no further works analyze this issue.

Chemical reactions to Melzer’s reagent have not been considered as diagnostic

characters for structures other than ascospores or asci. Although excipulum

cells in other ascomycete genera (Perrotia, Hymenoscyphus, Epibryon) show a

hemiamyloid reaction, its taxonomic value is uncertain (Baral 1987).

Helvella ascospores have limited taxonomic value. In fact, their size is a

key feature only for H. pocillum and H. crassitunicata (Weber 1975; Harmaja

1976), while the fusoid to subfusoid shape (differing from the typically ellipsoid

ascospores found elsewhere) is found only in H. macropus (Pers.) P. Karst.

(Abbott & Currah 1997) and H. terrestris (Velen.) Landvik (Landvik et al.

1999). Ascospore ornamentation is highly complex and a subject of controversy.

Dissing (1964) and Dissing & Nannfeldt (1966) regarded ornamented ascospores

in Helvella as immature, while Weber felt (1972) that they corresponded

to the mature condition. Eckblad (1968) described ascospores with “false

ornamentation, and Schumacher (pers. com.) mentioned that during the final

stage of ascospore development, some secondary wall remnants might adhere

to the ascospore surface, producing a highly inconsistent pattern; typically,

a number of ascospores do not get such adherences at all and consequently

remain smooth. On the other hand, Abbott & Currah (1997) gave to this

character a high value, because they considered verruculose ascospores unique

to the subgenus Macropodes (Dissing) S.P. Abbott.

Species concepts are similarly varied. Some authors with a narrow species

concept, such as Harmaja (1976, 1977a,b, 1979), segregated species based

on subtle differences. Others with a wider species concept (e.g., Abbott &

Currah 1997) use fewer diagnostic characters to separate Helvella species. The

purpose of our study was to review selected Helvella type specimens to delimit

morphospecies and to determine which characters could be used in taxonomic

and phylogenetic studies of the genus. We present five sets of types belonging

to species with problematic delimitations: 1) H. costifera vs. H. hyperborea

Helvella type studies — 1... 39

and H. griseoalba; 2) H. crassitunicata vs. H. pocillum; 3) H. subglabra vs.

H. atra; 4) H. solitaria vs. H. queletii and H. ulvinenii; and 5) H. verruculosa

vs. H. dryadophila. We also add to the earlier descriptions of H. leucopus var.

populina, H. maculata, H. paraphysitorquata, and H. robusta. Finally, we suggest

a set of standard terms to clarify descriptions of Helvella species.

Materials & methods

Sixteen type as well as some additional specimens from eight herbaria (DAOM, FH,

H, MA, MICH, O, S, UPS) were studied. Herbarium abbreviations follow Holmgren

et al. (1990). Free-hand sections from apothecium and stipe were made with a razor

blade directly from dry specimens. Sections and mycelium were placed first in 70% ethyl

alcohol and subsequently in water to rehydrate the tissues. Specimens were examined

under a Zeiss Axioskop 2 plus microscope with a drawing tube. Microphotographs were

taken under a Zeiss Axioskop 40 microscope using Axio Vision 4 software. Ascospore

measurements were made on mature free spores outside the asci, or on those attached

to the hymenial surface, stipe, or basal mycelia. Mounts from apothecium, stipe, and

mycelium were also made in Melzer’s reagent (without pre-treatment in KOH) and

cotton blue. We follow in part the terminology used by Baral (1987) to describe the

hemiamyloid reaction in Melzer’s reagent (‘Melzer’s’): “rr+” for hemiamyloid (“solely

red”) reaction and “rr—” for negative reaction. All measurements and features recorded

are from type specimens. Where additional non-type specimens were studied for some

species, observations that differ from those of the type are indicated under “Remarks”

but not included in the technical description.

Terms used to refer to the surface of the apothecial ectal excipulum and stipe outer

layer are: GLABROUS (FIG. 1), SUB-PUBESCENT (for fascicled hyphal chains < 50 um long;

Fic. 2), and PUBESCENT (for fascicled hyphal chains > 50 um long; Fic. 3). As the hairs’

length can vary in the last case, their measurements are given in the descriptions. See

also “Results & discussion’.

Results & discussion

We studied sixteen type specimens and present only new information or

data that differs from the protologue or previous publications. In addition

to the characters established by previous authors (Dissing 1966b; Calonge &

Arroyo 1990; Abbott & Currah 1997), we found that pigmentation of apothecial

cells, stipe tissue anatomy, and the reddish (hemiamyloid) reaction in Melzer’s

reagent (Fics 10-11) are important characters that contribute to species

recognition in Helvella. Because we found no variation in the basal mycelium

of the stipe, which is universally composed of smooth-walled, hyaline, rr-,

interwoven hyphae, we do not include it in the species descriptions. Nor do

we make further mention of the fact that ascospores of all species have both

cyanophilic cytoplasm and ornamentation (when present).

Regarding the value of pigmentation in the structures, we think it is important

to study it from two perspectives: a) location and b) intensity. Concerning

40 ... Landeros, Iturriaga & Guzman-Davalos

location, pigment may occur in the cytoplasm, in the wall, or encrusted on

the wall (Fics 19-21). We observed that while the first two are constant within

species, encrusted pigments are variable. For example, in H. costifera there are

specimens with encrusted paraphyses and others without such incrustations.

On the matter of pigment intensity, we return to cotton blue used first by

Dissing (1964) to note that there are species where the cytoplasmic pigment

color is so intense that it is still visible even after adding cotton blue, while

others have lighter pigment no longer visible when adding the colorant (Fics

12-17). We realized that this characteristic is constant among specimens of the

same species. This also happens with the pigment in the cell wall.

As indicated in the introduction, only Weber (1972, 1975) described the

stipe tissue. We consider this character taxonomically valuable for the following

reasons: 1) the hyphal fascicle lengths among specimens of the same species

are more constant than in the apothecial sterile surface (see H. hyperborea

remarks); 2) the pigments in the ectal excipulum and stipe outer layer can

differ in water and cotton blue (see H. ulvinenii remarks; Fic. 22); and 3) the

hemiamyloid reaction can occur in stipe tissue but not in the excipulum (e.g.

H. leucopus var. populina).

As Baral (1987) noted, we know little about hemiamyloid reaction, and there

indeed are few records of this reaction in tissues other than the hymenium. We

report this reaction in different apothecial and stipe tissues for H. leucopus var.

populina, H. robusta, and H. subglabra; we have also observed this reaction in

specimens of H. albella Quél., H. connivens Dissing & M. Lange (holotype),

H. elastica Bull., and H. stevensii Peck (unpublished data).

Concerning ascospore ornamentation, we make the following points. 1)

Ornamented ascospores are not restricted to H. subgen. Macropodes (here

represented only by H. macropus, Fic. 18a) as Abbott & Currah (1997)

proposed. We have also observed ornamented ascospores (generally only in

mature ascospores outside the asci) in the type specimens of H. leucopus var.

populina (Fic. 18b), H. paraphysitorquata (Fic. 18c), H. solitaria, H. subglabra,

and H. ulvinenii. 2) We agree with Weber (1972) that it is the mature Helvella

ascospores that are verrucose, contrary to the assumption of Dissing (1964)

and Dissing & Nannfeldt (1966) that ornamented ascospores are the immature

ones. 3) When citing ascospore sizes, it is important to indicate what kind of

spores (smooth vs. ornamented) were measured, because in some species they

have different size range (Fic. 9) (see remarks under H. queletii).

As noted previously, many different terms have been used to describe the

apothecial and stipe outer layer surfaces, giving the impression that there were

different types of hairs. However, in all cases these hairs are developmentally

the same: they consist of hyphal clusters that differ only in length. The tiny hairs

have been especially problematic; Weber (1972) and Abbott & Currah (1997)

Helvella type studies — 1... 41

Figs. 10-14. 10: Reaction of the apothecial excipulum to Melzer’s reagent—a: negative (Helvella

solitaria), b: hemiamyloid in medullary excipulum (H. robusta). 11: Reaction of stipe tissues to

Melzer’s reagent—a: negative (H. hyperborea), b: hemiamyloid in inner layer (H. subglabra).

12: Apothecium hyphal fascicles (H. costifera)—a: brown pigments visible in water, b) pigments

not visible in cotton blue. 13: Apothecium hyphal fascicles (H. griseoalba)—a: only terminal

cells are pigmented, b: pigments visible in cotton blue. 14: Apothecium hyphal fascicles

(H. dryadophila)—a: brown pigments visible in water, b: brown pigments visible in cotton blue.

Scale bars: 10a, 11a, 11b = 200 um, 10b = 100 um, 12-14 = 20 um.

42 ... Landeros, Iturriaga & Guzman-Davalos

Fics. 15-18. 15: Paraphyses of Helvella costifera—a: brown pigments visible in water, b: pigments

not visible under cotton blue. 16: Paraphyses of H. paraphysitorquata—a: brown pigments visible

only in the thickened apices in water, b: pigments visible in cotton blue. 17: Paraphyses of H.

dryadophila—a: brown pigments visible in water, b: pigments visible in cotton blue. 18: Ornamented

ascospores in cotton blue—a: H. macropus, b: H. leucopus var. populina, c: H. paraphysitorquata.

Scale bar: 15, 17 = 20 um, 16a = 100 um, 18 = 10 um.

Helvella type studies — 1... 43

Fics. 19-22. 19: Hyaline ectal excipulum cells in water (Helvella griseoalba). 20: Wall pigments

in ectal excipulum cells (H. hyperborea). 21: Wall and cytoplasmic pigments in ectal excipulum

cells (H. dryadophila). 22: H. ulvinenii—a: apothecium thin section in water, b: ectal excipulum

in cotton blue, notice the brown pigments, c: stipe layers in water, d: stipe outer layer under

cotton blue. Scale bar: 19-21 = 10 um, 22a, 22b = 100 um, 22c = 200 um, 22d = 20 um.

used indistinct terms such as “finely pubescent,’ “furfuraceous,’ “granulose,”

or “granulose-roughened” — all with very different meaning. Here we propose

to the term “subpubescent” to indicate that they have the same microscopic

structure (i.e. are hyphal fascicles) of the hairs found on a “pubescent” surface

but are simply shorter (Fic. 2). Likewise, longer hairs have been described

as “pubescent” or “villose,’ without considering that the terms may refer to

different types of hairs. We use “pubescent” (Fic. 3) to emphasize that Helvella

has only one hair type that varies only in length. This agrees with Kaushal’s

(1991) terminology.

44 ... Landeros, Iturriaga & Guzman-Davalos

Reasons to retain H. subglabra as independent of H. atra and H. griseoalba

separate from H. costifera are discussed below. We agree with Harmaja

(1977a) that H. queletii is a synonym of H. solitaria and with Abbott & Currah

(1997) that H. dryadophila is a synonym of H. verruculosa. We also propose

H. hyperborea as a synonym of H. costifera.

We present below a synoptic key of the 12 Helvella species considered in

this paper. Although not all Helvella species are included, we feel that this

key provides additional morphological insights into the treated species and

contributes to a better understanding of the genus.

Synoptic key to species of Helvella included in this work

1) HH. costifera (= H. hyperborea)

2) — H. crassitunicata

3) — H. griseoalba

4) —H. leucopus var. populina

5) H. maculata

6) — H. paraphysitorquata

7) H. pocillum

8) H. robusta

9) — H. solitaria (= H. queletii)

10) H. subglabra

11) H. ulvinenii

12) H. verruculosa (= H. dryadophila)

APOTHECIUM SHAPE (MATURE)

a. Cup-shaped 1, 2, 3, 7, 8, 9, 11, 12

b. Lobed 4, 5, 6, 8, 10

APOTHECIAL STERILE SURFACE

a. Glabrous 2, 4

b. Subpubescent (hyphal fascicles < 50 um long) 2, 7, 8, 9, 10, 11, 12

c. Pubescent, with hyphal fascicles 50-150 um long 1, 3, 5, 6, 7, 8, 9, 11, 12

d. Pubescent, with hyphal fascicles > 150 um long 1, 5, 6

RIBS (APOTHECIAL STERILE SURFACE)

a. Missing or reaching up to % of the surface 2, 4, 6, 7, 9, 10, 11, 12

b. Reaching from % to % of the surface 1, 3

c. Reaching from % to the edge of the apothecium 1, 5, 8

d. Simple 1,5

e. Bifurcated 1, 3, 5, 8

f. Anastomosed 1, 8

STIPE (SURFACE CONFIGURATION)

a. Even 4, 10

b. Only ribbed 1, 2, 3, 7, 9, 11, 12

c. Ribbed and lacunose 5, 8

d. Consisting of 2-3 strands free or apically partly fused 6

Helvella type studies — 1...

RIB (STIPE)

a. Sharp-edged 5, 8

b. Blunt-edged 1, 2, 3, 7, 9, 11, 12

STIPE (STERILE SURFACE)

a. Glabrous 2, 3, 4

b. Subpubescent (hyphal fascicles < 50 um long) 2, 3, 8, 9, 10, 12

c. Pubescent (hyphal fascicles 50-150 um long) 1, 5, 6, 7, 8, 9, 11, 12

STIPE COLOR

a. Light:tones-1; 2, 3;-5,.6, 7, 8,9, 10, 11, 12

b. Dark tones 4

STIPE CONTEXT

a. Solid 1, 2, 3, 6, 7, 9, 10, 11, 12

b. Hollow 4, 5, 8

ASCUS TYPE

a. Aporhynchous 2

b. Pleurorhynchous 1, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12

ASCOSPORE SIZE

a. Usually < 23 um long 1, 3, 4, 5, 6, 8, 9, 10, 11, 12

b. Usually > 23 um long 2, 7

THICK-WALLED PARAPHYSES

a. Absent 1, 3, 4, 5, 7, 8, 9, 10, 11, 12

b. Present 2

c. Thickness restricted to the apex, forming a collar or hood 6

MELZER’S REACTION (MEDULLARY EXCIPULUM)

a. Negative 1, 2, 3, 4,5, 6, 7, 9, 11, 12

b. Hemiamyloid (reddish) 8, 10

MELTZER’S REACTION (STIPE INNER LAYER)

a. Negative 1, 2, 3, 5, 6, 7,9, 11, 12

b. Hemiamyloid (reddish) 4, 8, 10

MELZER’S REACTION (STIPE OUTER LAYER)

a.Negative 1,2; 3,.4, 5, 65:7; 9,105.11, 12

b. Hemiamyloid (reddish) 8

PARAPHYSES (PIGMENT TOPOGRAPHY)

a. Absent 3

b. In the cell wall 1, 7, 8, 10, 11

c. In the cytoplasm 1, 2, 4, 5,7, 9, 12

d. Restricted to the apical collar or hood 6

PARAPHYSES (PIGMENT IN COTTON BLUE)

a. Not visible 1, 5, 8, 10

b. Visible in all 2, 4, 7, 9, 11, 12

c. Visible only in thickened apices 6

46 ... Landeros, Iturriaga & Guzman-Davalos

PIGMENT (ECTAL EXCIPULUM)

a. Absent 3, 4

b. In the cell wall 1, 2, 5, 6, 7, 8, 9, 10, 11, 12

c. In the cytoplasm 2, 5, 7, 9, 11, 12

PIGMENT (ECTAL EXCIPULUM IN COTTON BLUE)

a. Not visible 1, 5, 6, 8, 9, 10

b. Visible 2, 7, 11, 12

PIGMENT (STIPE OUTER LAYER)

a. Absent 1, 3, 11

b. In the cell wall 2, 5, 6, 7, 8, 9, 10, 12

c. In the cytoplasm 2, 4, 5, 12

PIGMENT (STIPE OUTER LAYER IN COTTON BLUE)

a. Not visible 2, 4, 6, 7, 8, 9, 10, 12

b. Visible in the cytoplasm 5

PIGMENT (APOTHECIAL HYPHAL FASCICLES)

a. Absent 5

b. In the cell wall 1, 2, 6, 8, 9, 10, 12

c. In the cytoplasm 2, 7, 11, 12

d. Restricted to the terminal cells (in the wall cell and cytoplasm) 3

PIGMENT (APOTHECIAL HYPHAL FASCICLES IN COTTON BLUE)

a. Not visible 1, 6, 8, 9, 10

b. Visible in the cytoplasm 7, 11, 12

c. Visible only in the cytoplasm in apical cells 3

PIGMENT (STIPE HYPHAL FASCICLES)

a. Absent 1, 3, 5, 11

b. In the cell wall 2, 6, 7, 8, 9, 10, 12

c. In the cytoplasm 2, 12

Taxonomy of type specimens

Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suec.,

Fasc. 41-42: 37, 1953 Figs 3, 12, 15, 23

= Peziza costata Fr., Nova Acta Reg. Soc. Sci. Uppsal., Ser. 3, 1: 120, 1851

Type: Sweden: Uppland, Uppsala, Botanic Garden, E.P. Fries s.n. Peziza costata

(Lectotype UPS [not seen]; isolectotypes S [studied], K [not seen]).

= Helvella hyperborea Harmaja, Karstenia 18: 57, 1978

Asct 280-370 x 14-18 um. PARAPHYSES 4.5-7.5 um wide at the apex, thin-

walled, hyaline, pale brown to brown in mass, pigment deposits in the cytoplasm

and cell walls, few with brown pigment encrusted on the wall. MEDULLARY

EXCIPULUM hyaline to light brown. ECTAL ExcIPULUM light brown, pigment

deposits in the cell walls. STIPE INNER AND OUTER LAYERS hyaline. APOTHECIAL

HYPHAL FASCICLES 100-230 um long, light brown in mass, pigment deposits in

the cell walls; sTIPE HYPHAL FASCICLES 50-120 um long, hyaline. The brown

Helvella type studies — 1... 47

pigments of the ectal excipulum are not visible in cotton blue or Melzer’s while

those of the paraphysis cell walls are visible in Melzer’s, but inconspicuous in

cotton blue. All tissues rr-.

ADDITIONAL SPECIMENS EXAMINED: FINLAND: LAPLAND, Enontekid, 6 Aug 1985,

J. Vauras 1932 (H); NORTHERN OSTROBOTHNIA, Kuusamo, 4 Aug 1994, T. Ulvinen

FO23453 (OULU); Uusimaa, Lohja, 19 Jul 1998, U. Nummela-Salo & P. Salo 4924 (H).

NORWAY: NorRTHERN Norway, FINNMARK, 19 Aug 1995, I. Kylévuori 95-744 (H).

SWEDEN: UPPLAND, 22 Jun 1948, H. Smith (UPS); 7 July 1948, A. Melderis (UPS); 9

Jul 1948, Exsiccata J.A. Nannfeldt no. 9956 (K).

REMARKS — When Nannfeldt (Lundell & Nannfeldt 1953) transferred Peziza

costata to Helvella, he had to publish a nom. nov., H. costifera, because the

epithet “costata” was already occupied in Helvella by H. costata Schwein. In

the protologue of the replaced synonym Peziza costata, Fries (1851: 120) had

presented two syntypes:

“In Ohio Americae borealis (Lindblom) — etiam copiose in Horto Botanico

Upsaliensi una cum Pez. Helvelloidis var. minori, P. sepulta (Cfr. Summ. Veg. Sc.)

Augusto pluvio. El. Fries, filius [= Elias Petrus Fries]. ... Descripti ad specimina

Upsaliensia viva, a quibus exsiccatis Fungus Ohioensis non distinguendus.”

Nannfeldt (1937: 64) noted that “Peziza costata Fr. is represented in the Uppsala

Museum by beautiful authentical material,’ but Lundell & Nannfeldt (1953)

cited none of Fries’s specimens, noting only that the North American specimen

cited by Fries is lost. Subsequently, a UPS syntype specimen collected by

E.P. Fries was designated as lectotype by Dissing (1966b, as “holotype’), and

accepted by Harmaja (1979, as “lectotype?”) and Abbott & Currah (1997, as

“holotype”) — these errors in type terminology are correctable (McNeill et al.

2006: Art. 9.8).

The lectotype was not available for study because of the bad condition of

the specimen at UPS. Dissing (1966b) described this species with narrower

asci (12-15 um) and paraphyses (3-4 um) than we observed. Our isolectotype

measurements are closer to those made by Abbott & Currah (1997; asci

313-381 x 13.3-17.1 um, paraphyses 3.6-6.2 um at the apex). Helvella costifera

differs from H. acetabulum (L.) Quél. in being less ribbed, with blunt edged

ribs, a non-lacunose stipe, grayish hymenium, and pubescent apothecium.

In H. acetabulum the stipe is highly ribbed to lacunose with sharp edged ribs

(Dissing 1966b), the hymenium has brownish tones (Weber 1972), and the

sterile part of the apothecium is subpubescent to pubescent. See also discussion

under H. griseoalba and H. hyperborea.

Helvella crassitunicata N.S. Weber, Beih. Nova Hedwigia 51: 30,1975 Fics 5, 8, 24

Type: U.S.A.: Washington, Mount Rainier National Park, Narada Falls, on soil along a

path, 10 Aug 1948, A.H. Smith 30052 (Holotype MICH, Barcode 11561).

48 ... Landeros, Iturriaga & Guzman-Davalos

APOTHECIUM sterile surface subpubescent. Stipe glabrous to subpubescent.

Asct 290-340 x 18-24 um. AscosporEs (21.5—)23-26(-28) x (12-)13-14.5

(-15.5) um, ellipsoid to oblong, smooth. PARAPHYSEs of two types: a) 5-7.5 um

wide at the apex, thin-walled, septate, hyaline or light brown; b) 7-9 um wide

at the apex, thick-walled, non-septate, light brown, pigment deposits always

in the cytoplasm. ECTAL EXCIPULUM brown, pigment deposits in the cell wall

and cytoplasm. STIPE OUTER LAYER slightly lighter than the ectal excipulum of

the apothecium, pigment deposits in the cell wall and cytoplasm. APOTHECIAL

HYPHAL FASCICLES < 50 um long, brown pigment deposits in cell walls and

cytoplasm; STIPE HYPHAL FASCICLES < 10 um long, tan pigment deposits in cell

walls and cytoplasm. The brown pigments of the paraphyses, ectal excipulum,

and apothecial hyphal fascicles are visible in cotton blue or Melzer’s; light brown

pigments of the stipe outer layer and hyphal fascicles inconspicuous in cotton

blue or Melzer’s. All tissues rr-.

REMARKS — Helvella crassitunicata and H. leucomelaena (Pers.) Nannf. are the

only two taxa with aporhynchous asci (Fic. 5) (Abbott & Currah 1997). Helvella

leucomelaena is differentiated by mature paraphyses that are never thick-

walled, slightly smaller (20-23(-25) x 10.5-14 um) ascospores, and ascomata

that fruit in spring or early summer. [However, although Weber (1972) cited

H. crassitunicata as fruiting in the fall, Abbott & Currah (1997) noted it

might fruit from May to October.] Also in H. leucomelaena the stipe is poorly

developed or absent and the color of the paraphyses is lost in cotton blue.

Neither Weber (1972) nor we observed the following features cited for

H. crassitunicata by Abbott & Currah (1997): 1) ectal excipulum “pubescent

to densely pubescent’, 2) stipe “finely pubescent to pubescent’, and 3) asci

350-400 x 17-20 um. It is uncertain whether the differences in wall thickness of

the paraphyses should be interpreted as dimorphic (Abbott & Currah 1997) or

developmental (young = thin-walled and mature = thick-walled; Weber 1975).

Since a hymenium develops first as a palisade of paraphyses within which the

asci develop, it would be difficult to have mature asci and immature paraphyses

at the same time (Pfister pers. com.). On the other hand, we observed thin and

thick-walled branches borne on the same paraphyses. See also remarks under

H. pocillum.

Helvella dryadophila Harmaja, Karstenia 17: 58, 1977 Figs:i4, 17, 34,35

Type: NORWAY: prov. Oppland, par. Lom, fjled Hoyrokampen, alt. 1400-1440 m, Dryas

octopetala assoc., 29 Aug 1957 FE. Eckblad, p.p. (Holotype O).

= Helvella verruculosa (Sacc.) Harmaja, Karstenia 18: 57, 1978

AscI 230-280 x 14-18 um. PARAPHYSES 4-7 um wide at the apex, thin-walled,

brown, pigment deposits in the cytoplasm. MEDULLARY EXCIPULUM hyaline.

ECTAL EXCIPULUM brown, pigment deposits in the cell wall and cytoplasm.

Helvella type studies — 1... 49

STIPE INNER LAYER hyaline. STIPE OUTER LAYER brown, pigment deposits in

cell wall and cytoplasm. APOTHECIAL HYPHAL FASCICLES 40-100 um, brown,

pigment deposits in the cell wall and cytoplasm; sTIPE HYPHAL FASCICLES

30-70 um, brown, pigment deposits in cell walls and cytoplasm. The brown

pigments of the paraphyses, ectal excipulum, and apothecial hyphal fascicles

are visible in cotton blue or Melzer’s; brown pigments of the stipe outer layer

and hyphal fascicles inconspicuous in cotton blue. All tissues rr-.

REMARKS — We agree with Abbott & Currah (1997) that H. dryadophila is

a synonym of H. verruculosa, because Harmaja (1977b, 1979) used variable

characters to distinguish them. 1) Both ascomata appear the same: Harmaja’s

own illustration (1977b: 51) (Fic. 5) shows both species with a cup-shaped

apothecium and a ribbed stipe with ribs that never reach the apothecial

sterile surface. 2) For the excipulum, Harmaja cited, “the outermost layer

has somewhat larger cells with slightly thicker wall as H. arctoalpina,” but

gave no measurements, and we did not see such differences. 3) The paler and

homogeneous contents of paraphyses in H. dryadophila did not appear paler to

us; in fact, in cotton blue, the brown pigments are visible in the paraphyses of

both species. The homogeneous content is variable and the pigment tends to be

less homogeneous in immature ascomata. 4) As the presence of a fairly distinct

layer of textura angularis in the excipulum is a generic character for Helvella, its

occurrence should not be used to distinguish between species.

Abbott & Currah (1997) mentioned that H. arctoalpina Harmaja (Harmaja

1977) should also be considered a synonym of H. verruculosa, but as the type

collection of H. arctoalpina is not in O, we unfortunately could not examine

the holotype.

Helvella griseoalba N.S. Weber, Michigan Bot. 11: 162, 1972 Figs 2, 13, 19, 26

Type: U.S.A.: Michigan, Cheboygan Co., Grapevine Point, Douglas Lake, Univ. of

Michigan Biol. Sta., 10 Jun 1968, N.J. Smith 982 (Holotype MICH, Barcode 14379).

Asci 220-265 x 12.5-16 um. MEDULLARY & ECTAL EXCIPULA hyaline. STIPE

INNER AND OUTER LAYERS hyaline. APOTHECIAL HYPHAL FASCICLES 70-120

um long, hyaline with brown terminal cells, pigment deposits in the cell wall

and cytoplasm; STIPE HYPHAL FASCICLES < 30 um long, completely hyaline. The

brown pigments of terminal cells of the apothecial hyphal fascicles are visible in

cotton blue or Melzer’s. All tissues rr-.

REMARKS — Although in fresh ascomata the hymenium and sterile surface of

the apothecium are concolorous (gray to cinnamon; Weber 1972), the dried

sterile surface is lighter, cream to light brown, and the hymenium is slightly

darker, grayish brown. This may be one reason why dry H. griseoalba material

can be easily confused with H. costifera. Both Haffner (1987) and Abbott &

Currah (1997) synonymized H. griseoalba under H. costifera, noting that color

50 ... Landeros, Iturriaga & Guzman-Davalos

differences of the hymenial surface and absence of pigments in the apothecial

ectal excipulum in H. griseoalba were not enough to separate them. However,

we observed additional features in the types of H. costifera and H. griseoalba

that support keeping them as different taxa: 1) H. costifera presents pubescent

hyphal fascicles in the stipe while H. griseoalba does not (glabrous) or has

shorter (subpubescent) hyphal fascicles; 2) H. costifera can have light brown

paraphyses while H. griseoalba has only hyaline ones, and 3) in H. griseoalba

the terminal cells of the apothecial hyphal fascicles have brown pigments that

are visible in cotton blue (Fic. 13) while in H. costifera the light brown pigments

in the cell walls are not visible in cotton blue (Fic. 12).

Helvella hyperborea Harmaja, Karstenia 18: 57, 1978 Fics 11a, 20, 27

Type: FINLAND: prov. Kuusamo, par. Kuusamo, Juuma, western part of the gorge

Jakalavuoma, alt. ca. 205 m, on a shady shelf in the basal part of a steep dolomitic

rock, among the moss Distichium capillaceum (etc.), accompanied by Salix reticulata,

Saxifraga aizoides, S. nivalis, Woodsia glabella, Gerronema albidum, 27 Aug 1970, H.

Harmaja (Holotype H).

= Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suec.,

Fasc. 41-42: 37, 1953

Asct 240-315 x 15-18 um. PARApHYSES thin-walled, hyaline, pale brown to

brown in mass, pigment deposits in cytoplasm and cell walls, some with brown

encrusted pigment on walls. MEDULLARY EXCIPULUM hyaline to light brown.

STIPE INNER AND OUTER LAYERS hyaline. APOTHECIAL HYPHAL FASCICLES

50-160 um long, light brown in mass, pigment deposits in cell walls; sTIPE

HYPHAL FASCICLES 50-100 um long, hyaline. The brown pigments of all

structures are not visible in cotton blue. All tissues rr-.

ADDITIONAL SPECIMENS EXAMINED: FINLAND: LAPLAND, Kemi, 7 Aug 1998, U.

Nummela-Salo & P. Salo 5318 (H); NORTERN SAVONIA, Kuopio, 22 July 1984, J. Vauras

1638F (H); 23 Aug 1987, J. Vauras 2856 (H); VARSINAIS-SUOMEN, 4 Jul 1996, J. Vauras

11192F (H). NORWAY: NorTHERN Norway, Troms, 16 Aug 1992, I. Kyt6vuori 92-352

(H).

REMARKS — Helvella hyperborea was proposed as a new species by Harmaja

(1978) and accepted as distinct by Abbott & Currah (1997). After comparing

its type with H. costifera, including eleven additional specimens from Herbaria

O and UPS (labeled either H. costifera or H. hyperborea), we conclude these

two species are the same. Harmaja (1979, Table 1) listed 12 characters for

differentiating H. hyperborea from H. costifera but explained that only a

combination of all features should be used for its identification. In fact, we

think that his table is an excellent presentation of variability in H. costifera.

Abbott & Currah (1997) used only the ribs of the apothecial sterile surface to

separate the species, distinguishing H. costifera by branched anastomosed ribs

extending up to the marginal area from H. hyperborea with simple unbranched

Helvella type studies — 1... 51

ribs extending onto the basal half. However, both isolectotype and exsiccata

(J.A. Nannfeldt no. 9956) of H. costifera possess the same features used to

identify H. hyperborea (see Fics 23, 27).

We provide below our evaluation of what we consider the five most important

features used by Harmaja (1979) to separate these species.

1) A brown-grey hymenium color that is paler in H. costifera — we have observed

that in fresh material, the color intensity of the hymenium may depend on

whether the specimen is growing in a forest clearing or in a shaded area.

2) Ribs extending onto the sterile surface, especially in dried specimens —

Although Harmaja (1979; Figs. 2-3) illustrated this feature, we found that the

H. costifera isolectotype also has ribs on the sterile surface as, for that matter,

does H. acetabulum.

3) Sterile surface of the apothecium: “with fine but almost always + distinct

hyaline to pale brown pubescence or villosity” (H. costifera) versus “more or

less delicate brown (mostly dark) pubescence usually appearing glabrous to bare

eye” (H. hyperborea) — Under the microscope, the hairs of all studied specimens

have the same color. However, we measured 100-230 um long hyphal fascicles in

the H. costifera type vs. 50-160 um long fascicles in H. hyperborea. The hairs as

measured in the additional specimens range from 50-240 um long, depending

on the maturity of the ascoma. In the stipe hair lengths are similar for both types

— 50-120 um long in H. costifera, 40-100 um long in H. hyperborea — and

remains relatively constant (40-120 um) in the additional specimens.

4) Pigmentation of the paraphyses: “medium (rarely pale) brown wall

encrustation” and more or less brown contents (H. hyperborea) versus hyaline,

inconspicuous or pale brown encrustation and pale brown to practically hyaline

contents (H. costifera) — We observed the same color in the paraphyses for both

species, and in cotton blue the cytoplasmic pigment is inconspicuous; the cell

wall pigment is variable among specimens in both species.

5) Distribution, “middle boreal to lower oroarctic (low alpine), optimal area

apparently northern boreal zone” (H. hyperborea) versus “temperate to middle

(oro)boreal with preference for southern and low-lying areas” (H. costifera)

— Sequences of the 28s large subunit ribosomal DNA from three specimens

identified as H. costifera (unpublished) from Scandinavia (two from northern

boreal specimens and one from the middle boreal region) show only one or two

base differences. Notably, both species were described from the Scandinavian

peninsula.

Helvella leucopus var. populina 1. Arroyo & Calonge, in Calonge, Bol. Soc. Micol.

Madrid 25: 302, 2000 Fics 1, 18b, 28

TyPeE: Spain: Guadalajara, Sigiienza, 8 May 1988, M.J. Rodriguez 888 (Holotype MA

Fungus 22870).

APOTHECIUM sterile surface glabrous, even. STIPE tapering toward the apex,

hollow, glabrous. Asci pleurorhynchous. AscosPorEs verrucose, smooth within

the ascus, hyaline, uniguttulate. PARAPHYSES 3.5-4.5 um wide at the apex, thin-

52 ... Landeros, Iturriaga & Guzman-Davalos

walled, hyaline or brown, pigment deposits in the cytoplasm. MEDULLARY AND

ECTAL EXCIPULA hyaline. STIPE INNER LAYER hyaline. STIPE OUTER LAYER light

brown, pigment deposits in the cytoplasm. HyPHAL FASCICLES absent in the

apothecium and stipe. The brown pigments of the paraphyses are visible and

those of the stipe outer layer are not visible in cotton blue or Melzer’s. Stipe

inner layer rr+; medullary and ectal excipula and stipe outer layer rr-.

REMARKS — Persoon (1822) described H. leucopus Pers. as having a deflexed

apothecium at both ends (lobate), bay to black in color, and a 30-40 mm long

glabrous white stipe. Arroyo & Calonge (1990) distinguished variety populina

(validated by Calonge 2000) by the darker stipe color and larger ascoma

size (apothecium 40-60 x 30-45 mm; stipe 50-120 x 15-35 mm). However,

Moravec (1980) previously cited larger specimens of H. leucopus var. leucopus

(ascomata < 150 mm high) from Slovakia. Ascoma size can be quite variable

and so should be considered with caution; for example, H. crispa (Scop.) Fr. and

H. lacunosa ascomata may range from 50 to 200 mm high. Arroyo & Calonge

(1990) described smooth ascospores and < 12 um wide paraphyses apices,

while we observed verrucose mature ascospores (more evident in cotton blue

or Melzer’s) and narrower paraphyses.

Helvella maculata N.S. Weber, Beih. Nova Hedwigia 51: 27, 1975 Fig. 29

Type: U.S.A.: Idaho, Bonner Co., south side of Hoodoo Mountain, 5 Oct 1968, H.V.

Smith et N.J. Smith (N.J. Smith 2124) (Holotype MICH, Barcode 5635).

APOTHECIUM sterile surface pubescent. AscosporEes 18-22.5 x 10.5-12

um. ECTAL EXCIPULUM yellowish brown, pigment deposits in cell walls and

cytoplasm. STIPE INNER LAYER hyaline. STIPE OUTER LAYER brown, pigment

deposits in cell walls and cytoplasm. APOTHECIAL HYPHAL FASCICLES 107-205

um long, hyaline; sTIPE HYPHAL FASCICLES 60-130 um long, hyaline. The

brown pigments of the paraphyses and ectal excipulum of the apothecium are

inconspicuous and those of the stipe outer layer are very conspicuous in cotton

blue or Melzer’s. All tissues rr-.

REMARKS — We measured smaller ascospores than those cited by Weber (1975,

20-23 x 12-13.5 um). Helvella maculata might be confused with H. crispa,

but the mottled apothecium, grayish stipe ribs, pubescent sterile surface, and

incurved and unrolling margin of H. maculata are diagnostic (Weber 1975;

Abbott & Currah 1997). Furthermore, the brownish pigment of the stipe

outer layer is evident in cotton blue in H. maculata but not in H. crispa. For

Weber (1975) ascospore size could help differentiate the two species, but the

size range we observed for H. maculata overlaps with that seen in H. crispa

(16-20.5 x 9.8-12 um). Another similar species, H. fusca Gillet, has a dark

red-brown hymenium, distinctly ribbed sterile surface with some ribs reaching

the apothecium margin, and vernal fructification associated with Populus,

Helvella type studies — 1... 53

Fics. 23-29: Ascomata of type specimens (except when indicated) of Helvella. 23: H. costifera

(Exsiccata J.A. Nannfeldt no. 9956). 24: H. crassitunicata. 25: H. dryadophila. 26: H. griseoalba.

27: H. hyperborea. 28: H. leucopus var. populina. 29: H. maculata. Scale bar: 1 cm.

while H. maculata fruits in autumn and is associated with conifers (Weber

1975); furthermore, the apothecium sterile surface of H. fusca is subpubescent.

According to Abbott & Currah (1997), H. maculata could also be confused

with H. lacunosa, which differs in having an apothecium margin attached to the

stipe, black hymenium, and glabrous sterile surface of apothecium and stipe.

54 ... Landeros, Iturriaga & Guzman-Davalos

Helvella paraphysitorquata 1. Arroyo & Calonge, in Calonge & Arroyo, Mycotaxon

39: 210, 1990 FIGS 7, 16, 18c, 30

TyPE: Spain: within the province of Teruel, Albarracin, growing in soil under Populus, 3

Jun 1988, I. Arroyo & ED. Calonge (Holotype MA-Fungi 24512).

APOTHECIUM ‘sterile surface pubescent, even. STIPE pubescent. ASCI

pleurorhynchous. AscosporEs 16.5-19 x 11-13.5 um, ovoid to ellipsoid,

verrucose, smooth within the ascus. PARAPHYSES 5-9 um wide at the apex,

thin-walled, hyaline, with a brown thick-walled cap or collar at the apex.

MEDULLARY EXCIPULUM hyaline. ECTAL EXCIPULUM hyaline to light brown,

pigment deposits in cell walls. StipE INNER AND OUTER LAYERS hyaline to

light brown, pigment deposits in cell walls. APOTHECIAL HYPHAL FASCICLES

60-220 um long, hyaline to light brown in mass, pigment deposits in the cell

wall; sTIPE HYPHAL FASCICLES 60-140 um, hyaline to light brown in mass,

pigment deposits in the cell wall. The brown pigments of the paraphyses are

visible in cotton blue, and pigments of the ectal excipulum, apothecial hyphal

fascicles, and stipe outer layer and hyphal fascicles are not visible in cotton blue

or Melzer’s. All tissues rr-.

REMARKS — Calonge & Arroyo (1990) described H. paraphysitorquata with

smooth ascospores, but we also found ornamented ascospores on the hymenial

surface, stipe, and basal mycelium. We observed narrower paraphyses than

those recorded by Calonge & Arroyo (1990; 7-10(-15) um), possibly due

to their rehydration in 2% KOH or 10% NH,. Helvella paraphysitorquata

is morphologically similar to H. pezizoides Afzel., which differs in its totally

black even stipe and paraphyses lacking the brown collar. On the other hand,

in a specimen determined as H. macropus from Mexico (J.M. Rodriguez

Canseco 12, IBUG), we also observed a few or occasional brown capped or

collared paraphyses, so apparently collared paraphyses are not unique to

H. paraphysitorquata.

Helvella pocillum Harmaja, Karstenia 15: 30, 1976 FIGS 4, 31

TyPeE: Sweden: prov. Torne Lappmark, par. Jukkasjarvi, ffeld Laktatjakko, on bare soil, 17

Aug 1946, L. Holm 472 (Holotype UPS).

APOTHECIUM sterile surface subpubescent to pubescent, concolorous with the

hymenial surface, stipe ribs only reaching the apothecium basis. Stipe 1-5 x

1-3 mm, solid, costate, ribs blunt, pubescent, concolorous with the hymenial

surface. Asc1 320-380 x 18-24 um, pleurorhynchous. AscosporEs 23-28

x 12-15.5 um, smooth. PARAPHysEs thin-walled. MEDULLARY EXCIPULUM

hyaline. ECTAL EXCIPULUM brown, pigment deposits in cytoplasm. STIPE INNER

LAYER hyaline. STIPE OUTER LAYER light brown, pigment deposits in cell walls.

APOTHECIAL HYPHAL FASCICLES 40-100 um long, brown, pigment deposits in

cytoplasm; STIPE HYPHAL FASCICLES 50-80 um, hyaline to light brown in mass,

Helvella type studies — 1... 55

pigment deposits in cell walls. The brown pigments of the paraphyses, ectal

excipulum, and apothecial hyphal fascicles are visible and those of the stipe

outer layer and hyphal fascicles are not visible in cotton blue or Melzer’s. All

tissues rr-.

REMARKS — Helvella pocillum is characterized by its very small ascoma;

only H. rivularis Dissing & Sivertsen is as small, but with an even stipe. For

Harmaja (1976) H. pocillum could be separated from H. crassitunicata by its

larger ascospores, the width of the paraphyses (we found no differences in

either of these characters), and the absence of thick-walled paraphyses. Besides,

H. crassitunicata has aporhynchous asci (Fic. 5) and the sterile surfaces of

apothecium and stipe are glabrous to subpubescent. Another similar species is

H. leucomelaena, also with aporhynchous asci.

Helvella queletii Bres., Fungi trident. 1(3): 39, 1882 FIG. 32

Type: Italy: Bosee di lance sopra Terzolaj, May 1882, J. Bresadola (Holotype S).

= Helvella solitaria P. Karst., Bidr. Kann. Finl. Nat. Folk 19: 37, 1871

STIPE solid, costate, ribs blunt. Asc1 260-310 x 15-20 um, pleurorhynchous.

Ascospores 17.5-21.5 x 10.5-12.5 um, broadly ellipsoidal, smooth.

PARAPHYSES 3.5-7 um wide at the apex, thin-walled, brown, pigment deposits

in the cytoplasm. MEDULLARY EXCIPULUM hyaline. ECTAL EXCIPULUM brown,

pigment deposits in cell walls and cytoplasm. STIPE INNER LAYER hyaline. STIPE

OUTER LAYER light brown, pigment deposits in cell walls. APOTHECIAL HYPHAL

FASCICLES 25-60 um long, light brown in mass, pigment deposits in the cell

wall; STIPE HYPHAL FASCICLES 40-100 um, light brown, pigment deposits in

cell walls. The brown pigments of the paraphyses are visible and pigments of

the ectal excipulum, stipe outer layer, and apothecial and stipe hyphal fascicles

are not visible in cotton blue or Melzer’s. All tissues rr-.

ADDITIONAL SPECIMENS EXAMINED: SPAIN: MADRID, Madrid, 7 Apr 2001, A. Gonzalez,

J.C. Campos et al., G.P. 1395 (MA-Fungi 73836); CASTILE AND LEON, Valladolid, 31

Mar 2001, Herrera de Duero 1294 (MA Fungi 54822).

REMARKS — Bresadola (1882), who described H. queletii as having a cup-

shaped apothecium, sulcate-lacunose stipe (“costate” according to Weber 1972),

and ascospores measuring 17-20 x 12 um, noted its proximity to H. lacunosa

but without mentioning the differences with H. solitaria. Dissing (1966b)

differentiated H. queletii from H. solitaria by the following: 1) ascospore size —

17-19.1-21 x 11-12.3-13.5 um (H. queletii) vs. 19-21.7-24 x 12-13.1-15 um

(H. solitaria); 2) ascoma size — 20-80 mm broad apothecium, 20-60 mm

high stipe (H. queletii) vs. 10-40 mm broad apothecium, 10-25 mm high

stipe (H. solitaria); 3) hymenium color (pale brownish to dark greyish-brown

(H. queletii) vs. greyish to greyish-brown (H. solitaria); 4) stipe rib number

— 4-7 in H. queletii vs. 2-5 in H. solitaria; 5) fruiting time — April-October in

H. queletii vs. February—June in H. solitaria.

56 ... Landeros, Iturriaga & Guzman-Davalos

Harmaja (1977a) suggested that H. queletii be recognized as a synonym of

H. solitaria based on the variability of ascoma size, color, pubescence, and stipe

rib number. Abbott & Currah (1997) also considered them synonymous (but

see below under H. solitaria). The following list summarizes our observations

of both holotypes and other original descriptions.

1) The H. solitaria holotype has smooth ascospores and a spore range (18-20 x

9.5-12 um) matching that of the H. queletii holotype and as reported by Dissing

(1966b). We also found smaller (15-17.5 x 9.5-11.5 um) verrucose ascospores

more closely matching measurements given by Karsten (1871, 14-16 x 10 um)

but never matching that cited by Dissing (1966b).

2) Ascoma size, highly variable in Helvella, is not a good taxonomic criterion for

a widespread species,

3) Ascoma color is highly variable in H. queletii with tones very similar to

H. solitaria, as noted in Dissing (1966a) and Dissing & Nannfeldt (1966),

4) Among materials cited as H. queletii by Dissing (1966b) is a specimen

collected in March (Velenovsky s.n., Mar 1923, PRC), while among his

H. solitaria materials are specimens collected in July (Eftesal s.n., 8 Jul 1956, O;

Berg s.n., 15 Jul 1957, O), August (Bresadola s.n., Aug 1893, S; Hakelier s.n., 28

Aug 1962, UPS), September (P. Karsten s.n., 21 Sep 1866, H), October (Toft s.n.,

21 Oct 1965, C), and November (Bresadola s.n., Nov 1896, S; Arwidsson s.n.,

1 Nov 1943, S). Thus, the time of fruiting is not diagnostic.

5) Both holotypes are anatomically similar, except that the brown cytoplasmic

pigment in the paraphyses is lighter in H. solitaria.

Helvella robusta S.P. Abbott, in Abbott & Currah, Mycotaxon 33: 242, 1988

Fics 10b, 33

Type: CANADA: Calgary, Nose Hill area, 30 Aug 1972, R.M. Danielson 459 (Holotype

DAOM-143869).

APOTHECIUM irregularly cupulate with a large central depression to irregularly

bi-lobed, margin free, covering the stipe; sterile surface subpubescent

to pubescent, ribbed, ribs sharp. Stipe gradually expanding toward the

apothecium, lacunose, ribs sharp, subpubescent to pubescent. PARAPHYSES

hyaline to light brown, pigment deposits in cell walls. MEDULLARY EXCIPULUM

hyaline. ECTAL EXCIPULUM light brown, pigment deposits in cell walls. STIPE

INNER LAYER hyaline. STIPE OUTER LAYER light brown, pigment deposits in

cell walls. APOTHECIAL HYPHAL FASCICLES 38-80 um long, hyaline to light

brown, pigment deposits in cell walls; srrpE HYPHAL FASCICLES 40-90 um long,

hyaline to light brown, pigment deposits in cell walls. The brown pigment of the

paraphyses, ectal excipulum, apothecial hyphal fascicles, and stipe outer layer is

not visible in cotton blue or Melzer’s. Medullary excipulum and stipe inner and

outer layers rr+; ectal excipulum rr-.

Helvella type studies — 1... 57

RemARKS — Abbott & Currah (1988, 1997) described H. robusta with

apothecium that is irregularly cupulate or centrally depressed with reflexed

margins; in addition to this type, we observed an irregularly bi-lobed

apothecium covering the stipe, with the apothecial edges curved towards the

stipe, thus fully exposing the hymenium. The peculiar shape of the ascoma

makes H. robusta easy to recognize, although Abbott & Currah (1988) noted

that it could be confused with some species of section Acetabulum, although

members of that section have a well-defined cup-shaped apothecium in all stages

of development. In addition to its peculiar apothecial shape, H. robusta has a

strong reddish reaction in the apothecial medullary excipulum and stipe inner

layer and moderately reddish reaction in the stipe outer layer in Melzers.

Helvella solitaria P. Karst., Bidr. Kann. Finl. Nat. Folk 19: 37,1871 Fras 9, 10a, 34

Type: Finland: Mustiala, in horto, ad marg. rivula, 21 Sep 1866, P. Karsten PAK 3288

(Holotype H).

= Helvella queletii Bres., Fungi trident. 1(3): 39, 1883

ASCOSPORES a) smooth: 18-20 x 9.5-12 um, broadly ellipsoidal; b) verrucose:

15-17.5 x 9.5-11.5 um, ellipsoidal. PARAPHysES 4-7 um wide at the apex,

thin-walled, light brown, pigment deposits in the cytoplasm. MEDULLARY

EXCIPULUM hyaline. ECTAL EXCIPULUM brown, pigment deposits in cell walls

and cytoplasm. STIPE INNER LAYER hyaline. STIPE OUTER LAYER light brown,

pigment deposits in cell walls. APOTHECIAL HYPHAL FASCICLES 40-90 um long,

light brown, pigment deposits in cell walls; strpE HYPHAL FASCICLES 35-90

um, light brown in mass, pigment deposits in cell walls. The brown pigments

of the paraphyses are visible, and pigments of the ectal excipulum, stipe outer

layer, and apothecial and stipe hyphal fascicles are not visible in cotton blue or

Melzer’s. All tissues rr-.

REMARKS — Helvella solitaria can be confused with H. pocillum, H. ulvinenii,

and H. verruculosa because of its cup-shaped apothecium, costate stipe with

ribs extending only to the base or a short distance onto the apothecium, and

pleurorhynchous asci. Helvella pocillum has larger ascospores (22-30 x 13-17

um). Helvella ulvinenii has brown, sepia to black apothecial sterile surface (never

with grayish tones), brown pigment of the apothecial hyphal fascicles visible in

cotton blue and hyaline stipe hyphal fascicles. Helvella verruculosa differs in

the visibility of the brown cytoplasmic pigments in the ectal excipulum and

apothecial hyphal fascicles in cotton blue, and its arctic and alpine distribution

contrasts with the boreal and temperate distribution for H. solitaria (Abbott &

Currah 1997).

Paraphysis widths cited here match those by Karsten (1871) but not those

by Abbott & Currah (1997), who cited paraphyses with 7-8 um wide apices.

Karsten (1871) gave 14-16 x 10 um for H. solitaria ascospores but did not

58 ... Landeros, Iturriaga & Guzman-Davalos

mention whether they were smooth or ornamented. We agree with Harmaja

(1977a,b) that mature and immature ascospore sizes differ in some species, but

in view of the greater size variation in H. solitaria, we consider it important to

list the sizes of both immature and mature ascospores.

Although Dissing (1966b) recognized H. solitaria and H. queletii as different

species, we agree with Harmaja (1977a) that they are synonyms, with the name

H. solitaria having priority (see discussion under H. queletii).

According to Abbott & Currah (1997), H. solitaria sensu Dissing (1966b)

corresponds to H. leucomelaena. We do not agree with this because of the

following: 1) H. leucomelaena has aporhynchous asci; Dissing (1966b) did not

mention the ascus base, and 2) the specimens studied by Dissing do not seem

to correspond to a single species. Harmaja (1977a) designated one Dissing

H. solitaria specimen as holotype of H. confusa Harmaja (also considered a

synonym under H. leucomelaena by Abbott & Currah, 1997), while other Dissing

specimens correspond to different species, among them H. leucomelaena and

H. solitaria (Harmaja 1979). As Dissing’s concept of H. solitaria was very broad,

it cannot be attributed to a particular taxon.

Helvella subglabra N.S. Weber, Michigan Bot. 11: 179, 1972 Fics 6, 11b, 35

Type: U.S.A.: Michigan, Washtenaw, Co., Stinchfield Woods, near Dexter, 13 Oct 1968,

N.J. Smith 2145 (Holotype MICH, Barcode 14381).

Asci 224-324 x 15-17 um, pleurorhynchous. Ascospores 15-19 x 10-11.5

um, smooth, few verrucose. PARAPHYSES thin-walled, hyaline to pale brown,

pigment deposits in the cell wall. Stipe OUTER LAYER brown, pigment deposits

in the cell wall. APOTHECIAL & STIPE HYPHAL FASCICLES < 30 um long,

scattered; hyaline to brown, pigment deposits in cell walls. The brown pigments

of all structures are not visible in cotton blue. Medullary excipulum and stipe

inner layer rr+; ectal excipulum and stipe outer layer rr-.

REMARKS — Haffner (1987) and Abbott & Currah (1997) synonymized

H. subglabra under H. atra J. Konig. They considered that the differences listed

by Weber (1972) were not enough to separate them, because other species

(H. lacunosa, H. albella, H. latispora Boud.) also accommodate such variability.

Distinctions noted by Weber (1972) are: 1) hymenium surface color— dark

gray to brownish gray or drab (H. subglabra) vs. black to grayish black (H. atra);

2) color of stipe and apothecial sterile surface — drab to gray (H. subglabra)

vs. black or dark gray (H. atra); 3) pigments in cells of the ectal excipulum

— scattered hyphal cells with brown walls (H. subglabra) vs. most hyphal

cells with brown walls (H. atra); and 4) apothecial sterile surface texture

— subpubescent (H. subglabra) vs. completely glabrous (H. atra). We add

here two more characters that separate H. subglabra from H. atra: 1) tissues

in Melzer’s — ascoma negative (H. atra) vs. apothecial medullary excipulum

Helvella type studies — 1... 59

Fics. 30-37: Ascomata of type material of Helvella. 30: H. paraphysitorquata. 31: H. pocillum.

32: H. queletii. 33: H. robusta. 34: H. solitaria (scanned image of the picture contained in the type

specimen). 35: H. subglabra. 36: H. ulvinenii. 37: H. verruculosa. Scale bar: 1 cm.

weakly reddish and stipe inner layer strongly reddish (H. subglabra); 2) the

brown cell wall pigment in the paraphyses, ectal excipulum, and stipe outer

layer is visible in cotton blue in H. atra.

Helvella ulvinenii Harmaja, Karstenia 19: 42, 1979 Fics 22, 36

TyPE: Finland: prv. Enontekion Lappi, par. Enonteki6, Kilpisjarvi, NE slope of W peak of

fjeld Pikku- Malla, alt. ca. 650 m, H. Harmaja, 25 Aug 1979 (Holotype H).

60 ... Landeros, Iturriaga & Guzman-Davalos

APOTHECIUM Sterile surface subpubescent to pubescent. STIPE solid, costate,

ribs blunt, pubescent. Ascr 240-300 x 14-17 um. Ascospores 14.5-18.5 x

10.5-12(-13.5) um, broadly ellipsoid, smooth and verrucose. STIPE INNER

& OUTER LAYERS hyaline. APOTHECIAL HYPHAL FASCICLES 40-105 um long,

brown, pigment deposits in the cytoplasm; STIPE HYPHAL FASCICLES 60-140

um, hyaline. The brown pigments of the paraphyses, ectal excipulum, and

apothecial hyphal fascicles are visible in cotton blue or Melzer’s. All tissues

rr-.

REMARKS — Harmaja (1979) did not mention ascospore ornamentation for

H. ulvinenii; we observed both smooth (immature), and verrucose (mature)

ascospores with the same size and shape. Helvella ulvinenii shares cup-shaped

apothecia and costate stipes with ribs that do not reach the sterile apothecial

surface with H. solitaria and H. verruculosa. We agree with Harmaja (1979)

that H. ulvinenii is distinguished from H. solitaria by 1) a darker sterile surface

that is never grayish, 2) shorter ascospores, and 3) ectal excipular hyphae with

distinctly dark pigments, present only in the cell wall and not in the cytoplasm.

Additional distinctions include: 1) the stipe outer layer and hyphal fascicles

are hyaline in H. ulvinenii and light brownish (on the cell walls) in H. solitaria,

and 2) the apothecial hyphal fascicle pigments are deposited in the wall in

H. solitaria and occur both in the cytoplasm and in the cell wall in H. ulvinenii.

Harmaja (1979) differentiated H. verruculosa (as H. dryadophila) from

H. ulvinenii based on 1) apothecial shape, 2) longer ascospores, and 3) very

long paraphysis end cells. Although we did measure longer ascospores (17.5-22

x 10.5-13 um) for H. verruculosa, both types had cup-shaped apothecia and

ribbed stipes and the paraphysis end cells appeared the same. In addition, we

observed that H. verruculosa has pigmented stipe hyphal fascicles. Probably,

H. ulvinenii is a variety of H. verruculosa.

The H. ulvinenii type demonstrates particularly well the differences between

the structure of the ectal excipulum and apothecial hyphal fascicles and of the

stipe outer layer and hyphal fascicles; cells are pigmented in the apothecium

and hyaline in the stipe (Fic. 22).

Helvella verruculosa (Sacc.) Harmaja, Karstenia 18: 57, 1978 Fic. 37

Type: Russia: Madagan Obl., Chukotski Peninsula, Arakamchene Island by the Bearing

Strait, Stony Hills, 11 Aug ca. 1885, Wright, Herb. U.S.N. Pacif. Expl. Exped. No. 369

(Holotype K, Isotype FH, both studied).

= Geopyxis verruculosa Sacc., Sylloge fungorum 8: 68, 1889

= Helvella dryadophila Harmaja, Karstenia 17: 58, 1977

PARAPHYSES 4-8 um wide at the apex, thin-walled, brown, pigment deposits

in the cytoplasm. MEDULLARY EXCIPULUM hyaline. ECTAL EXCIPULUM brown,

pigment deposits in cell walls and cytoplasm. STIPE INNER LAYER hyaline.

Helvella type studies — 1... 61

STIPE OUTER LAYER brown, pigment deposits in cell walls and cytoplasm.

APOTHECIAL HYPHAL FASCICLES 70-150 um, brown, pigment deposits in cell

walls and cytoplasm; sTIPE HYPHAL FASCICLES 50-130 um, brown, pigment

deposits in cell walls and cytoplasm. The brown pigments of the paraphyses,

ectal excipulum, and apothecial hyphal fascicles are visible and those of the

stipe outer layer and hyphal fascicles are not visible in cotton blue or Melzer’s.

All tissues rr-.

RemMarKs — Abbott & Currah (1997) described this species with a “villose

margin” of the apothecium (sometimes with distinct white hairs), the stipe as

having few chambers, and with stipe ribs extending onto basal half of apothecial

sterile surface. We did not observe the marginal hairs, perhaps because the

hairs have been lost due to age or handling. Furthermore, we observed neither

stipe chambers nor ribs on the sterile surface of the apothecium. In fact,

H. verruculosa is more easily confused with H. solitaria and H. ulvinenii, which

do not have ribs on the sterile surface (see differences under the remarks of

these species) than with H. acetabulum, H. costifera, and H. griseoalba.

Acknowledgments

We appreciate the thorough reading and helpful comments from Donald H. Pfister

(Harvard University, USA) and Trond K. Schumacher (University of Oslo, Norway)

who acted as presubmission reviewers. Shaun Pennycook, Mycotaxon Nomenclature

Editor, kindly helped us to make clearer the H. costifera nomenclatural remarks. We

especially thank Lorelei Norvell, Mycotaxon Editor-in-Chief, who made an excellent

work to improve the text. Thanks are given to the curators of the herbaria DAOM, FH,

H, K, MA, MICH, S and UPS for the loan of type specimens. The first author thanks the

Universidad Auténoma de Querétaro for its support and CONACYT for a scholarship

grant for his doctoral studies. Virginia Ramirez Cruz (Universidad de Guadalajara,

Mexico) is acknowledged for her valuable help inking the drawings, Greg Bonito (Duke

University, USA) for literature on Helvella, and Dick Korf for his advice on the typification

of H. costifera. Funds were obtained from Universidad de Guadalajara (projects 72640,

88682, 108721, PIFI-2008-2009).

Literature cited

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northwestern North America. Mycotaxon 62: 1-125.

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populina Arroyo & Calonge, nuevas para Espajia. Bol. Soc. Micol. Madrid 14: 197-199.

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the ascus wall. Mycotaxon 29: 399-450.

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62 ... Landeros, Iturriaga & Guzman-Davalos

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Dissing H, Nannfeldt JA. 1966. Helvella cupuliformis sp. nov., H. villosa (Hedw. ex O. Kuntze) comb.

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valle de México. Bol. Soc. Bot. Méx. 29: 1-18.

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Mykologie 7: 1-165.

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17: 40-44.

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Karstenia 17: 45-58.

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world. 8" edition. New York Botanical Garden. New York. 693 p.

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Folk 19: 1-263.

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Publishing House. New Delhi.

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fungi. 10" ed. CABI Bioscience, Oxon. 771 p.

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278-285. http://dx.doi.org/10.2307/3761373

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+ 2 plates.

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MYCOTAXON

http://dx.doi.org/10.5248/119.65

Volume 119, pp. 65-81 January-March 2012

Psilocybe s.s. in Thailand: four new species and

a review of previously recorded species

GASTON GUZMAN”, FLORENCIA RAMIREZ GUILLEN’,

KEVIN D. HyDE*? & SAMANTHA C. KARUNARATHNA”*

"Instituto de Ecologia, Apartado Postal 63, Xalapa 91070, Veracruz, Mexico

? School of Science, Mae Fah Luang University,

333 Moo 1, Tasud, Muang, Chiang Rai 57100, Thailand

*Botany and Microbiology Department, College of Science, King Saud University,

Riyadh, Saudi Arabia

*Mushroom Research Foundation, 128 M.3 Ban Pa Deng T: Pa Pae,

A. Mae Taeng, Chiang Mai 50150, Thailand

* CORRESPONDENCE TO: gaston.guzman@inecol.edu.mx

Axsstract — Psilocybe deconicoides, P. cubensis, P. magnispora, PB. samuiensis, and

P. thailandensis (previously known from Thailand) are revisited, and P thaiaerugineomaculans,

P. thaicordispora, P. thaiduplicatocystidiata, and P. thaizapoteca are described as new species.

These new species are bluing and belong to sections Cordisporae, Stuntzae, and Zapotecorum.

Following the recent conservation of Psilocybe as the generic name for bluing species,

P. deconicoides, which does not blue upon bruising, is transferred to Deconica, while the bluing

taxa P. cubensis (sect. Cubensae), P. magnispora and P. thailandensis (sect. Neocaledonicae),

and P. samuiensis (sect. Mexicanae) remain in Psilocybe.

Key worps — hallucinogenic mushrooms, richness mycobiota, Strophariaceae, tropics

Introduction

The hallucinogenic fungal species in Thailand, as in most tropical countries,

are poorly known, which is in direct contrast with the large fungal diversity

that occurs throughout the tropics. Moreover, with considerable destruction

of tropical habitats for use as agricultural or cattle farms, many species will

likely disappear before being documented. We have been surveying several

mushroom genera in northern ‘Thailand (e.g. Le et al. 2007, Sanmee et al. 2008,

Wannathes et al. 2009, Zhao et al. 2010, Karunarathna et al. 2011); but studies

of Psilocybe are rare. Despite the recreational use of hallucinogenic mushrooms

in the country (see Allen & Merlin 1992), we have found only a few references

66 ... Guzman & al.

in papers by Heim (1958, 1962), Guzman et al. (1993), Chamratpan (2003),

Klinhom et al. (2003), and Horak et al. (2009). Fortunately, Thailand now has

a mycological research station, Mushroom Research Centre, where local and

visiting mycologists can explore, gather and study fungi. We report Psilocybe

spp. collected in 2010 in the area surrounding the Research Centre. We

describe four species as new and discuss five species previously reported from

Thailand.

Materials & methods

For microscopic study, thin hand sections of the basidiome, mainly in the pileus,

including context and lamellae, were rehydrated with 96% alcohol and then mounted in

5% KOH and 1% Congo Red solutions (the latter added to tissues previously mounted

in KOH). Basidiospore dimensions indicate length and width (face view) and thickness

(side view). At least 25 measurements were taken for each microscopic feature. All

specimens are deposited at Mae Fah Luang University herbarium (MFLU), Chiang

Rai, Thailand, with duplicates in the Fungus Collection at XAL Herbarium, at Xalapa,

Mexico.

Results

Five species of Psilocybe s.l. known from Thailand are discussed here

(P. cubensis, P. deconicoides, P. magnispora, P. samuiensis, and P. thailandensis)

and four new species are proposed.

Psilocybe cubensis (Earle) Singer, Sydowia 2: 37, 1948

= Stropharia cubensis Earle, Estudios Agronémicos, Cuba 1: 240, 1906

This is an important hallucinogenic mushroom with a pantropical

distribution, although the records from Africa (Cullinan et al. 1945, Vedcourt

& Trump 1969) are still under study to determine whether they represent

P. aquamarina (Pegler) Guzman (= Stropharia aquamarina Pegler), as

discussed by Heim (1958, 1962), Pegler (1977), and Guzman (1995). The Thai

material (identified as Stropharia cubensis) studied by Heim, based on two

collections (one by Apaiwongse, the second by Champagne) from the Bangkok

region in 1957, agree well with the Mexican specimens studied by Heim.

Recently Chamratpan (2003) and Klinhom et al. (2003) reported P cubensis

from Thailand as a “magic” and/or “soporific” mushroom. This coprophilous

fungus, common on cattle and horse dung in meadows, probably grows also in

Thailand on wild mammal dung. The African mentioned above was reported by

Vedcourt & Trump (1969) on monkey dung, and Guzman & Ramirez-Guillén

(2010) reported the closely related P subcubensis Guzman on rhinoceros dung

in Nepal. Nonetheless, Guzman did not find P. cubensis nor P. subcubensis

in meadows explored in Thailand in 2010, where he found only Copelandia

cyanescens (Berk. & Broome) Singer and Panaeolus spp.

Psilocybe s.s. in Thailand ... 67

Psilocybe cubensis as shown in the key (below) has a well developed annulus,

hyaline pleurocystidia, and thick walled basidiospores — (12—)13-15(-17) x

8-10(-11) x 7-9 um — that are subhexagonal in face-view and subellipsoid in

side-view (Guzman 1983).

Deconica deconicoides (E. Horak, Guzman & Desjardin) Guzman, comb. noy.

MycosBank MB563261 FIGURES 1-3

= Psilocybe deconicoides E. Horak, Guzman & Desjardin, Sydowia 61(1): 32, 2009

According to Horak et al. (2009) this is a non-bluing species, growing on rotting

branches in Chiang Mai Prov. characterized by small mycenoid basidiomata

with broadly adnate to decurrent lamellae. The thick-walled basidiospores

— (5.5-)6-6.5 x (4.5-)5-5.5 x 3.5-4 um — are rhomboid in face-view and

ellipsoid in side-view. Basidia 17-23 x (5-)6.5-7 um, 4-spored, hyaline,

ventricose-clavate. Pleurocystidia absent. Cheilocystidia (14.5-)16-19(-24) x

5-6(-7) um, polymorphic, ventricose, clavate, fusoid or sublageniform.

MATERIAL STUDIED — THAILAND. CHIANG Mat PrRov., Chiang Mai, Dui Suthep, 19

km marker, June 6, 2002, Horak 10156 (isotype XAL).

REMARKS: Moncalvo et al. (2002) showed two Psilocybe s.l. clades, one

containing the bluing species and the other containing the non-bluing species.

Redhead et al. (2007) proposed to conserve the name Psilocybe for the bluing

clade and accommodate the non-bluing clade under Deconica. This proposal

was supported by the Nomenclature Committee for Fungi (Norvell 2010)

and accepted on July 31, 2011, by the 13" International Botanical Congress,

Melbourne, necessitating the transfer of P. deconicoides to Deconica.

Psilocybe magnispora E. Horak, Guzman & Desjardin, Sydowia 61: 25, 2009

FIGURES 4-7

This bluing species grows on old rotting elephant dung in a tropical

montane forest and is only known from Pong Chang Province at Kha Yai

(Horak et al. 2009). The basidiomata have a convex to obtusely papillate pileus

(20-)30-45 mm diam, adnate lamellae, and stipe 35-60 x 2.5-4(-6) mm,

with a submembranous annulus. Basidiospores are 7-8(-8.5) x (5.5-)6-7 x

5-5.5 um, thick walled, and rhomboid in face-view. Pleurocystidia 21-28(-31)

x (7-)9-13 um, more or less common, ventricose-claviform or subfusoid,

sometimes submucronate, hyaline, frequently with refringent incrustations, as

chrysocystidia. Cheilocystidia (13—) 16-26(-30) x 5-7(-10) um, polymorphous,

fusoid, ventricose or sublageniform, hyaline. The chrysocystidia and rhomboid

spores place P. magnispora in sect. Neocaledonicae Guzman.

MATERIAL STUDIED — THAILAND, Kua Yat, Pong Chang, Princess Trail, July 6, 2002,

Horak 10171 (isotype XAL).

68 ... Guzman & al.

O) :

FIGURES 1-9. Deconica deconicoides (type) - 1. basidiomata; 2. basidiospores; 3. cheilocystidia.

Psilocybe magnispora (type) — 4. basidiomata; 5. basidiospores; 6. pleurocystidia; (chrysocystidia);

7. cheilocystidia. P. samuiensis (type) — 8. basidiomata; 9. basidiospores. Scale bars: 1, 8 = 10 mm;

4 = 20 mm; 2-3, 5-7 = 5.5 um; 9 = 10 um.

Psilocybe s.s. in Thailand ... 69

Psilocybe samuiensis Guzman, Bandala & J.W. Allen, Mycotaxon 46: 156, 1993

FIGURES 8-11

This species, known only from ‘Thailand, grows on rich soil in meadows

(Guzman et al. 1993). It presents basidiomata like P. mexicana R. Heim, with

pileus 7-15 mm diam., lamellae adnexed and stipe 40-65 x 1.5-2 mm, without

annulus. Basidiospores 10-13 x 6-8 um, rhomboid to subrhomboid in face-

view, thick-walled. Pleurocystidia 16-20 x 5-6.5 um, scattered, sublageniform,

hyaline and cheilocystidia 18-28(-30) x (5-)5.5-7(-8) um, sublageniform,

sometimes branched.

Basidiospore morphology supports P. samuiensis within sect. Mexicanae

Guzman. Molecular analyses by Borovicka et al. (2011) that show this species

as separate from sects. Cyanescens and Semilanceatae support sect. Mexicanae

as independent.

MATERIAL STUDIED — THAILAND, SuRAT THANI PRov., Koh Samui, Thailand, 2 km

west of Ban Hua Thanon, August 8, 1991, Allen s.n. (holotype XAL).

Psilocybe thailandensis E. Horak, Guzman & Desjardin, Sydowia 61: 30, 2009

FIGURES 12-15

This bluing species grows on rotten wood in a tropical montane forest

(Horak et al. 2009). Pileus (4-)6-15 mm, conic, brown to pale brown, turning

paler after drying, with veil remnants at the margin. Lamellae adnate. Stipe

(10-)15-30 x 1-2 mm, subbulbous, with a fibrillose-membranaceous annulus,

sometimes ephemeral. Basidiospores (4.5-)5-6(-6.5) x 4.5-5.5 x 3-4 um,

rhomboid in face-view, ellipsoid in side-view, thick-walled. Pleurocystidia of

the chrysocystidia type with a refringent content, common, 16-25(-30) x (6.5-

)7-9(-10), broadly clavate or vesiculose. Cheilocystidia (11-)20-30 x 3.5-6 um,

hyaline, sublageniform, ventricose-rostrate or irregularly slender subfusoid. Its

chrysocystidia and rhomboid thick-walled spores place P. thailandensis in sect.

Neocaledonicae with P. magnispora.

MATERIAL STUDIED — THAILAND, CutaAnc Mat Prov., Doi Suthep, Monthatharn

Waterfall Trail, July 4, 2002, Horak 10128 (isotype XAL); trail to Huai Kok, Horak 10115

(paratype XAL).

Psilocybe thaiaerugineomaculans Guzman, Karunarathna & Ram.-Guill., sp. nov.

MycoBank MB 561575 FIGURES 30-35

Differs from Psilocybe aerugineomaculans in producing pyriform yellowish brown

pseudocystidia from the hymenophoral trama.

Type: Thailand, Chiang Mai Prov., Chiang Mai University Park, 18°48’N 98°53’E,

elevation 300 m, July 27 2010, Karunarathna NTS-121 (holotype MFLU-10-0851;

isotype XAL.

Erymo.oey: The epithet refers to Thailand and to the similar species P aerugineo-

maculans.

70 ... Guzman & al.

PitEus 10-20 mm diam., conic to subumbonate, finally convex, smooth,

glabrous, margin glabrous, striate when dry, hygrophanous, dark reddish

brown to pale yellowish brown to whitish, bluing, finally blackish chocolate

when dry. LAMELLAE broadly adnate to subadnexed, brownish violaceous

to chocolate-brown, with white edges. StrpE 25-45 x (2-)3-4 mm, equal,

subbulbous, smooth above, sub-floccose below, base fibrillose, pale yellowish

brown to white or brownish black to reddish brown, hollow to stuffed, with

whitish conspicuous mycelium at the base. ANNULUS membranous, white to

off white, thick, persistent. CONTEXT white to pale rose, bluing upon exposure.

ODOR & TASTE not checked, in herbarium specimens none, but taste strongly

pungent after some minutes. SPORE PRINT unknown.

BASIDIOSPORES 9-12 x 7-8(-9) x 5.5-7 um, subrhomboid or subhexagonal

in face-view, subellipsoid in side-view, thick-walled, wall 1-1.5 wu thick,

yellowish brown, with a slightly truncate germ pore at the apex and a short,

acute apiculus at the base. Basrp1a 21-31(-36) x (6-)7-10 um, 4-spored,

hyaline, cylindric-vesiculose to subclaviform, the sterigmata up to 6 um long.

PLEUROCYSTIDIA (11-)14-22(-24) x 5-11(-13) um, hyaline, thin-walled,

more or less common, ventricose-mucronate to sub-rostrate or fusiform-

globose or obovate. PSEUDOCYSTIDIA (25-)26-33(-35) x 11-16 um, pyriform,

content oily, yellowish brown, growing from the hymenophoral trama.

CHEILOCYSTIDIA 15-20 x (5—)6-8(-10) um, hyaline, ventricose-mucronate to

sub-rostrate or sublageniform, some fusiform or sub-ovate, rarely irregularly

branched. SUBHYMENIUM subcellular, hyaline to yellowish pale, the elements

2-9 um wide, thick-walled. HYMENOPHORAL TRAMA regular, with hyphae

2-23 um wide, hyaline, rarely pale yellowish, thin- to thick-walled. PILEIPELLIS

AN IXOCUTIS, 21-30 um thick, with hyaline hyphae, 2-6 um wide. CONTEXT

with hyaline or some yellowish hyphae, 2-12 um wide. CLAMP CONNECTIONS

present.

HABIT, HABITAT AND DISTRIBUTION — Solitary or gregarious at 300 m on

soil with rotting cow dung, in an open grassland below the shadow of trees.

Known only from the type locality.

DiscussIon. Possession of a stipe annulus on the stipe and thick-walled

subrhomboid spores place P. thaiaerugineomaculans in sect. Stuntzae Guzman.

The pseudocystidia easily separate this species from all others in the section,

mainly with the similar P aerugineomaculans (Hohn.) Singer & A.H. Sm. from

Java. That species, which has an ixocutis < 10 um thick and basidiospores

8.5-10.5 x 7-8 x 5.5-6.5 um, lacks pseudocystidia and its cheilocystidia are

18-22 x 6-8 um (Singer & Smith 1958). Another similar species, also from

Java, is P subaeruginascens Hohn., grows on dung, has 25-33 x 9-12 um

pleurocystidia and 20-33 x 6-9 um cheilocystidia, lacks pseudocystidia, and

its pileipellis is not gelatinous (Singer & Smith 1958). Guzman (1983) referred

Psilocybe s.s. in Thailand ... 71

FIGURES 10-19. Psilocybe samuiensis (type) - 10. pleurocystidia; 11. cheilocystidia. P. thailandensis

(type) - 12. basidiomata; 13. basidiospores; 14. pleurocystidia (chrysocystidia); 15. cheilocystidia.

P. thaizapoteca (type) - 16. basidiomata; 17. basidiospores; 18. basidia; 19. pleurocystidia. Scale

bars: 12, 16 = 10 mm; 10-11 = 8 um; 13-15, 17-19 = 5.5 um.

72. ...Guzman & al.

both Hohnel species to P subaeruginascens but later (Guzman 1995) separated

them following Singer & Smith (1958). Recently Guzman & Yang (2010)

described P. taiwanensis Zhu L. Yang & Guzman from Taiwan, also in the sect.

Stuntzae, with conic to subumbonate pilei, (5.5-)6-7 x (3.5-)4-4.5 x 3.5-4 um

basidiospores, pileipellis with 8-10 x 4—5.5 um cystidioid elements, and lacking

pseudocystidia. We conclude that P thaiaerugineomaculans is a well-defined

Asiatic species independent from P aerugineomaculans, P. subaeruginascens,

and P. taiwanensis. A final note: Guzman would refer the unidentified annulate

Psilocybe from northern Thailand depicted in Stamets (1996: 171, 4" picture at

bottom of color plate) to P thaiaerugineomaculans based on form and color of

the basidiomata, which agree well with the species here described.

Psilocybe thaicordispora Guzman, Ram.-Guill. & Karunarathna, sp. nov.

MycoBank MB 561576 FIGURES 21-29

Differs from Psilocybe thaicordispora in narrower longer branched cheilocystidia and

presence of pleurocystidia.

Type: Thailand, Chiang Mai Prov., approximately 100 km from Chiang Mai, near Huai

Nam Dang National Park, (19°16’N 98°16’E), elevation 1500 m, rainforest dominated by

Pinus kesiya, July 12, 2010; Guzman 38349 (holotype MFLU-10-0804, isotype XAL).

Erymo.oey: The epithet refers to Thailand and to the sect. Cordisporae.

PrLEus 10-30 mm diam., conic to subcampanulate, finally convex or subconcave-

umbonate, sometimes irregularly lobulate, sub-hygrophanous, yellowish brown

to brownish orange, smooth, glabrous, without veil remnants, cuticle not

separable. LAMELLAE subadnexed, yellowish pale to dark brown-violaceous,

with whitish edges. Stipe 35-50 x 1-4 mm, uniform or subbulbous, smooth

above to subfloccose toward the base or the latter is smooth, whitish above

to reddish brown or reddish black. VEIL poor development. CONTEXT white

and thick below pileus, yellowish and fibrous in stipe. The entire basidiome

intensely bluing and becoming blackish. Dried specimens are black-chocolate.

ODOR AND TASTE Slightly farinaceous. SPORE PRINT unknown.

BASIDIOSPORES (6.5—)7-8(-9) x (4.5-)5-5.5(-6.5) x 4.5-5 um, subrhomboid

or ovoid in face-view, subellipsoid in side-view, thick-walled, wall 1-1.5 um

thick, yellowish brown, with a truncate germ pore at the apex and a short acute

apiculus at the base. Basip1A 20-25 x 6-7 um, 4-spored, hyaline, ventricose-

subcylindric or subclaviform, with a middle constriction. PLEUROCYSTIDIA

12-18(-24) x 3.5-7 um, rare or more or less common, hyaline, thin-walled,

ventricose, mucronate to rostrate. CHEILOCYSTIDIA 12—20(-28) x (4.5-)5-7

(-8) um, hyaline, thin-walled, polymorphous, ventricose, mucronate or rostrate,

sometimes with a long cylindric base, apex frequently and irregularly bifurcate

or branched. SUBHYMENIUM subcellular, with hyaline elements to incrusted

by yellowish brown pigment, also with blue-green pigment. HYMENOPHORAL

Psilocybe s.s. in Thailand ... 73

FIGURES 20-32. Psilocybe thaizapoteca (type) - 20. cheilocystidia. P thaicordispora (21-25 =

type; 26-29 = Guzman 38346) - 21. basidiomata; 22. basidiospores; 23. basidia; 24. cheilocystidia;

25. pleurocystidia; 26. basidiomata; 27. basidiospores; 28. pleurocystidia; 29. cheilocystidia.

P. thaiaerugineomaculans (type) - 30. basidiomata; 31. basidiospores; 32. basidia. Scale bars: 21,

26, 30 = 10 mm; 20, 22-25, 27-29, 31-32 = 5.5 um.

74 ... Guzman & al.

TRAMA regular, with hyphae (2—)3-20(-25) um wide, thin- or thick-walled, up

to 0.8 um thick, branched, hyaline. PILEIPELLIS AN IXOCUTIS, 12-20 um thick,

with hyphae, 2-5 um wide, hyaline, branched. CONTEXT subcellular, hyaline to

pale brownish through the pileipellis, with hyphae 3-9(-23) um wide, hyaline,

some branched. CLAMP CONNECTIONS present.

HABIT, HABITAT AND DISTRIBUTION — Solitary or gregarious, on grassy soil,

below the shadow of trees, in an open subtropical forest, at 1500 m elevation.

Known only from the type locality.

ADDITIONAL MATERIAL STUDIED — THAILAND, CHIANG Mat PROv., approximately

100 km from Chiang Mai, near Huai Nam Dang National Park, (19°16’N 98°16’E),

elevation 1500 m, rainforest dominated by Pinus kesiya, July 12, 2010; Guzman 38346

(MFLU- 10-0801; XAL).

Discussion. Psilocybe thaicordispora belongs to sect. Cordisporae Guzman

because of its small spores, which are ovoid or subrhomboid in face-view,

thick-walled and have a broad germ pore, as well as in its tropical habitat. It is

somewhat similar to P papuana Guzman & E. Horak known only from Papua

New Guinea, and also has polymorphous cheilocystidia, but these are wider,

shorter and un-branched, (6.5-)10-18 x 6.5-13 um, and apparently has no

pleurocystidia (Guzman & Horak 1978). Another similar species is P dumontii

Singer ex Guzman known only from Panama, but this species has un-branched

cheilocystidia which are not so variable in form, 10-18(-—26) x 3-6(-8) um,

hyaline to somewhat brownish (Guzman 1978, Guzman & Piepenbring 2011).

Psilocybe thaiduplicatocystidiata Guzman, Karunarathna & Ram.-Guill., sp. nov.

MycoBAank MB 561577 FIGURES 36-42

Differs from Psilocybe caerulescens in the possession of pleurocystidia, polymorphic

cheilocystidia, and longer basidiospores.

Type: Thailand, Chiang Mai Prov., Chiang Mai University Park, 18°48’N 98°53’E,

elevation 300 m, July 27, 2010, SC Karunarathna NTS-120 (holotype MFLU-10-0850,

isotype XAL).

Erymo.ocy: The epithet refers to Thailand and the dimorphic pleurocystidia and

cheilocystidia.

PiLEus 20-40 mm diam., convex to subumbonate or subumbilicate, glabrous,

smooth but margin slightly striate, also sometimes sub-lobulate, sub-

hygrophanous, pale brown or brownish rose to brownish dull orange at the

center, pale yellowish brown when dry, bluing to becoming irregularly blackish

chocolate. LAMELLAE adnate to seceding, black-violaceous to blackish chocolate,

with white edges. Stipe 45-75 x 4-6 mm, equal, subbulbous, probably with a

pseudorhiza (not gathered), surface white to pale to dark reddish brown, or

pale grayish brown when dry, smooth above to sub-floccose below, hollow to

stuffed. CONTEXT white to pale rose, bluing. ODOR AND TASTE not checked

Psilocybe s.s. in Thailand ... 75

FIGURES 33-39. Psilocybe thaiaerugineomaculans (type) - 33. pleurocystidia type A;

34. pleurocystidia type B; 35. cheilocystidia. P. thaiduplicatocystidiata (type) - 36. basidiomata;

37. basidiospores; 38. basidia; 39. pleurocystidia type A. Scale bars: 36 = 10 mm; 33-35, 37-39 =

5.5 um.

76 ... Guzman & al.

FIGURES 40-43. Psilocybe thaiduplicatocystidiata (type) - 40. pleurocystidia type A; 41. pleuro-

cystidia type B; 42. cheilocystidia type A; 43. cheilocystidia type B. Scale bar = 5.5 um.

when fresh, without any odor or taste in herbarium specimens, but the taste

after some minutes is strongly pungent. SPORE PRINT unknown.

Psilocybe s.s. in Thailand ... 77

BASIDIOSPORES 9-10(-11) x 6.5-7(-8) x 5.5-6.5 um, subrhomboid or

obscurely subovoid in face-view, subellipsoid in side-view, thick-walled, wall

1-1.5 um thick, yellowish brown, with a truncate germ pore at the apex and

a short acute apiculus at the base. Basip1A 25-36 x 7-8(-9) um, 4-spored,

hyaline, ventricose-subcylindric or subclaviform, with a middle constriction,

sterigmata up to 12 um long. PLEUROCysTIDIA polymorphous, arising from the

subhymenium, of two types: (A) 20-37(-44) x 7-9.5 um, hyaline, ventricose-

subcylindric, short rostrate or with a long narrow and cylindric neck,

sometimes branched; (B) 22-29 x 10-15 um, as PsEUDOCYSTIDIA, hyaline or

yellowish pale, subglobose, subfusiform or ovoid, thin-walled. CHEILOCYSTIDIA

polymorphous, of two types: (A) 16-22(-27) x 6.5-9.5 um, fusiform, ventricose,

rostrate, regular or irregularly, apex short or long, sometimes lobulate; (B) 18-24

x 9.5-13 um, rare, as PPEUDOCYSTIDIA, grayish, arising from the hymenophoral

trama. SUBHYMENIUM thick, hyaline, subcellular, globose elements 5.5-13

(-20) um wide. HYMENOPHORAL TRAMA regular, with hyaline to yellowish

brown hyphae, 2-16 um wide. PILEIPELLIs subgelatinous, with hyaline hyphae,

2-4 um wide. CoNTEXT with hyaline to yellowish hyphae, 3.5-34 um wide,

thin-walled. CLAMP CONNECTIONS present.

HABIT, HABITAT AND DISTRIBUTION — Gregarious, on soil with rotting

dung, in an open grassland, at 300 m elevation. Known only from the type

locality.

Discussion. Psilocybe thaiduplicatocystidiata with its small, subrhomboid,

thick-walled spores is a special member of the sect. Cordisporae as it has two

types of both pleurocystidia and cheilocystidia, one as pseudocystidia, a feature

not previously observed on any species of sect. Cordisporae (Guzman 1983,

1995). This species may also possess a pseudorhiza, but this was lost during

picking. The basidiome is similar to P caerulescens Murrill, but that species lacks

pleurocystidia, the cheilocystidia are not polymorphous and its basidiospores

are (6-)6.5-8(-8.5) um long. P. thaiduplicatocystidiata is also similar to

P. wrightii Guzman, but the latter has simple pleurocystidia and cheilocystidia,

and also smaller spores. These two similar species are only known from the

neotropics.

Psilocybe thaizapoteca Guzman, Karunarathna & Ram.-Guill., sp. nov.

MycoBankK MB 561578 FIGURES 16-20

Differs from Psilocybe angustipleurocystidiata in a thick subgelatinous pilleipellis and

presence of branched, lobed pleurocystidia.

Type: Thailand, Chiang Mai Prov., close to Huai Nam Dang National Park (19°16'N

98°16'E), elevation 1500 m, rainforest dominated by Pinus kesiya, July 12, 2010, Guzman

38342 (holotype MFLU-10-0797, isotype XAL).

Erymo.ocy: The epithet refers to Thailand and similarity of the fungus to the P zapo-

tecorum-group.

78 ... Guzman & al.

PILEus (20-)30-50 mm diam., conic-umbonate to campanulate, finally convex

to plano-convex, sometimes irregularly lobulate, glabrous overall, smooth,

lubricous to soon dry, cuticle not separable, sub-hygrophanous, brownish red

with chocolate tints, at the margin without veil remnants. LAMELLAE adnexed

or short sinuate, brownish chocolate, edges whitish. Stipe 70-120 x 4-8 mm,

uniform but narrow toward the base, hollow, sub-fleshy to fibrous, whitish

above, brownish red or brownish purple to whitish gray or dark vinaceous

at the base. PSEUDORHIZA as a long irregular whitish cordon. VEIL poorly

developed as fibrillar remnants. CONTEXT white to whitish, fleshy, bluing to

blackish. ODOR AND TASTE slightly farinaceous at first, but the taste also a little

pungent in fresh specimens. SPORE PRINT unknown.

BASIDIOSPORES 6-6.5(-7) x (3-)3.5-4(-4.5) x 3-3.5 um, subellipsoid or

obscurely subrhomboid in face-view, subellipsoid on side-view, thin-walled,

wall 0.5-0.8 um thick, pale yellowish brown, with a truncate germ pore

at the apex and a short acute apiculus at the base. Bastp1a 19-25 x (4.5-)

5-5.5(-6) um, 4-spored, hyaline, ventricose, subcylindric, or claviform with

a middle constriction. PLEUROCyYSTIDIA 13-20(-30) x (3-)4-6(-7.5) um,

common, hyaline, subventricose or subcylindric regular or irregularly but

acute mucronate, frequently irregularly branched or lobed. Apparently there

are two kind of pleurocystidia, smaller and bigger, but these latter are very

rare. PSEUDOCYSTIDIA absent. CHEILOCYSTIDIA 15-22(-25) x 4-7(-8) um,

hyaline, subventricose, somewhat claviform, all of them mucronate, regular or

irregularly, also irregularly lobulate or subbranched. SUBHYMENIUM subcellular,

hyaline, with thin-walled, somewhat incrusted hyphae. HYMENOPHORAL

TRAMA regular, with 3-30 um wide, hyaline to yellowish or pale brownish

hyphae, some of them swollen. PILEIPELLIs as thin layer, not gelatinous, with

hyaline hyphae, 1-2(-5) um wide, without pileocystidia. CoNTEXT like the

hymenophoral trama hyphae. CLAMP CONNECTIONS present.

HABIT, HABITAT AND DISTRIBUTION — Gregarious, on muddy soil with

few herbs, mosses and grasses, in a tropical-subtropical vegetation, with

Dipterocarpaceae trees and some Pinus. Known only from the type locality.

DiscussIon. Its thin-walled, subellipsoid spores place P. thaizapoteca in

sect. Zapotecorum Guzman, reported the first time from Thailand. Psilocybe

angustipleurocystidiata Guzman (known only from Colombia and Mexico),

which also lacks pseudocystidia, differs in its thick (10-15(-20) um)

subgelatinous pileipellis and pleurocystidia that are neither branched nor

lobed. Psilocybe aucklandiae Guzman et al. (known only from New Zealand)

is differentiated by the subgelatinous pileipellis and unbranched cheilocystidia

and pleurocystidia. Psilocybe kumaenorum R. Heim (known only from Papua

New Guinea) is also similar to P. thaizapoteca but is distinguished by small

basidiomata (pileus 5-7 mm diam, stipe 15-30 x 1-2 mm) and non-lobed

cheilocystidia (Heim 1966).

Psilocybe s.s. in Thailand ... 79

Key to species of Psilocybe s.|. reported from Thailand

(bold = new species described in the present paper)

la. Non-bluing species (excluded from Psilocybe s.s.) ......... Deconica deconicoides

LB BUTS PeCles se fan. ccre tA wapd a swt le cota tea Bila Sette tbo ses goatee nenah des ethantes done 2

2a. Spores thin-walled, rhomboid or subellipsoid in face-view.

Cire VSO ys wise ho usa 4. Sonsay a ttewcal ts Pebad clas Bg chs Zotay dit peters zed P. thaizapoteca

2b. Spores thick-walled, subrhomboid or subhexagonal in face-view.

Not On mudd ySOilrrg ox stn sve f opdtew le aig Te Tall. te sella ole berkeley OH gale 8 3

Soe S TESOL igo 45 Gls 2s ele o> ole Acetone la ala we nln ng dla wtng aie Sy dle 3 4

SB eAMITU ISAS EINEM, FN a Fe uta Me be ed oN tS AOR aoe SA oes eRe oan ts

4a. Spores (12—)13-15(-17) um long, subhexagonal in face-view.

LOH Acs e1e5 1118 kel £)-1 9 01 ae MRR A penn aR EMAL Ata lk RO SP MES Se P. cubensis

4b. Spores smaller, up to 12 um long. Chrysocystidia present or absent............. 5

5a. Chrysocystidia absent. Spores 9-12 um long. Growing on litter

mn otienggt lematbat, cematdan Meotan da acer ber seas cat tides tuo. aeons P. thaiaerugineomaculans

SY CHEVsOCyStia PLOSEN™ iat Ak cet Ak cit neue tua enced mtu ln memiarete ain 6

6a. Spores 7-8 um long. On elephant dung ......................00. P. magnispora

6b. Spores (4.5-)5-6(-6.5) um long. On rotten wood ............... P. thailandensis

7a. Pleurocystidia and cheilocystidia of two types. Growing on soil

entails Houten ds ivouiag tri Mabagd rinonnegdrtezed-2 4d hngetes Sc sehen ator P. thaiduplicatocystidiata

7b. Pleurocystidia and cheilocystidia of one type. On grassy soil .................. 8

Say Spores OA Sapiin OMe or la ee clears hake Rad aad ae Wad be Bead P. samuiensis

8b. Spores (6.5-)7-8(-9) um long ...... 1. ee eee ee eee P. thaicordispora

Final considerations

It is interesting that four new species were found during such a short

excursion time (five days) in Thailand, while none of the five species

previously reported for the country were found. We suggest this demonstrates

the complexity of mushroom fructification and at same time the richness

of the mycota in the tropics. We confirmed that species of Psilocybe s.s.

sects. Cordisporae, Mexicanae, Stuntzae, and Zapotecorum are tropical or

subtropical. We also observed that bluing species have a greatly varied habitat,

for example, litter, muddy soil, rotten wood or dung. Although the odor and

taste of basidiomata are generally farinaceous for all bluing Psilocybe species,

for Thai species odor and taste are either not distinctive or acidulous to only

very slightly farinaceous, with the taste of dried material in all four new

species (as well of fresh basidiomata in P. thaizapoteca) becoming strongly

pungent after some minutes.

Acknowledgments

Partial funding support for the fieldwork to Guzman was provided by the Instituto de

Ecologia at Xalapa, Veracruz (Mexico). Guzman and Ramirez-Guillén also received the

80 ... Guzman & al.

help of Manuel Hernandez, Juan Lara Carmona and Alonso Cortés-Pérez in computing,

herbarium duties and microscopy observations, respectively. The Global Research

Network for Fungal Biology, King Saud University and the Mushroom Research

Foundation are thanked for various support. We would also thank Michael Pilkington,

Phongeun Sysouphanthong, and Ma Tao for their help in field explorations. Finally we

express our acknowledgements to Drs. L. Guzman-Davalos, C.L. Ovrebo, T. Baroni, and

J. Trappe for their critical review.

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http:/dx.doi.org/10.1007/s13225-010-0050-4

MY COTAXON

http://dx.doi.org/10.5248/119.83

Volume 119, pp. 83-88 January-March 2012

A new Corynesporella species and two first records from China

L1-Guo Ma, JIAN Ma, YI-DONG ZHANG & XIU-GUO ZHANG

Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China

*CORRESPONDENCE TO: zhxg@sdau.edu.cn, sdau613@163.com

ABSTRACT — A new species, Corynesporella obclavata, is described and illustrated from dead

branches of Pistacia chinensis collected from tropical forests in Yunnan Province, China. A

synopsis of Corynesporella species is provided. Stephembruneria elegans and Diplococcium

pulneyense are recorded for the first time from China.

KEY worps — anamorphic fungi, taxonomy

Introduction

Numerous fungi on wood have been discovered in tropical and subtropical

forests during recent research on the fungus flora of South China (Dai et al.

2009, Dai & Li 2010, Ma et al. 2010, 2011). In our continuing study of conidial

fungi on dead wood, a hitherto undescribed species of Corynesporella and

two new records were found. These three species are described, illustrated,

and compared with similar species. The specimens are deposited in HSAUP

(Herbarium of the Department of Plant Pathology, Shandong Agricultural

University) and HMAS (Mycological Herbarium, Institute of Microbiology,

Chinese Academy of Sciences).

Taxonomy

Corynesporella obclavata L.G. Ma & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 563273

Conidiophora macronematica, ramosa, 230-635 um longa, ad apicem 3.5-6.5 ym lata, ad

basem 8.5-15 um lata. Cellulae conidiogenae monotreticae, in conidiophoris incorporatae,

terminales, 7.5-16 x 3.5-7.5 um. Conidia solitaria, acrogena, obclavata, brunneae, cellula

apicali pallide brunnea, laevibus, 33-55 x 6.5-11 um, 5-7-pseudoseptata.

Type: China, Yunnan Province: the Forbidden Forest of Banna, on dead branches of

Pistacia chinensis Bunge (Anacardiaceae), 17 Oct. 2008, L.G. Ma (holotype, HSAUP

H0033; isotype, HMAS 146152).

Erymo toy: obclavata, in reference to the conidial shape.

84 ... Ma & al.

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Corynesporella obclavata sp. nov. (China) ... 85

Fic. 1. Corynesporella obclavata. a, b. Branched conidiophores, conidiogenous cells and

conidia. c. Conidiophores arising from wood. d. Conidiogenous cells. e. Conidia.

COLONIES on natural substrate effuse, brown to dark brown. Mycelium partly

immersed, partly superficial, composed of branched, septate, pale brown, smooth

hyphae, 2-4 um wide. CONIDIOPHORES macronematous, mononematous, erect,

branched, straight or flexuous, cylindrical, brown, thick-walled, multiseptate,

smooth, 230-635 um long, 3.5-6.5 um wide at the apex, 8.5-15 um wide at

the base. CONIDIOGENOUS CELLS monotretic, integrated, terminal, cylindrical,

smooth, brown, thick-walled, 7.5-16 x 3.5-7.5 um. Conrp1a solitary,

acrogenous, obclavate, brown, apical cells pale brown, smooth, thick-walled,

5-7-distoseptate, 33-55 x 6.5-11 um.

All seven Corynesporella species are presented in TABLE 1. In conidial shape,

C. obclavata is most similar to C. helminthosporioides (Holubova-Jechova 1987),

which can be easily separated by its larger (50-75 x 8.8-12.8 um) subhyaline

conidia with more septa (9-13-distosepta).

Stephembruneria elegans R.F. Castafieda, Fungi Cubenses III: 14, 1988. Fic. 2

COLONIES on natural substrate effuse, brown. Mycelium partly superficial,

partly immersed, composed of branched, septate, smooth, pale brown to brown

hyphae, 1-2 um wide. CONIDIOPHORES macronematous mononematous,

unbranched, erect, straight to slightly flexuous, smooth, dark brown, cylindrical,

thick-walled, paler and narrower at the apex, multiseptate, 170-290 um long,

5-11.5 um wide. CONIDIOGENOUS CELLS monophialidic, cylindrical, integrated,

86 ... Ma & al.

terminal, percurrent, smooth, pale brown to subhyaline, with a minute collar at

the apex, 2.5-3.3 um wide. ConrpiA solitary, phialidic, navicular to fusiform,

rounded at the apex, with a narrow hilum at the base, smooth, 6-7-euseptate,

simple, end cells subhyaline, central cells dark brown, 26.5-33.5 um long,

10-11.5 um wide.

SPECIMEN EXAMINED: CHINA, HAINAN PROVINCE: the National Natural Reserve of

Bawangling, on dead branches of Ficus benjamina L. (Moraceae), 23 Nov. 2008, L.G. Ma

(HSAUP H0514, HMAS 146149).

a b

20um

20um 20um

Fic. 2. Stephembruneria elegans. a. Conidiophores, conidiogenous cells and conidia.

b. Conidiogenous cells with immature conidia. c. Conidia.

Corynesporella obclavata sp. nov. (China) ... 87

The genus Stephembruneria was erected by Castafieda Ruiz (1988) to

accommodate a single species, S. elegans. Currently Stephembruneria

remains monotypic and is characterized by macronematous, mononematous

conidiophores, monophialidic, terminal, integrated, conidiogenous cells that

narrow to a slightly cupulate collarette at the apex, and solitary, euseptate,

navicular to fusiform, versicolored conidia. The type material differs from our

material by its larger conidia (30-43 x 10-13 um) and longer conidiophores

(250-350 um). This is the first record from China.

20um 20um

Fic. 3. Diplococcium pulneyense.

a. Branched conidiophores, conidiogenous cells and conidia. b. Conidia.

Diplococcium pulneyense Subram. & Sekar, Kavaka 15: 91, 1989. Fic. 3

COLONIES on natural substrate effuse, dark brown, hairy. Mycelium partly

superficial, partly immersed, composed of branched, septate, pale brown

to brown, smooth hyphae, 2-3 um wide. CONIDIOPHORES macronematous,

mononematous, branched, branches often short, erect, cylindrical, straight

88 ... Ma & al.

or flexuous, smooth, brown, thick-walled, 6-12-septate, 110-265 um long,

8.5-10 um wide at the base, 5-7.5 um wide at the apex. CONIDIOGENOUS

CELLS integrated, polytretic, terminal and intercalary, cylindrical, brown,

smooth, thick-walled. Conip1a in acropleurogenous chains, up to 7 in number,

ellipsoidal, subcylindrical, 1-(2)-euseptate, rounded at both ends, concolored,

brown, smooth, thick-walled, 10-18.5 x 6-7.5 um.

SPECIMEN EXAMINED: CHINA, GUANGDONG PROVINCE: the National Forest Park of

Shimen, on dead branches of Bauhinia blakeana Dunn (Caesalpiniaceae), 20 Oct. 2010,

L.G. Ma (HSAUP H1787, HMAS 146150).

Diplococcium pulneyense is most similar to D. spicatum in conidial shape and

septation (Grove 1885), but D. spicatum can be distinguished by its smaller

conidia (6-9 x 3-4 um). The type material differs from the Chinese specimen

by its longer conidia (12-30 um) and narrower conidiophores (3-5.5 um). This

is a new record from China.

Acknowledgments

The authors are grateful to Dr Eric H.C. McKenzie and Dr W.B. Kendrick for serving

as pre-submission reviewers and for their valuable comments and suggestions. This

project was supported by the National Natural Science Foundation of China (Nos.

31093440, 30499340, 30770015) and the Ministry of Science and Technology of the

People’s Republic of China (Nos. 2006FY120100, 2006FY110500-5).

Literature cited

Castaneda Ruiz RF. 1988. Fungi Cubenses II: 1-27.

Dai YC, Li HJ. 2010. Notes on Hydnochaete (Hymenochaetales) with a seta-less new species

discovered in China. Mycotaxon 111: 481-487. http://dx.doi-org/10.5248/111.481

Dai YC, Cui BK, Yuan HS. 2009. Trichaptum (Basidiomycota, Polyporaceae) from China with a

description of three new species. Mycol. Prog. 8: 281-287.

http://dx.doi.org/10.1007/s11557-009-0598-0

Grove WB. 1885. New or noteworthy fungi. Part II. J. Bot. 23: 161-169.

Holubova-Jechova V. 1987. Studies on hyphomycetes from Cuba VI. New and rare species with

tretic and phialidic conidiogenous cells. Ceska Mykologie 41: 107-114.

Ma J, Wang Y, O’Neill NR, Zhang XG. 2011. A revision of the genus Lomaantha, with the description

of a new species. Mycologia 103: 407-410. http://dx.doi.org/10.3852/10-176

Ma IG, Ma J, Zhang YD, Zhang XG. 2010. A new species of Spadicoides from Yunnan, China.

Mycotaxon 113: 255-258. http://dx.doi.org/10.5248/113.255

MY COTAXON

http://dx.doi.org/10.5248/119.89

Volume 119, pp. 89-94 January-March 2012

Queletia mirabilis, a rare gasteroid species in Gran Canaria, Spain

G. Moreno’ , M.A. R1IBEs?, V. EscosBio? & A.N. KHALID4

' Department of Biologia Vegetal, University of Alcald,28871 Alcala de Henares, Madrid, Spain

? Avenida Pablo Neruda 120 F, 2° D, 28018 Madrid

° Sociedad Micologica de Gran Canaria, Apartado 609, 35080 Las Palmas de Gran Canaria

* Department of Botany, University of the Punjab, Lahore, Pakistan

* CORRESPONDENCE TO: gabriel.moreno@uah.es

ABSTRACT — This paper provides the first report of a rare gasteroid taxon, Queletia mirabilis,

from the Canary Islands, accompanied by color photographs of its macro- and microscopic

characteristics and scanning electron micrographs of spore ornamentation and capillitium.

A comparison with other similar species of Queletia and Schizostoma is also given.

KEY worps — Agaricaceae, Basidiomycota, taxonomy

Introduction

Queletia mirabilis is a very rare species. According to Lloyd (1904) and

Azema (1990), it was first collected by the mycologist, Perdrizet of Vaudancourt

(Oise), in Port de Sochaux (France) around 1868 and sent to Lucien Skeleton,

a doctor in Hérimoncourt (Doubs), who eventually passed it to Fries, who

classified and named it. This species has very few citations and is considered

by some mycologists as “an alien species.” Queletia mirabilis is known only

from France (Dumée & Maire 1913, Mornand 1989, Azema 1990), England

(Ramsbottom 1953, Pegler et al. 1995), Italy (Cetto 1980, Sarasini 2005), and

USA (White 1901, Lloyd 1904, Coker & Couch 1928). The species has also been

described from different authors’ herbarium material such as Fischer (1900),

Lloyd (1903), Pilat (1955), Moravec (1958) and Jiilich (1983). For the first time

below, we describe Q. mirabilis from the island of Gran Canaria, Canary Islands

(Spain), in the Macaronesian region and provide SEM illustrations of its spore

and capillitial ultrastructure.

Materials & methods

The studied specimens have been deposited in the herbaria of AH (University of

Alcala, Spain) and the Botanical Garden “Viera y Clavijo” (Gran Canaria, Spain) and

90 ... Moreno & al.

the personal herbarium of M.A. Ribes (MAR). Glebal tissue was mounted in Hoyer’s

medium and studied with a Nikon microscope. Spore measurements were made using

an oil immersion objective. The scanning electron microscope (SEM) photographs were

made according to Moreno et al. (1995).

Taxonomy

Queletia mirabilis Fr., Ofvers. Forhandl. K. Svenska Vetensk.-Akad. 28: 171 (1872)

Figs. 1-12

BASIDIOMATA scattered to gregarious, stipitate. PiLeus globose to

subglobose, 3-4 cm in diam., 1.5-2 cm high. PErrp1um mostly missing, only

near the base in contact with foot, composed of a membrane, 3-5 mm thick,

with smooth, whitish outer surface; rough inner surface, irregular, brown

to ochre, with abundant remains of capillitium and spores. GLEBA ocher,

abundant, with filaments of whitish capillitium. DEHISCENCE apical, irregular,

without formation of a stoma or defined structure. STIPE 0.7-1.2 x 6-10 cm,

cylindrical, curved, fading towards base, not twisted, with a strongly folded

surface and appearing torn, ochre and brown, woody. COLUMELLA absent.

Basip1ospores yellow-ochraceous, globose to subglobose, (4.9-)5.4-6.6

(-7.8) x (4.4-)4.9-6.2(-6.8) um, excluding ornamentation; Q = 1.0-1.2(-1.4),

N = 94, I = 6.0 x 5.5 um, Q = 1.1, with a prominent hilar appendix, strongly

verrucose, with verrucae grouped into conical structures (0.8-)1.1-1.5(-1.7)

um, N = 52, I = 1.3 um high. Basrpra not seen. CapPILLITIUM abundant,

cylindrical, twisted, wavy-helical without fibulae, ocher, thick-walled, poorly

branched and rarely septate, that breaks easily, probably as a result of handling

(4.4—)5.6-8.9(-10.3) um, I = 7.1 um wide.

SEM revealed spore ornamentation consisting of long rods that join together

to form pyramidal structures. This type of spore ornamentation matches

Q. mirabilis from France deposited at PC (Jiilich 1983).

MATERIAL EXAMINED: SPAIN: CANARY ISLANDS, GRAN CANARIA, Tejeda, close to the

“Presa de la Data” dam, (27°55’52.35"N, 15°39°17.32” W) in sandy soil in grass near dam,

flooded all year round with seasonal variations, surrounding vegetation scattered Pinus

canariensis C. Sm. ex DC., a thicket of legumes [Teline microphylla (DC.) PE Gibbs &

Dingwall and Chamaecytisus proliferus ssp. meridionalis Acebes] and “tabaibas” bitter

(Euphorbia regis-juba Webb & Berthel.), with “verodes” (Kleinia neriifolia Haw.), 16-X-

2010, leg. M. Cardona Sosa, det. Gabriel Moreno, AH 39331, MAR 161010-25, duplicate

in Botanical Garden “Viera y Clavijo” SMGC (Soc. Mycol. Gran Canaria) 11043.

ITALY: FRIULI-VENEZIA GIULIA, UDINE, Gorizia, Rindenabfallen von auf bei Laubholz

Torviscosa, with substrate temperature 38-40° C, 25-VIII-1980, leg. Tarcisio Figar misit

B. Cetto, det. Tarcisio Figar, Hanns Kreisel ex Herbarium, duplicate in AH 25235.

OTHER MATERIAL STUDIED: Queletia andina —ARGENTINA, MENDOzaA, San

Martin, leg. J. Semper, 15-VII-1950, Ruiz Leal 16204 (holotype, BAFC 28711;, isotype,

AH 39332). Queletia mundkurii —PAKISTAN, Punyjas, Ladhar, on ground, 18-VIII-

1947, LAH, duplicate in AH 39333.

Queletia mirabilis in Gran Canaria... 91

Fic. 1-3: Queletia mirabilis (AH 39331).

1. Fruiting bodies. 2-3. Details of gleba (2) and capillitial remnants (3). Scale bars = 0.5 cm.

Comments: Although Queletia mirabilis isa rare species, it is easily distinguished

by its irregular peridium dehiscence, absence of columella, filamentous,

helical-wavy, winding capillitium, and strongly ornamented basidiospores.

Development of the fruiting bodies, details of peridium dehiscence, basidia

92 ... Moreno & al.

Fics 4-12: Queletia mirabilis (AH 39331).

4-6. Details of capillitium (LM, 4-5) and spores (LM, 6) in distilled water).

7-12. SEM photos: 7-9. Capillitium and branches; 10-12. Spore ornamentation.

Scale bars: 4-6 and 8-9 = 10 um, 7 = 20 um, 10-12 = 1 um.

formation, and development of the spores have been described in detail by

Dumée & Maire (1913).

De Toni’s (1888: 65) description of Q. mirabilis differs from those of other

mycologists because he described comparatively larger basidiospores (10-15

um diam.). In general, the basidiospores are smaller (7.5-9.2 um including

ornamentation) than given by De Toni (1888). The size of the fruiting bodies

also differs. According to De Toni (1888), the stipe is 1-2 cm long with a base

up to 4.5 cm wide and apex 2.5 cm broad.

This gasteroid fungus is usually found fruiting on bark and wood debris

(Cetto 1980, Azéma 1990, Pegler et al. 1995, Sarasini 2005). It rarely produces

fruiting bodies on the ground after rains (De Toni 1888), an important

character. The sample from the Canary Islands grew in sandy soil that remains

waterlogged all year round but with seasonal variations.

Queletia mirabilis in Gran Canaria ... 93

The genera Queletia Fr. and Schizostoma Ehrenb. ex Lév., are closely related

by the wavy-helical, aseptate or rarely septate, shortly branched capillitium.

However, Schizostoma differs in possessing smooth basidiospores and a

peridium that becomes irregular and stellate at maturity.

The only other species currently accepted in this genus is Queletia andina

J.E. Wright, which is characterized by its filamentous, branched capillitium

with abundant septa (Tulostoma type) that do not break under the pressure

and smooth basidiospores. The capillitium of Schizostoma mundkurii

(S. Ahmad) Long & Stouffer [ = Queletia mundkuriiS. Ahmad] is similar to that of

Q. mirabilis but presence of smooth spores and other generic characters of the

former differentiate these two species.

Acknowledgements

We express our gratitude to Dr. M. Esqueda and Lda. M?.M. Dios for reviewing the

manuscript and adding a number of useful comments. We also thank Mr. Luis Monje

of the “Cabinet of Drawing and Scientific Photography’, at the University of Alcala de

Henares for help in the digital preparation of the photographs and to Dr. J. Rejos, curator

of the AH herbarium. We express our gratitude to D.W. Mitchell and Dr. Amy Rossman,

Research Leader, Systematic Mycology & Microbiology Laboratory, USDA-ARS, for

their assistance in correcting the English and providing valuable comments.

Literature cited

Azéma RC. 1990. Queletia mirabilis. Bull. Fed. Mycol. Dauphine-Savoie 119: 20-22.

Cetto B. 1980. Guia de los hongos de Europa 3. Ediciones Omega S.A. Barcelona. 645 p.

Coker WC, Couch JN. 1928. The Gasteromycetes of the eastern United States and Canada. University

of North Carolina Press, Chapel Hill. 201 p.

De Toni GB. 1888. Gasteromyceteae. Fam. 3. Lycoperdaceae. Sylloge Fungorum 7(1): 48-154.

Dumeée P, Maire R. 1913. Note sur le Queletia mirabilis Fr. et sa découverte aux environs de Paris.

Bulletin Société Mycologique de France. 29: 495-502.

Fischer E. 1900. Plectobasidiineae. In: A Engler, K Prantl (eds). Nat. Pflanzenfam. 1, Abt. 1, Lief.

195%

Julich W. 1983. Sporenstruktur bei Gasteromycetes. 2. Queletia. International Journal of Mycology

and Lichenology. 1: 169-174.

Lloyd CG. 1903. 237-Queletia. 238-Queletia mirabilis. [Mycological Notes n°. 14]. Mycological

Writings 1: 135-136.

Lloyd CG. 1904. 278-Queletia mirabilis. [Mycological Notes n°. 17]. Mycological Writings 1: 185.

Mornand J. 1989. Les Gasteromycetes de France (5 Tulostomatales). Documents Mycologiques

19(76): 1-18.

Moravec Z. 1958. Queletia Fr. 615-617, in: A. Pilat (ed.) Gasteromycetes. Flora C.S.R. Prague.

Moreno G, Altés A, Ochoa C, Wright JE. 1995. Contribution to the study of the Tulostomataceae in

Baja California, México. |. Mycologia 87: 96-120. http://dx.doi.org/10.23 07/3760953

Pegler DN, Lzessge T, Spooner BM. 1995. British puffballs, earthstars and stinkhorns. An account

of the British Gasteroid Fungi. Royal Botanic Gardens, Kew. 255 p.

Pilat A. 1955. Vénujte v roce 1955 zvySenou pozornost houbam brichatkovitym - Gasteromycetes.

Ceska Mykologie 9(2): 91-94.

94 ... Moreno &al.

Ramsbottom J. 1953. Mushrooms & toadstools. A study of the activities of Fungi. London. Collins

New Naturalist Series. doi:10.1079/PNS19530011

Sarasini M. 2005. Gasteromiceti epigei. Assoc. Micol. Bresadola. Fondazione Centro Sttudi

Micologici del A.M.B. Vicenza. 406 p.

White VS. 1901. The Tylostomaceae of North America. Bulletin Torrey Botanical Club 28: 421-444.

http://dx.doi.org/10.2307/2478590

MY COTAXON

http://dx.doi.org/10.5248/119.95

Volume 119, pp. 95-102 January-March 2012

New species and a new record of Solicorynespora

from southern China

JIAN Ma, Li-Guo Ma, Y1-DONG ZHANG, JI-WEN XIA & XIU-GUO ZHANG

Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China

*CORRESPONDENCE TO: zhxg@sdau.edu.cn, sdau613@163.com

ABSTRACT — Four species of Solicorynespora were collected on dead branches from tropical

or subtropical forests in southern China. The new species S. ligustri on Ligustrum quihoui,

S. machili on Machilus chinensis, and S. cryptocaryae on Cryptocarya chinensis are described,

illustrated, and compared with closely related taxa. Solicorynespora pseudolmediae is recorded

for the first time from China.

KEY worpDs — anamorphic fungi, taxonomy

Introduction

Solicorynespora R.F. Castafieda & W.B. Kendr., erected by Castaneda &

Kendrick (1990) with S. zapatensis R.F. Castafieda & W.B. Kendr. as the type

species, was mainly characterized by conspicuous, single, determinate or

percurrently extending conidiophores and integrated, terminal, monotretic

conidiogenous cells with solitary, obclavate or pyriform, euseptate

phragmoconidia. These characters separate Solicorynespora from similar

genera including Corynespora Giissow (Gtissow 1906), Corynesporella Munjal

& H.S. Gill (Munjal & Gill 1961), Hemicorynespora M.B. Ellis (Ellis 1972), and

Corynesporopsis P.M. Kirk (Kirk 1981). Keys to Solicorynespora species have

been given by Castafieda et al. (2004) and Ma et al. (2011b), based mainly

on conidial morphology. Until now, 15 species have been included within

Solicorynespora, with species circumscribed primarily based on conidial shape,

size, septation, ornamentation, lateral appendage and presence or absence of

a rostrum (Castaneda 1996, Castafieda et al. 2004, McKenzie 2010, Ma et al.

2011b).

The forests of southern China have a rich mycota, and many lignicolous

fungi have been discovered there (Dai & Cui 2006, Yuan & Dai 2009, Zhang et

al. 2009, Cui et al. 2011, Ma et al. 2011a). During ongoing surveys of conidial

96 ... Ma & al.

fungi associated with woody debris in tropical or subtropical forests of southern

China, several hyphomycete species were collected on dead branches. The

present paper deals with four species clearly related to the genus Solicorynespora.

Three are new to science while the fourth is a new record for China.

A B C D

bee |/ I,

6866

6666 .

20m

wiigz

Fic. 1. Solicorynespora ligustri. A. Colonies on natural substratum. B. Conidiophores

with developing conidia. C. Conidiophore with terminal conidium. D. Conidiophore

apices showing conidiogenous cells and percurrent extensions. E. Conidia.

Solicorynespora ligustri Jian Ma & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 563282

Differs from Solicorynespora zapatensis and S. calophylla in smooth conidia that lack a

constriction at the septa.

Type: China, Hainan Province: Natural Reserve of Liulianling, on dead branches of

Ligustrum quihoui Carriere (Oleaceae), 5 Dec 2009, J. Ma, (Holotype HSAUP H5247;

isotype HMAS 146078).

EryMoLoey: in reference to the host genus, Ligustrum.

Conidial fungi. Colonies on natural substrate effuse, brown to dark brown, hairy.

Mycelium partly superficial, partly immersed in the substratum, composed of

branched, septate, pale brown to brown, smooth-walled hyphae, 1-2.5 um

thick. Conidiophores distinct, single or in groups, erect, straight or flexuous,

Solicorynespora spp. nov. (China) ... 97

unbranched, smooth, septate, brown to dark brown, 52-97 um long, 2.5-4.5

um wide. Conidiogenous cells monotretic, integrated, terminal, cylindrical,

brown, smooth, 11-17 um long, 2-3.5 um wide, with 0-2 cylindrical percurrent

extensions. Conidia solitary, dry, acrogenous, obclavate or obpyriform, mostly

3-euseptate, rarely 2-euseptate, smooth, the lower two cells brown, other cells

pale brown to subhyaline, 13-22 um long, 4.5-6 um wide in the widest part,

tapering to 1.5-2.5 um near the apex, 1.5-2.5 um wide at the truncate base.

ComMENTs - Among the known Solicorynespora species, S. ligustri bears some

resemblances to S. zapatensis and S. calophylli (Hol.-Jech. & R.E Castafieda) R.E.

Castafieda & W.B. Kendr. (Castafieda & Kendrick 1990) in conidial shape, but

the conidia of S. ligustri are smaller than in S. zapatensis (20-27 x 15-17 um)

and longer and with more septa than in S. calophylli (11-16 um long, 2-septate).

Solicorynespora zapatensis and S. calophylli are additionally distinguished by

having roughened conidia with constrictions at the septa.

Solicorynespora machili Jian Ma & X.G. Zhang, sp. nov. FIG. 2

MycoBank MB 563283

Differs from Solicorynespora zapatensis in producing smaller conidia with pigmented

lower cells, from S. calophylli in having larger conidia, and from S. ligustri by its wider

conidia with verrucose wall.

Type: China, Guangdong Province: Mount Dinghu, collected on dead branches of

Machilus chinensis (Benth.) Hemsl. (Lauraceae). 18 Oct 2010, J. Ma, (Holotype HSAUP

H5427; isotype HMAS 146079).

EryMoOLoGey: in reference to the host genus, Machilus.

Conidial fungi. Colonies on natural substrate effuse, brown to dark brown, hairy.

Mycelium partly superficial, partly immersed in the substratum, composed of

branched, septate, pale brown to brown, smooth-walled hyphae, 1-2.5 um

thick. Conidiophores distinct, singly or in groups, erect, straight or flexuous,

unbranched, smooth, septate, brown to dark brown, 76-110 um long, 3.5-4.5

um wide. Conidiogenous cells monotretic, integrated, terminal, brown, smooth,

10-16 um long, 3.5-4.5 um wide, with 0-1 lageniform percurrent extension.

Conidia solitary, dry, acrogenous, obclavate or obpyriform, 3-euseptate, the

lower two cells verrucose, brown to dark brown, other cells smooth, pale brown

to subhyaline, 16-24 um long, 7-8.5 um wide in the widest part, tapering to

1.5-2.5 um near the apex, 2-2.5 um wide at the truncate base.

ComMENTSs - Solicorynespora machili resembles S. zapatensis, S. calophylli

and S. ligustri in conidial shape. However, S. zapatensis can be distinguished

by its distinctly larger conidia (20-27 x 15-17 um) with unpigmented lower

two cells, while S. calophylli has smaller (11-16 x 5-7 um) 2-euseptate conidia.

Solicorynespora ligustri is differentiated by its wider (4.5-6 um) conidia with

verrucose walls.

98 ... Ma & al.

A B C

wngz

20m

20nm

Fic. 2. Solicorynespora machili. A. Conidiophores with terminal conidia. B. Conidiophore

apices showing conidiogenous cells and percurrent extension. C. Conidia.

Solicorynespora cryptocaryae Jian Ma & X.G. Zhang, sp. nov. Fic. 3

MycoBank MB 563284

Differs from Solicorynespora mulanjeensis and S. pseudolmediae in slightly wider conidia

with more septa, and further differs from S. mulanjeensis and S. pseudolmediae in having

shorter or longer conidia, respectively.

Type: China, Guangdong Province: Liuxihe National Forest Park, on dead branches

of Cryptocarya chinensis Hemsl. (Lauraceae). 20 Oct 2010, J. Ma, (Holotype HSAUP

H5367-2; isotype HMAS 146087).

ETryMoLoey: in reference to the host genus, Cryptocarya.

Conidial fungi. Colonies on natural substrate effuse, brown, hairy. Mycelium

mostly immersed in the substratum, composed of branched, septate, pale brown

Solicorynespora spp. nov. (China) ... 99

to brown, smooth-walled hyphae, 1.5-3 um thick. Conidiophores distinct,

single or in groups, erect, straight or flexuous, unbranched, smooth, septate,

brown to dark brown, 86-140 um long, 4.5-6.5 um wide. Conidiogenous

cells monotretic, integrated, terminal, brown, smooth, 13-21 um long, 4-5.5

uum wide, with 0-1 cylindrical percurrent extension. Conidia solitary, dry,

acrogenous, obclavate or ellipsoidal, gradually tapered to an obtuse paler apex,

6-12-euseptate, smooth, brown, 32-55 um long, 10-13 um wide in the widest

part, 3.5-4 um wide at the truncate base.

ComMENTS - Solicorynespora cryptocaryae superficially resembles S.

mulanjeensis (B. Sutton) R.F. Castafieda et al. (Castafieda et al. 2004) and S.

pseudolmediae (Castafteda & Kendrick 1990). However, the conidia of S.

mulanjeensis are longer (56-71 um), and those of S. pseudolmediae are shorter

(16-29 um). Moreover, S. mulanjeensis (conidia 10-12.5 um wide, 5-8-

euseptate) and S. pseudolmediae (conidia 8.5-12 um wide, 2—-5-euseptate) are

further distinguished by their slightly narrower conidia with fewer septa.

__20nm

Fic. 3. Solicorynespora cryptocaryae. A. Conidiophores with terminal conidia, and conidiophore

apices showing conidiogenous cells and percurrent extension. B. Conidiophore. C. Conidia.

A B

100 ... Ma & al.

Solicorynespora pseudolmediae (R.F. Castafieda) R.E. Castafieda & W.B. Kendr.,

FIG. 4

Univ. Waterloo Biol. Ser. 33: 43, 1990.

= Sporidesmium pseudolmediae R.F. Castaneda, Rev. Jard. Bot. Nac. 5: 66. 1984.

= Corynespora pseudolmediae (R.F. Castafieda) Hol.-Jech., Ceska Mykol. 40: 145. 1986.

20um

Fig. 4. Solicorynespora pseudolmediae. A. Conidiophores with terminal conidia.

B. Conidiophore and conidium. C. Conidiophore with terminal conidium, and conidio-

phore apex showing conidiogenous cells and percurrent extension. D. Conidia.

Solicorynespora spp. nov. (China) ... 101

Conidial fungi, hyphomycetes. Colonies on natural substrate effuse, dark

brown, hairy. Mycelium partly superficial, partly immersed in the substratum,

composed of branched, septate, pale brown to brown, smooth-walled hyphae,

1.5-3.5 um thick. Conidiophores distinct, single or in groups, erect, straight or

slightly flexuous, unbranched, smooth, brown, 40-170 um long, 4-5.5 um wide

at the base, 4—5 um wide in the middle, 3.5—4.5 um wide at the apex, 2—5-septate,

sometimes with 1-2 enteroblastic percurrent extensions. Conidiogenous

cells monotretic, integrated, terminal, cylindrical, brown, smooth, 14-26 um

long, 3.5-5 um wide. Conidial secession schizolytic. Conidia solitary, dry,

acrogenous, obclavate or obpyriform, 2-5-euseptate (usually 4), sometimes

slightly constricted at the speta, smooth, brown, with 1-2 upper cells pale

brown or subhyaline, 22-32 um long, 9-13 um wide in the widest part, tapering

to 3-4.5 um wide near the apex, 2-3 um wide at the truncate base.

SPECIMEN EXAMINED: CHINA, YUNNAN PROVINCE: Tropical Botanical Garden of

Xishuangbanna, on dead branches of unidentified plant, 12 Oct 2007, J. Ma, (HSAUP

H5286, HMAS 146088).

ComMENTs -'This species was originally assigned to Sporidesmium Link and

subsequently transferred to Corynespora (Holubova-Jechova & Mercado

1986) due to its monotretic conidiogenous cells. Castafieda & Kendrick (1990)

established the genus Solicorynespora and moved this species into the new genus

based on its euseptate conidia. Solicorynespora pseudolmediae is morphologically

similar to S. zapatensis in conidial shape, but differs from the latter in conidial

size, wall ornamentation, and the number of septa. Comparing our collection

with the type material in Castafieda's description (Castaneda 1984), the conidia

of our specimen are slightly larger (22-32 x 9-13 um vs. 16-29 x 8.5-12 um)

and the conidiophores are shorter (40-170 um vs. 50-275 um). Despite these

minor differences, we believe they are basically the same species.

Acknowledgments

The authors express gratitude to Dr Bryce Kendrick and Dr Eric H.C. McKenzie for

serving as pre-submission reviewers and for their valuable comments and suggestions.

This project was supported by the National Natural Science Foundation of China (Nos.

31093440, 30499340, 30770015) and the Ministry of Science and Technology of the

People’s Republic of China (Nos. 2006FY120100, 2006FY110500-5).

Literature cited

Castafieda Ruiz RF. 1984. Nuevos taxones de Deuteromycotina: Arnoldiella robusta gen. et sp. nov.;

Roigiella lignicola gen. et sp. nov.; Sporidesmium pseudolmediae sp. nov.; y Thozetella havanensis

sp. nov. Rev. Jard. Bot. Nac. (La Habana, Cuba) 5: 57-87.

Castaneda Ruiz RF. 1996. New species of Haplotrichum and Solicorynespora from Cuba. Mycotaxon

59: 449-452.

Castaneda Ruiz RF, Kendrick B. 1990. Conidial fungi from Cuba: II. Univ. Waterloo Biol. Ser. 33:

1-61.

102 ... Ma &al.

Castafieda Ruiz RF, Heredia GP, Arias RM, Saikawa M, Minter DW, Stadler M, Guarro J, Decock

C. 2004. Two new hyphomycetes from rainforests of México, and Briansuttonia, a new genus to

accommodate Corynespora alternarioides. Mycotaxon 89: 297-305.

Cui BK, Du P, Dai YC. 2011. Three new species of Inonotus (Basidiomycota, Hymenochaetaceae)

from China. Mycol. Prog. 10: 107-114. http://dx.doi.org/10.1007/s11557-010-0681-6

Dai YC, Cui BK. 2006. Two new species of Wrightoporia (Basidiomycota, Aphyllophorales) from

southern China. Mycotaxon 96: 199-206.

Ellis MB. 1972. Dematiaceous hyphomycetes. XI. Mycol. Pap. 131: 1-25.

Giissow HT. 1906. Uber eine neue Krankheit an Gurken in England (Corynespora mazei, Giissow

gen. et sp. nov.). Z. Pflkrankh.16: 10-13.

Holubova-Jechova V, Mercado SA. 1986. Studies on hyphomycetes from Cuba IV. Dematiaceous

hyphomycetes from the Province Pinar del Rio. Ceska Mykol. 40: 142-164.

Kirk PM. 1981. New or interesting microfungi II. Dematiaceous hyphomycetes from Esher

Common, Surrey. Trans. Br. Mycol. Soc. 77: 279-297.

http://dx.doi.org/10.1016/S0007-1536(81)80031-9

Ma J, Wang Y, Ma LG, Zhang YD, Castafieda Ruiz RE, Zhang XG. 2011a. Three new species of

Neosporidesmium from Hainan, China. Mycol. Prog. 10: 157-162.

http://dx.doi.org/10.1007/s11557-010-0685-2

Ma J, Zhang YD, Ma LG, Ren SC, Castafieda Ruiz RF, Zhang XG. 2011b. Three new species of

Solicorynespora from Hainan, China. Mycol. Prog.

http://dx.doi.org/10.1007/s11557-011-0775-9

McKenzie EHC. 2010. Three new phragmosporous hyphomycetes on Ripogonum from an ‘ecological

island’ in New Zealand. Mycotaxon 111: 183-196. http://dx.doi.org/10.5248/111.183

Munjal RL, Gill HS. 1961. Corynesporella: a new genus of hyphomycetes. Indian Phytopath. 14:

6-9.

Yuan HS, Dai YC. 2009. Hydnaceous fungi of China 2: Mycorrhaphium sessileum sp. nov. Nova

Hedwigia 88: 205-209. http://dx.doi.org/10.1127/0029-5035/2009/0088-0205

Zhang K, Ma J, Wang Y, Zhang XG. 2009. Three new species of Piricaudiopsis from southern China.

Mycologia 101: 417-422. http://dx.doi.org/10.3852/08-147

MY COTAXON

http://dx.doi.org/10.5248/119.103

Volume 119, pp. 103-107 January-March 2012

Two new species of Endophragmiella from southern China

Jian Ma’, Y1-DoONG ZHANG ', LI-Guo Ma’,

RAFAEL F, CASTANEDA-RUIZ 2 & XIU-GUO ZHANG "

"Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China

? Instituto de Investigaciones Fundamentales en Agricultura Tropical “Alejandro de Humboldt”

(INIFAT), Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200

*CORRESPONDENCE TO: zhxg@sdau.edu.cn, sdau613@163.com

AsBstTRAct —Endophragmiella gardeniae sp. nov., on dead branches of Gardenia hainanensis,

and E. machili sp. nov., on dead branches of Machilus grijsii, are described and illustrated.

They differ from other described Endophragmiella species in their obclavate or fusiform

conidia with branches that developed a Selenosporella-like synanamorph. The specimens are

deposited in Herbarium of Shandong Agricultural University, Plant Pathology (HSAUP) and

Mycological Herbarium, Institute of Microbiology, Chinese Academy of Sciences (HMAS).

KEY worpDs — anamorphic fungi, taxonomy

Introduction

Sutton (1973) established Endophragmiella for two species, E. pallescens

B. Sutton (the type species) and E. canadensis (Ellis & Everh.) B. Sutton.

Subsequently, the genus was emended by Hughes (1979), who gave a very

detailed account of conidiogenesis and generic concepts. Endophragmiella is

characterized by monoblastic, integrated, terminal, determinate or percurrently

extending conidiogenous cells on distinct, branched or unbranched

conidiophores, and solitary, acrogenous, septate conidia with rhexolytic

conidial secession. To date, 79 species and one variety have been recognized

within the genus, all from temperate to tropical areas (Wu & Zhang 2005,

Chen et al. 2008, Brackel & Markovskaja 2009, Castafieda Ruiz et al. 2010,

Leao-Ferreira & Gusmao 2010). The species are differentiated primarily by

conidial shape, size, septation, pigmentation, ornamentation, and presence or

absence of a rostrum (Sutton 1973, Hughes 1979, Wu & Zhuang 2005).

During ongoing surveys of saprobic microfungi from forests of

southern China, two interesting anamorphic species were collected from

dead branches. The characters of their conidia and conidiogenesis suggest

104 ... Ma & al.

they are appropriately disposed in the genus Endophragmiella, although

morphologically distinct from all previously known species. Therefore, they

are described here as new to science.

Taxonomy

Endophragmiella gardeniae Jian Ma & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 519223

Differs from all other Endophragmiella species except E. machili, in producing conidia

with branches that develop a Selenosporella-type anamorph.

Type: China, Hainan Province: tropical forest of Bawangling, on dead branches of

Gardenia hainanensis Merr. (Rubiaceae), 11 Dec 2009, J. Ma (holotype HSAUP H5149;

isotype HMAS 146102).

ETyMoLoey: in reference to the host genus, Gardenia.

Colonies on natural substrate effuse, brown, hairy. Mycelium partly superficial

and partly immersed in the substratum, composed of branched, septate, pale

20nm

Fic. 1. Endophragmiella gardeniae. a. Colonies on natural substratum. B. Rhexolytic

secession of conidium leading to a breakage of the conidiophore apex. c-E. Conidiophores

with terminal conidia. In p-E conidiophores showing conidiogenous cells and percurrent

extension. FE. Conidia with branches that develop a Selenosporella-like synanamorph.

Endophragmiella gardeniae & E. machili spp. nov. (China) ... 105

brown, smooth-walled hyphae, 1-3 um thick. Conidiophores distinct, single or

in groups, unbranched, cylindrical, erect, straight or flexuous, smooth, septate,

brown to dark brown, 85-210 um long, 4-7.5 um wide, sometimes with 1

cylindrical percurrent extension. Conidiogenous cells monoblastic, integrated,

terminal, indeterminate, cylindrical, smooth, pale brown to brown, 9-27 um

long, 4-5.5 um wide. Conidial secession rhexolytic. Conidia acrogenous,

solitary, obclavate or fusiform, 5-7-euseptate, slightly constricted at the septa,

smooth, brown, apical cell pale brown to subhyaline, 41-66 um long, 8-10.5

um wide in the widest part, 2.5-4.5 um wide at the truncate base, with a small

collar of conidiogenous cell well, apical portion with branches that developed a

Selenosporella-like synanamorph.

Fig. 2. Endophragmiella machili. 4. Colonies on natural substratum. B-c. Conidiophores

and conidia. p. Conidia with branches that develop a Selenosporella-like synanamorph.

Endophragmiella machili Jian Ma & X.G. Zhang, sp. nov. FIG. 2

MycoBank MB 519224

Differs from Endophragmiella gardeniae in its longer conidia with more septa.

Type: China, Fujian Province: subtropical forest of Mount Wuyi, on dead branches of

Machilus grijsii Hance (Lauraceae), 15 Aug 2009, J. Ma (holotype HSAUP H5108-3;

isotype HMAS 146103).

EryMo_oey: in reference to the host genus, Machilus.

106 ... Ma & al.

Colonies on natural substrate effuse, brown, hairy. Mycelium partly superficial

and partly immersed in the substratum, composed of branched, septate, pale

brown, smooth-walled hyphae, 1.5-3 um thick. Conidiophores distinct, single

or in groups, unbranched, determinate, cylindrical, erect, straight or flexuous,

smooth, septate, brown, 4-57 um long, 3-4.5 um wide. Conidiogenous cells

monoblastic, integrated, terminal, cylindrical, smooth, pale brown to brown,

7-14 um long, 3-4 um wide. Conidial secession rhexolytic. Conidia acrogenous,

solitary, obclavate, straight or curved, 6-10-euseptate, sometimes slightly

constricted at the septa, smooth, brown, apical cell pale brown to subhyaline,

93-135 um long, 6-9 um wide in the widest part, 2.5-4 um wide at the truncate

base, with a small collar of conidiogenous cell wall, apical portion with branches

that developed a Selenosporella-like synanamorph.

Discussion

Endophragmiella gardeniae and E. machili are unlike the taxa hitherto

placed in Endophragmiella in that their conidia have branches that develop a

Selenosporella-like synanamorph. The initial thought was to introduce a new

genus to accommodate these two species. However, Drs. Heredia and Arias

showed us pictures of an Endophragmiella sp. (on decaying wood, Mexico,

Veracruz, Acajete Municipal, Mesa de la Yerba, 19°33’N 97°01'W, 5 Dec. 2010,

coll. G. Heredia & R.M. Arias) with many states of conidial development that

are clearly the same as these two species. Therefore, we place them as new

species in the genus Endophragmiella.

Endophragmiella gardeniae and E. machili bear some resemblance to E. socia

(M.B. Ellis) S. Hughes (Hughes 1979) and E. fagicola P.M. Kirk (Kirk 1981)

in conidial shape. However, the conidia of E. machili are longer and narrower

than those of either E. socia (36-50 x 10.8-15.3 um, 6-11-septate, usually 7) or

E. fagicola (70-90 x 11-17 um, usually 5-septate). Endophragmiella gardeniae

also differs in conidial size and septation from E. socia and E. fagicola. In

addition, E. gardeniae and E. machili produce conidia with branches that

developed a Selenosporella-like synanamorph, a feature not found in E. socia

and E. fagicola. Endophragmiella gardeniae differs from E. machili in its shorter

conidia with fewer septa.

Acknowledgments

We are deeply indebted to Dr Bryce Kendrick and Dr Eric H.C. McKenzie for serving

as pre-submission reviewers and for their valuable comments and suggestions. We

thank Dr Gabriela Heredia Abarca and Dr Rosa Maria Arias for their generous and

valued assistance with the fungi described above. This project was supported by the

National Natural Science Foundation of China (Nos. 31093440, 30499340, 30770015)

and the Ministry of Science and Technology of the People’s Republic of China (Nos.

2006FY 120100, 2006FY110500-5).

Endophragmiella gardeniae & E. machili spp. nov. (China) ... 107

Literature cited

Brackel W von, Markovskaja S. 2009. A new lichenicolous species of Endophragmiella from Bavaria/

Germany. Nova Hedwigia 88: 513-519.

http://dx.doi.org/10.1127/0029-5035/2009/0088-0513

Castafieda Ruiz RF, Minter DW, Stadler M, Gené J, Guarro J, Cano J. 2010. Two new anamorphic

fungi from Cuba: Endophragmiella profusa sp. nov. and Repetoblastiella olivacea gen. & sp. nov.

Mycotaxon 113: 415-422. http://dx.doi.org/10.5248/113.415

Chen JL, Tzean SS, Lin WS. 2008. Endophragmiella multiramosa a new dematiaceous anamorphic

ascomycete from Taiwan. Sydowia 60: 197-204.

Hughes SJ. 1979. Relocation of species of Endophragmia auct. with notes on relevant generic names.

New Zealand J. Bot. 17: 139-188.

Kirk PM. 1981. New or interesting microfungi: I. Dematiaceous hyphomycetes from Devon. Trans.

Br. Mycol. Soc. 76: 71-87. http://dx.doi.org/10.1016/S0007-1536(81)80010-1

Leao-Ferreira SM, Gusmao LFP. 2010. Conidial fungi from the semi-arid Caatinga biome of Brazil.

New species of Endophragmiella, Spegazzina and new records for Brazil, South America and

Neotropica. Mycotaxon 111: 1-10. http://dx.doi.org/10.5248/111.1

Sutton BC. 1973. Hyphomycetes from Manitoba and Saskatchewan, Canada. Mycol. Pap. 132:

1-143.

Wu WP, Zhuang WY. 2005. Sporidesmium, Endophragmiella and related genera from China. Fungal

Divers. Res. Ser. 15: 1-351.

MY COTAXON

http://dx.doi.org/10.5248/119.109

Volume 119, pp. 109-116 January-March 2012

Alveariospora, anew anamorphic genus from trichomes

of Dimorphandra mollis in Brazil

MEIRIELE DA SILVA’, RAFAEL F. CASTANEDA-RUIZ?’,

OLINTO LIPARINI PEREIRA', ROBERT WEINGART BARRETO*

"Departamento de Fitopatologia, Universidade Federal de Vicosa, Minas Gerais, 36570-000, Brazil

?Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt,

Calle 1 Esquina 2 Santiago de Las Vegas, La Habana, C.P. 17200, Cuba

* CORRESPONDENCE TO: rbarreto@ufv.br

ABSTRACT — A dematiaceous fungus associated with trichomes on leaflets of Dimorphandra

mollis was collected during a survey of fungi associated with the genus Dimorphandra

(Fabaceae). The host tree is endemic to the Brazilian Cerrado. Although widely distributed

there, this plant has been neglected in mycological studies, since no fungus has ever been

recorded in association with it. The new fungus produces large alveariform (skep or beehive-

shaped), muriform, dictyosporous, distoseptate, verruculose conidia from large, thickened,

dark washer-like conidiogenous loci. Among previously described anamorphic genera,

this fungus is somewhat similar to Annellophragmia, Annellosympodia, Briansuttonia,

Dictyospiropes, and Veracruzomyces as well as Spiropes dictyosporus but has some significant

differences from those taxa. Hence, the new genus Alveariospora (type species: A. distoseptata

sp. nov.) is proposed for this fungus.

Key worps — fungal survey, taxonomy, tropical fungi

Introduction

Dimorphandra mollis Benth. (Fabaceae: faveira, faveiro-do-cerrado,

falso barbatim4o) is a plant native to the Cerrado. This species is adapted to

conditions of low rainfall, developing foliage, flowering, and fruiting in the wet

season and losing its leaves in the dry season (Lorenzi 2002). The fruits are

utilized to extract rutin and other glycoside flavonoids for the pharmaceutical

industry (Ferreira et al. 2001; Lorenzi 2002). It has potential for exploitation as

a source of gum for the food industry (Panegassi et al. 2000). Although D. mollis

is a common component of the cerrado flora — an ecosystem that in recent

decades has been intensively explored by Brazilian mycologists (e.g., Rezende

& Dianese 2003, Dornelo-Silva & Dianese 2004, Hernandez-Gutiérrez &

Dianese 2009, Pereira-Carvalho et al. 2010) — no attention has been paid to the

mycobiota of this host plant and no fungi have ever been recorded on D. mollis.

110... Silva & al.

In July 2009, an exploratory survey of the mycodiversity of Dimorphandra spp.

was initiated, and the first brief survey to the municipalities of Paraopeba and

Caetandpolis (Minas Gerais, Brazil) has already yielded several fungi of interest.

One interesting fungus, found colonizing D. mollis trichomes, is described and

discussed herein.

Material & methods

Samples of D. mollis foliage were collected, dried in a plant press, and taken to the

lab. After examination, selected leaflets bearing fungus colonies were deposited in the

local herbarium (Herbarium VIC). Isolations were attempted by direct spore transfer

onto plates containing VBA medium (Pereira et al. 2003) with the help of a fine needle.

Fungal structures were removed from fresh leaves and mounted in lactophenol.

Observations, measurements, and illustrations were carried out with an OLYMPUS BX

51 light microscope fitted with a digital camera (EVOLT E330) and a drawing tube.

Wherever possible, 30 measurements of each structure were taken.

Taxonomy

Alveariospora Meir. Silva, R.F. Castafieda, O.L. Pereira & R.W. Barreto, anam. gen.

nov.

MycoBank 518826

Cellulae conidiogenae primo monoblasticae, deinde polyblasticae, indeterminatae,

cum, cicatricibus primo apicalibus, deinde lateralibus, discoidibus, protuberantibus,

nigris, incrassatis. Conidia alveariformia, muriformia, distoseptata, cum appendicibus

cellularibus, praedita.

TYPE SPECIES: Alveariospora distoseptata Meir. Silva et al.

ErymMo_oey: Latin, alvearium, meaning beehive, skep; and spora, meaning spores.

Anamorphic fungi. Colonies on the natural substrate effuse, hairy, brown,

olivaceous or black. Mycelium partly superficial and immersed. Conidiophores

distinct, single, unbranched,septate, brownorolivaceous,smooth.Conidiogenous

cells integrated, terminal, at first with a single terminal conidiogenous locus,

then indeterminate, polyblastic, with successive sympodial but rectilinear

proliferation, rupturing the outer wall around each scar, resulting in a lateral

displacement of scars, leaving conspicuous circumferential annular fringes of

the torn wall. Conidiogenous loci evident, lenticular, protuberant, thickened

and black, conidial secession schizolytic. Conidia solitary, ellipsoidal, oval to

broadly navicular, dictyoseptate, distoseptate, verruculose or smooth, brown or

dark brown, conspicuously cicatrized at the base, with a cellular, cylindrical or

subulate, brown apical appendage.

Alveariospora distoseptata Meir. Silva, R.F. Castafieda, O.L. Pereira & R.W. Barreto,

anam. sp. nov. PLATEs 1-4

MycoBAank 518827

Differs from Spiropes dictyosporus in hyphal conidiomata and distoseptate conidia.

Alveariospora distoseptata gen. & sp. nov. (Brazil) ... L111

Type: Brazil, Minas Gerais, Paraopeba, Floresta Nacional de Paraopeba, on trichomes of

Dimorphandra mollis, 10 Jul. 2009, M. Silva & O.L. Pereira (Holotype, VIC 31399).

ErymMo_oey: Latin, distoseptata, meaning distoseptate, having the individual cells each

surrounded by a sac-like wall distinct from the outer wall.

Anamorphic fungus. Colonies on the natural substrate effuse, hairy, brown.

Mycelium partly superficial and partly immersed, composed of septate,

branched, smooth, brown hyphae. Conidiophores distinct, single, unbranched,

erect, straight, 3—-6-septate, dark brown, pale brown near the apex, smooth,

120-160 x 7-10 um. Conidiogenous cells integrated, terminal, 22—45 x 6-10

um, at first with a single terminal conidiogenous locus, then indeterminate,

polyblastic, with successive sympodial but rectilinear proliferation, rupturing

the outer wall around each scar, resulting in a lateral displacement of scars,

leaving conspicuous circumferential annular fringes of the torn wall.

Conidiogenous loci evident, lenticular, black, 6—8 um diam., 2—4 um thick, 3-7

per conidiophore. Conidial secession schizolytic. Conidia solitary, ellipsoidal

to broadly navicular, dictyoseptate, distoseptate, verruculose, brown, 62-85

x 26-45 um, with a conspicuous protuberant, melanized, lenticular hilum

at the base, 7-8 um diam., 3—4 um thick, and with a cellular, cylindrical or

slightly subulate, smooth terminal appendages, 27-42 x 2.5—4.5 um, brown

and pale brown to subhyaline at the end, sometimes surrounded by a hyaline,

mucilaginous sheath. Teleomorph unknown. In culture, no conidial germination

was observed, even after a period of four weeks.

CoMMENTS — The pattern of ontogeny in Alveariospora can be classified as

conidial development type 17 (Kirk et al. 2008) (holoblastic, delimitation by 1

septum, schizolytic secession, maturation by diffuse wall-building, percurrent

enteroblastic conidiogenous cell extension, followed by conidial ontogeny by

replacement apical wall-building; strongly melanized, each successive conidium

seceding before the next percurrent elongation of the conidiogenous cell), but

sometimes also holoblastic, sympodial proliferation occurring and two or more

conidia are produced, a pattern of ontogeny classified as conidial development

type 10 (holoblastic, regularly alternating with sympodial proliferation,

maturation by diffuse wall-building and secession schizolytic).

The mode of rectilinear proliferation and the combination of conspicuous

annular structures and thickened, darkened conidiogenous loci in Alveariospora

distoseptata is very unusual and comparable only with Annellophragmia Subram.

(Ellis 1971), Annellosympodia McTaggart et al. (McTaggart et al. 2007), and

Spiropes dictyosporus Seifert & S. Hughes (Seifert & Hughes 2000). However,

the last species, inhabiting a sooty mould in New Zealand, is characterized

by synnematous conidiomata and euseptate conidia. Annellophragmia is

differentiated by synnematous conidiomata and phragmosporous (but also

distoseptate) conidia. Annellosympodia is characterized by its sporodochial

112... Silva & al.

PLaTE 1. Alveariospora distoseptata. Macronematous conidiophores with polyblastic

conidiogenous cells (A); detail of protuberant lenticular loci on enteroblastic percurrent

proliferations (B); initial conidial formation (C) and initial conidial septation (D). Bars: 10

um.

conidiomata with ampulliform, doliiform to obovoid conidiogenous cells, 0-

l1-euseptate conidia, and rhexolytic conidial secession. Among other known

anamorphic genera, Alveariospora appears superficially similar to Briansuttonia

Alveariospora distoseptata gen. & sp. nov. (Brazil) ... 113

PLATE 2. Alveariospora distoseptata. Detail view of muriform, distoseptate conidia (left) with

verruculose wall (right) on young yellowish conidia (A) and reddish-brown mature conidia

(B). Bars: 10 um.

R.F Castaneda etal. (Castaneda et al. 2004), Dictyospiropes M.B. Ellis (Ellis 1976),

and Veracruzomyces Mercado et al. (Mercado-Sierra et al. 2002). Briansuttonia

is characterized by monotretic, terminal, determinate or indeterminate with

114... Silva & al.

PLATE 3. Alveariospora distoseptata. Conidial appendage ends, sometimes surrounded by a

hyaline, mucilaginous tunica (A); germination of the conidia, through hilum (B) and laterally

through wall (C). Bars: A = 5 um; B = 20 um; C = 10 um.

enteroblastic percurrent conidiogenous cells. Dictyospiropes is quite distinct

from Alveariospora by having polyblastic, sympodial conidiogenous cells with

strongly melanized, lenticular conidiogenous loci lacking percurrent extensions,

Alveariospora distoseptata gen. & sp. nov. (Brazil) ... 115

PiatE 4. Drawing of Alveariospora distoseptata conidia and conidiogenous cells. Bar: 10 um.

and euseptate-dictyoseptate conidia without strongly thickened and melanized

basal hila. Veracruzomyces is clearly distinguished from Alveariospora by its

monoblastic, percurrently proliferating conidiogenous cells without cicatrized

conidiogenous loci.

Acknowledgments

The authors wish to thank Bryce Kendrick (University of Waterloo) and Uwe

Braun (Martin-Luther- Universitat), for reviewing the manuscript. U. Braun is also

acknowledged for his help on formulating a name for the newly described genus. This

116... Silva & al.

work was conducted with the permission of ICMBIO (permit n° 30022). The authors

wish to thank the administration of Floresta Nacional de Paraopeba for the use of

facilities. Financial support from the Fundacao de Amparo a Pesquisa do Estado de

Minas Gerais - FAPEMIG and the Conselho Nacional do Desenvolvimento Cientifico e

Tecnoldgico (CNPq) is also acknowledged.

Literature cited

Castafieda Ruiz RF, Heredia GP, Arias RM, Saikawa M, Minter DW, Stadler M, Guarro J, Decock

C. 2004. Two new hyphomycetes from rainforests of México, and Briansuttonia, a new genus to

accommodate Corynespora alternarioides. Mycotaxon 89(2): 297-305.

Dornelo-Silva D, Dianese JC. New hyphomycete genera on Qualea species from the Brazilian

cerrado. 2004. Mycologia 96(4): 879-884. http://dx.doi.org/10.2307/3762120

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey.

Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew,

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MY COTAXON

http://dx.doi.org/10.5248/119.117

Volume 119, pp. 117-132 January-March 2012

Intraornatosporaceae (Gigasporales), a new family

with two new genera and two new species

BRUNO T. GOTO’, GLADSTONE A. SILVA”, DANIELE M.A. DE Assis’,

DANIELLE K.A. SILVA’, RENATA G. SOUZA’, ARAESKA C.A. FERREIRA’,

KHADIJA JOBIM', CATARINA M.A. MELLO’, HELDER E.E. VIEIRA’,

LEONOR C. MATA? & FRITZ OEHL?

"Departamento de Botanica, Ecologia e Zoologia, CB, Universidade Federal do Rio Grande do Norte,

Campus Universitario, 59072-970, Natal, RN, Brazil

*Departamento de Micologia, CCB, Universidade Federal de Pernambuco,

Av. Prof. Nelson Chaves s/n, Cidade Universitaria, 50670-420, Recife, PE, Brazil

*Federal Research Institute Agroscope Reckenholz-Tdanikon ART, Organic Farming Systems,

Reckenholzstrasse 191, CH-8046 Ziirich, Switzerland

“CORRESPONDENCE TO: brunogoto@hotmail.com & gladstonesilva@yahoo.com

AxBstTRAcT — A new family (Intraornatosporaceae), two new genera (Intraornatospora,

Paradentiscutata), two new species (PR. bahiana, P. maritima), and a new combination

(I. intraornata) are presented in the Gigasporales. The genera, both with diagnostic introverted

ornamentations on the spore wall, are distinguished by spore wall structure and germ

shield characteristics. The new species, detected in NE Brazil, can be differentiated by their

projections on the outer spore surface. Partial sequences of the LSU rRNA gene place both

species next to I. intraornata in a monophyletic major clade related to Gigasporaceae and

Dentiscutataceae.

Key worps — Glomeromycetes, Scutellospora, molecular phylogeny, rDNA

Introduction

The phylum Glomeromycota has been extensively revised in recent years

at all classification levels from class to species (e.g. Morton & Redecker 2001,

Sieverding & Oehl 2006, Palenzuela et al. 2008, Goto et al. 2010, 2011, Oehl

et al. 2011a,b,c). Major revisions were necessary within the order Gigasporales

when separating Gigasporaceae species into four families (Gigasporaceae,

Scutellosporaceae, Racocetraceae and Dentiscutataceae) and Scutellospora into

seven genera when concomitant morphological and molecular spore analyses

were undertaken (Oehl et al. 2008, 2010, 2011b,c). It was initially clear that

Dentiscutataceae, in particular, was a heterogeneous family that needed more

118 ... Goto & al.

detailed study (Oehl et al. 2008). Unfortunately, little progress has been made

due to only a limited number of available isolates.

During recent studies on the diversity of the arbuscular mycorrhizal

(AM) fungi (Glomeromycota) in Northeastern Brazil, Dentiscutata colliculosa

B.T. Goto & Oehl, Racocetra intraornata, and two undescribed species were

detected (Goto et al. 2009, 2010). All four species have introverted projections

on the spore wall. The phylogenetic position of R. intraornata, however,

remained unresolved while the two new species appeared to belong to the

Dentiscutataceae, based on their spore formation on sporogenous cells and the

presence of yellow brown to brown shields on the inner spore walls. Moreover,

two major morphological characters separated these two new species within

the Dentiscutataceae — the multiple-lobed structure of their openly organized

shields and the presence of a tuberculate ornamentation on the inner surface of

the hyaline middle spore wall. A multiple-lobed structure of openly organized

shields has been detected only in the Racocetraceae, where shields are hyaline to

sub-hyaline. Ornamentation on the inner surface of the middle wall is hitherto

a unique character shared with no other species within the Glomeromycota.

Racocetra intraornata, however, was understood to have a rather rudimentary

germination shield where germ tube initiations are rarely found. The question

therefore arose whether this species truly belongs to Racocetra, or might have

a different phylogeny.

The main aim of this study was thus to analyze thoroughly and describe the

two new species and to elucidate their phylogenetic position as well as that of

R. intraornata.

Materials & methods

Study areas, soil sampling and soil parameters

One of the two new AM species was recovered in Serra da Jibdia, a fragment of the

Atlantic Rainforest (“Mata Atlantica’) in Santa Terezinha, Bahia State. The other species

was found in several locations in the sand dunes of Mataraca, Paraiba State, (Souza et

al. 2010, 2012) and in Natal, Rio Grande do Norte State. In the Serra da Jibdia, the soils

were taken in September 2010, in Mataraca in January and August 2005, and in Natal in

February 2010, as described in (Souza et al. 2010, 2012).

The Serra da Jibdia (12°51'S 39°28'W) is a forest fragment characterized by high

plant diversity (Queiroz et al. 1996) located on a hillside surrounded by drier areas

in transition to Caatinga (dry forest with trees and shrubs, deciduous during the dry

season; Andrade-Lima 1964). Mean annual temperature is 17-28°C with 1200 mm

of rain falling between April and July (Tomasoni & Santos 2003). The sand dunes in

Mataraca are located at 6°28’-6°30'S 34°55'-34°57'W. ‘The vegetation is typical of

‘restinga, a transition ecosystem between the primary coastal sand dunes and the tropical

Atlantic Rainforest with physiognomy varying from tree-shrub to herbaceous plants

(Oliveira-Filho & Carvalho 1993). The predominant geological formation is sandy-clay

Intraornatosporaceae fam. nov. (Gigasporales) ... 119

sedimentary rocks, superimposed by fixed dunes. The climate is tropical-(semi-)humid

(type Am of Képpen-Geiger, Kottek et al. 2006) with four months of dry season. Average

annual temperature is 24—28°C and rainfall 1795 mm. The sand dunes in Natal represent

one of the largest urban conservation areas with dune vegetation in Brazil. The “Parque das

Dunas’ site (5°46’S and 35°12'W) also has typical ‘restinga’ vegetation. The climate is

tropical rainy (type Am of Koppen) with a short dry period of four months. The mean

annual temperature is 25.5 °C, and the mean annual precipitation is 1191 mm.

Soil pH (H,0) in Santa Terezinha was 4.6, organic C was 21.5 g kg’ and available P

(after Mehlich; Nelson et al. 1953) 2.0 mg kg’. In Mataraca, soil pH was 5.6, organic C

25.0 g kg’ and available P 6.5 mg kg’. In Natal, pH was 5.6, organic C 16.6 mg kg", and

available P was 3.0 mg kg". Soils in all three areas were sandy.

AM fungal bait cultures

The native AM fungal communities from Serra da Jibdia and Mataraca were cultured

in bait (= trap) cultures over three consecutive cycles (three months each) using corn

(Zea mays L.), peanut (Arachis hypogaea L.) and sunflower (Helianthus annuus L.) in

500 mL pots, filled with autoclaved sand-vermiculite substrate (1:1 w/w; 400 g per

pot) mixed with the natural field soil as AM fungal inoculum (50 g per pot), in the

greenhouse of the Department of Mycology, Universidade Federal de Pernambuco,

Recife. Additionally, multiple glomerospores of the new species were separated and

used as infective propagules in single species cultures on Sorghum bicolor (L.) Moench

as described in Palenzuela et al. (2010), Tchabi et al. (2010), and Goto et al. (2011). The

new Serra da Jibdia species was successfully propagated in a two-species culture together

with Ambispora appendicula, and the sand dune species in a bait culture together with

Racocetra tropicana (Goto et al. 2011). However, no single species cultures were obtained

from the two fungi. Spores isolated from the cultures or directly obtained from the field

samples were used for morphological and molecular analyses.

Morphological analyses

Spores of the two new species and R. intraornata were extracted according to

Sieverding (1991) and thereafter mounted in polyvinyl-alcohol-lacto-glycerin (PVLG),

in PVLG + Melzer’s reagent, and in water (Brundrett et al. 1994, Spain 1990). Over

100 spores were examined per species. The germination shields of each species were

carefully separated from their spores by applying pressure and movements to cover

slides in PVLG. Species descriptions utilize spore and germination terminology for

gigasporalean species defined by Oehl et al. (2008, 2010, 2011b), Silva et al. (2008), and

Goto et al. (2010, 2011).

Molecular analyses

DNA was extracted from single spores, with three single spore extractions for each

of the two new species and one for R. intraornata. Individual spores were placed on a

slide in a drop (5-10 ul) of ultrapure water, crushed with a needle, and used directly in

the PCR reactions.

The extracts served as templates for a semi-nested PCR using primers ITS3 (White et

al. 1990) - 28G2 (Silva et al. 2006) and LR1 (van Tuinen et al. 1998) - 28G2 consecutively.

The template for the second PCR reaction was a 1:50 dilution of the first product. PCR

reactions were carried out in a volume of 50 ul, containing 75 mM Tris-HCl pH 8.8, 200

120 ... Goto & al.

Pacispora scintillans FM876832

P. scintillans FM876831 100

400 Orbispora pernambucana JF965445

70| 100| O. pernambucana JF965446

ad 1.00! O. pernambucana HQ871519

Scutellospora calospora FJ461864

31 S. calospora FJ461865

0.89 S. dipurpurescens FJ461868

S. calospora EU346867

S. calospora EU252109

100 Racocetra fulgida FJ461870

gsi R. gregaria AJ510232

100|'— R. coralloidea FJ461866

1.00] R. persica FJ461880

R. alborosea JN689226

100 R. tropicana GU385898

100} gg R. castanea Y12076

By tg 81 R. verrucosa AY900507

i 91 R. verrucosa AY900508

4.00 Cetraspora nodosa FM876833

400

400! C. nodosa FM876836

700 C. helvetica HM565945

85| (0-91 C. helvetica HM565946

66) C. helvetica HM565944

4 C. pellucida AY639323

91- C. pellucida AY639261

89 | C. gilmorei FN547608

64 4 od|' C. gilmorei FN547606

6: ag C. gilmorei FN547603

75 ose 98 | C. gilmorei FN547618

63 99 - Dentiscutata nigra AY900495

aon 98 N09 | D. nigra AY900497

j 81 | Quatunica erythropus AM040354

véies | Q. erythropus AM040357

65 60 Fuscutata heterogama AY900503

91 aad F. heterogama FJ461877

1.00 i is F. heterogama FJ461871

M F. heterogama DQ273792

Gigaspora rosea Y12075

67| 94 G. albida FJ461861

0.95} 92 G rosea AM040350

9 G. gigantea AY900506

-00 G. margarita AF396782

G. decipiens FJ461862

7 G. gigantea AY900504

G. margarita AF396783

Intraornatospora intraornata JN971072

97 i |. intraornata JN971074

4.00 I. intraornata JN971073

Mie |. intraornata JN971075

64 100, Paradentiscutata bahiana JN971070

70 100! p pahiana JN971071

0.73) | 4.09 . bahiana JN971069

85| + P. bahiana JQ231202

97 P. bahiana JQ231201

4.00 P. bahiana JQ231203

85 P. maritima JN971081

82L}; P. maritima JN971077

85 |! P. maritima JN971078

P. maritima JN971079

P. maritima JN971080

P. maritima JN971076

0.1

Fic. 1. Phylogenetic tree of the Gigasporales based on LSU rDNA analysis and rooted by Pacispora

scintillans. Sequences are labeled with database accession numbers. Support values (from up to

down) are from neighbor-joining (NJ), maximum parsimony (MP), maximum likelihood (ML)

and Bayesian analyses, respectively. Intraornatospora intraornata, Paradentiscutata bahiana

and P. maritima sequences are in bold. Only topologies with 250% bootstrap values are shown.

(Consistency Index = 0.53; Retention Index = 0.79).

Intraornatosporaceae fam. nov. (Gigasporales) ... 121

mM (NH,),SO,, 0.01% Tween 20, 2 mM MgCl, 200 uM each dNTPs, 1uM of each primer

and 2 units of Taq™ DNA polymerase (Fermentas; Maryland, USA); cycling parameters

were 5 min at 95°C (1 cycle), 45s at 94°C, 1 min at 55°C, 1 min at 72°C (40 cycles), and

a final elongation of 7 min at 72°C followed the last cycle. The final amplicons (~690bp)

were purified with a PureLink PCR Purification Kit (Invitrogen), sequenced directly or

cloned with a CloneJET.,., PCR Cloning kit (Fermentas; Carlsbad, USA) following the

manufacturer's instructions and sequenced. Sequencing was provided by the Human

Genome Research Center (Sao Paulo, Brazil).

Through a BLASTn query of the National Center for Biotechnology Information

databases, we verified that the sequences obtained from P maritima, P. bahiana, and

R. intraornata afhliated with the Gigasporales (Glomeromycota) before phylogenetic

analysis. The AM fungal sequences (partial LSU rRNA) obtained were aligned with

other glomeromycotean sequences from GenBank using ClustalX (Larkin et al. 2007)

and edited with BioEdit (Hall 1999). The sequences were deposited at GenBank under

the accession numbers JN971069-JN971081 and JQ231201-JQ231203.

Maximum parsimony (MP) and neighbor joining (NJ) analyses with 1000 bootstrap

replications were performed using the Phylogenetic Analysis Using Parsimony (PAUP)

vers. 4 (Swofford 2003). Bayesian (two runs over 1 x 10° generations with a burnin value

of 2500) and maximum likelihood (1000 bootstrap) analyses were executed, respectively,

in MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003) and PhyML (Guindon & Gascuel

2003), launched from Topali 2.5. The model of nucleotide substitution (GTR + G) was

estimated using Topali 2.5 (Milne et al. 2004). Sequences from Pacispora scintillans were

used as an outgroup.

Results

Molecular analyses

The phylogenetic analyses generated by the LSU rRNA gene sequences

(Fic. 1) revealed that the three species grouped in a monophyletic major clade

most closely associated with the Gigasporaceae and Dentiscutataceae clade, with

75% bootstrap value for at least one phylogenetic method. The data further

confirmed that the new genera and two new species conferred bootstrap values

above 79%, thus supporting a new family and two new genera within the

Gigasporales (Oehl et al. 2011b). We hereafter describe the two new species in

one new genus and transfer R. intraornata from Racocetra to another, hitherto

monospecific genus as the type species of the new family.

Taxonomic analyses

Intraornatosporaceae B.T. Goto & Oehl, fam. nov.

MycoBank MB 563599

Sporae ad cellulas sporogeneas, tunicis duabus vel tribus; ornamentatione introvertita

in superficie interiore tunicae mediae (tribus tunicis) vel exterioris (duabus tunicis);

scutellum germinale multilobatum, flavum-brunneum ad brunneum in sporis tribus

tunicis, hyalinum vel subhyalinum duabus tunicis.

TYPE SPECIES: Intraornatospora intraornata (B.T. Goto & Oehl) B.T. Goto et al.

122 ... Goto & al.

Sporocarps are unknown. Spores formed singly on bulbous sporogenous cells,

terminally on a subtending hypha that arise from mycelial hyphae. Spores have

2 or 3 walls. The outer spore wall has 3-4 layers and inner wall 2-3 layers.

Middle wall of 3-walled spores has an expanding outer and an inner layer

with tuberculate ornamentation towards the inner wall. Outer wall of 2-walled

spores also has introverted tuberculate ornamentation. Germination shield

generally formed on the outer surface of the innermost wall or beneath a thin

outer layer of the inner wall. Shields of triple-walled spores yellow-brown to

brown, with 4-8(-10) wave-like lobed projections; folds separate the lobes on

the shield, and each lobe may have a germ tube initiation from where the germ

tubes arise and penetrate the outer wall. Shield in bi-walled spores, hyaline to

light yellow, with 4-8 wave-like lobed projections; folds separate the lobes on

the shield, which often appears immature, even in more aged spores.

TYPE GENUS: Intraornatospora B.T. Goto et al.

OTHER GENUS: Paradentiscutata B.T. Goto et al.

Intraornatospora B.T. Goto, Oehl & G.A. Silva, gen. nov. Fics 2-4

MycoBank MB 563600

Sporae ornamentatione introvertita in superficie interiore tunicae exterioris; scutellum

germinale superficie tunicae interioris, multilobatum hyalinumque.

TYPE SPECIES: Intraornatospora intraornata (B.T. Goto & Oehl) B.T. Goto et al.

EryMo_oey: Latin: intra (= inside), ornata (= ornamented), spora (= spore); referring to

the position of the ornamentation on the spore in the type species.

Spores formed singly in soils, rarely in roots, on bulbous sporogenous cells that

arise terminally on mycelial hyphae. The spores have two walls. The outer spore

wall is generally triple-layered and continuous with the wall of the sporogenous

cell. Inner surface of the outer spore wall has tuberculate or spiny projections.

The inner wall is hyaline, 2-3 layered and forms de novo. A germination shield

arises on the outer surface of the inner wall or beneath a thin outer layer of

the inner wall; shield is hyaline to light yellow, generally oval to ellipsoid or

subglobose, with several wave-like lobed projections forming the outer surface

of the shield; folds separate the lobes on the shield. Germ tube initiations within

the lobes often appear rudimentary and are rarely observed.

Intraornatospora intraornata (B.T. Goto & Oehl) B.T. Goto, Oehl & G.A. Silva,

comb. nov. FIGS 2-5

MycoBank MB 563601

= Racocetra intraornata B.T. Goto & Oehl, Mycotaxon 109: 485. 2009.

Paradentiscutata B.T. Goto, Oehl & G.A. Silva, gen. nov.

MycoBank MB 563602

Sporae ornamentatione introvertita in superficie interiore tunicae mediae; stratum exterius

tunicae mediae expansivum; scutellum germinale, multilobatum, (flavo-)brunneum.

Intraornatosporaceae fam. nov. (Gigasporales) ... 123

2g oe

Fics 2-4. Intraornatospora intraornata (isotype, ZT Myc 775): typical, multiple-lobed germination

shields; Germ pore (gp) as connection between shield and cell contents regularly visible when

shield is in planar view (Fics 2-3); germ tube initiations rarely if ever visible.

TYPE SPECIES: Paradentiscutata bahiana Oehl et al.

EryMmo ocy: Latin: para (= equal), dentata (= dentate), scutata (= with shield); referring

to the similarities with the germination shields of spores of Dentiscutataceae.

Sporocarps are unknown. Spores formed singly on bulbous sporogenous cells,

terminally on subtending hypha that arise from mycelial hyphae. Spores have

three walls; outer spore wall with 3-4 layers, middle wall with an expanding

outer and a tuberculate ornamentation towards the inner wall, and an inner

wall with 2-3 layers. Germination shields are yellow-brown to brown, as in

Dentiscutataceae, but with 4-8(-10) wave-like lobed projections forming the

outer surface of the shield; folds separate the lobes on the shield, and each lobe

may have a germ tube initiation from where the germ tubes arise and penetrate

the outer wall.

Paradentiscutata bahiana Oehl, Magna, B.T. Goto & G.A. Silva, sp. nov.

MycoBank MB 563603 FIGs 5-20

Sporae aurantio-brunneae; stratum exterius verrucis, < 1.0 um in distantia.

Type: BRAZIL. Bahia State, Santa Terezinha, Serra de Jibdia, tropical rainforest fragment,

[September 2010], [G.A. Silva], 92-9201 (Holotype, URM 83317); Isotypes: 92-9202

(URM 83318), 92-9203 (URM 83319), 9204-9212 (ZT Myc 7624).

EryMo.oey: referring to Bahia State (NE Brazil), where the fungus was first found.

GLOMEROSPORES formed singly in soil, terminally on a subterminal bulbous

suspensor cell (= ‘sporogenous’ cell). Glomerospores are dark orange brown to

brown, globose, 190-260 um in diameter, to subglobose, 190-265 x 190-240

um, and have three walls (an outer, middle and inner wall). The spores become

dark black brown to black when exposed to Melzer’s reagent.

OUTER WALL (Ow) is 3.6-6.0 um thick in total and consists of 2(-3) layers:

outermost wall layer (ow11) is hyaline to subhyaline to light yellow, semi-

124 ... Goto & al.

persistent to persistent and 0.9-1.5 um thick. It is covered with densely crowded

irregular warts that are 0.8-2.0(-2.5) um high and 1.0-2.1(-3.5) x 1.0-2.5(-5.0)

um wide. OWL2 is 2.1-3.4 um thick, orange brown to brown, becoming dark

orange brown to dark reddish brown when exposed to Melzer’s reagent. owL3

is concolorous and adherent with ow12, very thin (0.6-1.1 tm), and generally

not observed even in crushed spores.

MIDDLE WALL (Mw) is hyaline to rarely light yellow, bi-layered and 4.8-6.8

um thick when mounted in water. MWL1 is hyaline and 3.8-5.2 um thick when

mounted in water. In crushed spores, it expands to 9.0-16.0 um and regularly

expands further to 20-32(-56) um in lactic acid based mountants. MwL1] may

even become completely transparent making it difficult to detect due to even

stronger swelling processes. MwL2 is hyaline becoming light yellow in older

spores, 1.0-1.6 um thick and bears fine tubes on its inner surface that are 0.5-

1.1 um high, 0.4-1.0 um wide and 1.0-2.5 um spaced. MwL] stains purple black

to black in Melzer’s reagent.

INNER WALL (Iw) is triple-layered, 2.8-4.5 um thick bearing a germination

shield on the outer surface. The outer rw layer (IWL1) is hyaline, semi-flexible,

0.6-1.1 um thick and often wrinkles in crushed spores when separated from

IWw12. The second layer (Iw12) is semi-flexible to unite, rarely ‘amorph when

slightly expanding in PVLG based mounting, and is 1.6-2.2 um thick. The

innermost layer (tw13) is relatively thin (0.6-1.2 um thick), flexible, mostly

tightly adherent to 1w12, and therefore generally difficult to observe. rwL2 and

IWL3 stain yellow-pink to light purple in Melzer’s reagent.

SPOROGENOUS CELL (sc) is globose to elongate, 32-50 um long and 28-38 um

broad. It is concolorous with the spore wall. Two wall layers are visible on the

young sporogenous cell, which are continuous with owt] and with laminated

OWL2. OWL] at sc is 0.4-1.2 um thick, smooth and semi-persistent; Ow1L2 is

1.0-2.5 um thick and persistent as long as sc remains attached on the spore.

One to two ‘hyphal pegs’ may be rarely detected on the sporogenous cells. The

sporogenous hypha attached to the cell is also bi-layered, 12-21 um in diameter,

tapering to 7-11 um within 100-250 um distance from the sporogenous cell.

Within this distance, the sporogenous hyphal wall tapers from 1.5-2.2 um to

1.0-1.7 um, and 4-9 septa originating from OwL2 may be visible in variable

distances within the sporogenous hypha. The ornamentation on OwL1 does not

continue on the wall of the sporogenous cell.

GERMINATION SHIELD is yellow brown to brown, subglobose to oval to

cardioid, to rarely ellipsoid or oblong, 140-200 x 121-180 um in diameter, and

multiple-lobed, generally with (4—)6-10 lobes. The shield is open-organized, as

Fics 5-20. Paradentiscutata bahiana (Type URM 83317-83319, ZT Myc 7624): 5-10. Spores

(crushed) formed on sporogenous cells (sc) with surface ornamentation (orn), showing outer wall

(ow), middle wall (mw), inner wall (tw) and conspicuous, yellow brown to brown germination

shields (gs). Each two layers of Mw (MWL1-2) and Iw (IWL1-2) visible. MWLI expanding in lactic

Intraornatosporaceae fam. nov. (Gigasporales) ... 125

fit 2 MWe

gs rf 4 Wer ‘a

— 4 ‘ % De a : 4

i Agee

4 “ ae

% ‘ ¥ fo SD Swi

Iw ‘ ia gow

. . Orang,

200 ym 200 "fim -

WL 2am /

F iwi IWL2

wt 4 ~ 3 " ee eS . -* ;

ae! vy t We Ae Se t By ee. .

100.4m 100 pm we; 100 ay 2 f MWL1™ ~400-ym s

acid based mountants; MwL2 with needle-tuberculate ornamentation. 11-12. Verrucose surface

ornamentation (warts) in cross and planar view, respectively. 13. Ornamentation of MwL2 in planar

view. 14-16. Spore segments (crushed) in Melzer’s reagent; expanding MWL1 stains dark purple to

black purple. 1w triple-layered (IWwL1-3; 1w12 staining light purple to purple. 17-20. Characteristic

germination shields are open-organized with multiple wavy-lobes, mostly bearing, each, one germ

tube initiation (gti) from where a germ tube (gt) emerges; lobes are separated by large folds (f), and

a single germ pore (gp) connects the shield with the cell content.

126 ... Goto & al.

known for species of Racocetraceae. Large folds (~11-45 um long) arise from

the shield wall separating the lobes. The one-layered shield wall and the folds

are generally 1.2-1.8 um thick. The shield periphery regularly appears slightly

dentate until the germination has started. Each lobe may bear one rounded

germ tube initiation (gti), 6.5-12.0 um in diameter. The majority of the gti’s

may remain undetectable in young spores, becoming increasingly visible with

age of spores. Single germination tubes may emerge from one, two to rarely

three gti’s during early germination, penetrating the OW and branching in the

spore periphery within a short distance.

ARBUSCULAR MYCORRHIZA forming (as proofed in dual-species cultures

together with Ambispora appendicula, after staining in trypan blue).

DISTRIBUTION: Paradentiscutata bahiana has been recovered only from

natural tropical rainforest in Santa Terezinha, Serra da Jibdia, Bahia State.

Paradentiscutata maritima B.T. Goto, D.K. Silva, Oehl & G.A. Silva, sp. nov.

MycoBAnk MB 563604 FIGS 21-31

Differt ad Paradentiscutata bahiana in ornamentatione superficialis: stratum exterius

papillis rotundatis, 3.0-8.0 um in distantia.

Type: BRAZIL. Paraiba State, Mataraca, Atlantic Rainforest biome, ‘restinga’ vegetation

in sand dunes, [August 2005], [R.G. Souza], 93-9301 (Holotype, URM 83320); Isotypes

9302 (URM 83321), 9303 (URM 83322), 9304-9310 (ZT Myc 7625).

ErymMo.oey: Latin: maritima (=maritime, coastal), referring to occurrence in sand

dune systems of the Atlantic coast.

GLOMEROSPORES formed singly in soils terminally on a subterminal or

intercalary bulbous suspensor cell. Spores are globose (150-260 um in

diameter) to subglobose (145-250 x 165-280 um), bright (yellowish) brown,

orange brown to dark brown, with three walls: outer, middle, and inner (ow,

Mw, and Iw).

OUTER WALL (ow) is three-layered: outermost wall layer (OWL1) is 0.5-1.2

um thick and has a papillae ornamentation with projections that are 1.5-2.5

um high and 1.2-2.6 um wide at base, and 3.0-8.0 um apart. OwL2 is yellow to

orange, persistent, 2.5-3.8 um thick and laminate. Ow13 is 0.5-1.0 um thick,

tightly adherent to ow12 and especially difficult to distinguish in lactic acid-

based mountants, also due to expansive MWL1 beneath. ow12 stains red to red-

brown in Melzer’s reagent. The straight pore channel at the spore base is about

2.5-4.2 um broad and is generally closed by a plug formed by ow1z2.

Fics 21-31. Paradentiscutata maritima (Type URM 83320-URM83322, ZT Myc 7625): 21. Spore

(crushed) formed on sporogenous cell (sc) with three walls (ow, Mw, Iw) and germination shield

(gs); Iw staining purple in Melzer’s reagent. Two layers of Mw (MWLI1-2) visible. 22-23. Spore

wall structure: ow triple-layered (OWL1-3), bi-layered mw, and triple layered rw (IWwL1-3); spore

surface ornamentation of OWL1, with dispersed, blunt papillae. 24. Blunt ornamentation in planar

view. 25-26. MWLI expanding in lactic acid based mountants; Mw12 with needle-tuberculate

Intraornatosporaceae fam. nov. (Gigasporales) ... 127

, 7 1 a wi , wos “- IWL1

“ 90 pm Bot \ ; > 50pm wo" “<i

Pe =

~~ . mo

a 7

75 pm Al 120 ym

ra ss ; i . ——

ie th diaee Gti — 3

ZH a gti te ' Jere 28 gti rer iW i—.

: '

ornamentation. Yellow brown to brown germination shields with germ tube initiations (gti).

27-30. Germ shields (planar view) with 4-8 lobes, mostly bearing, each, one germ tube

initiation (gti) from where germ tubes (gt) emerge; single lobes are separated by large folds (f),

and a single germ pore (gp) connects the shield with the cell content. 31. gti of the germ shield

partly visible on Mw12.

128 ... Goto & al.

MIDDLE WALL (Mw) is hyaline to rarely light yellow, bi-layered and 5.2-7.6

(-8.9) um thick when mounted in water. MWL] is hyaline, and 4.0-5.1 um thick

in water. It expands to 7.5-14.6 um in crushed spores and regularly expands

further to 20-28(-51) um in lactic acid based mountants. Mw1z is hyaline to

light yellow, 1.2-2.5(-3.8) um thick, and densely packed with small tuberculate-

warty projections on the inner surface. Projections are 1.0-1.5(-2.5) um long

and 0.5-1.0(-1.1) um broad. Warts are about (0.5-)1.0 (-2.5) um apart from

each other.

INNER WALL (Iw) is triple-layered bearing a germination shield on the

outer surface. Outer layer of the inner wall (1w11) is hyaline, semi-flexible and

0.5-1.2 um thick. Second layer (IwL2) is unit, semi-flexible and 1.2-2.5 um

thick. Innermost layer (twL3) is thin (0.4-0.8 um thick), flexible, generally

tightly adherent to rw12, and difficult to observe. tw12 stains reddish-brown to

purple in Melzer’s reagent.

SPOROGENOUS CELL is subglobose to elongate, concolorous with the spore,

or slightly lighter in color than the spore, and 46-64 um long and 71-76

(-79) um broad. Two wall layers are generally visible on the sporogenous

cell, continuous with owLl and owL2. OwL1 on the sporogenous cell is about

0.5-1.6 um; adherent OwL?2 is about 2.5-7.6(—10.2) um thick. The ornamentation

on OWL1 does not continue on the wall of the sporogenous cell. The pore of the

sporogenous cell is generally closed at the septum to the attached ‘sporogenous

hypha’ by a septum arising from OwL2.

GERMINATION SHIELD is yellow-brown to brown, cardioid to oval or

ellipsoid, 181-189 x 166-171 um in diameter, and has several 4-8(-10) lobes,

that are easy to differentiate as they are often wavy in planar view and separated

by large folds. These folds (about 5-15 um long) arise from the shield wall. The

one-layered shield wall and the folds are generally only 0.5-1.7 um thick. Each

lobe regularly bears one rounded germ tube initiation, 1.2-2.5 um in diameter,

from where the germination tubes emerge.

ADDITIONAL MATERIAL EXAMINED: BRAZIL. R10 GRANDE DO Norte, Natal, from

sand dunes (URM 83483-83485).

DISTRIBUTION: Paradentiscutata maritima has been recovered from two sand

dune locations in NE Brazil (Mataraca, Paraiba State, and Natal, Rio Grande do

Norte State).

Spore development in /ntraornatospora and Paradentiscutata

In I. intraornata the outer spore wall is continuous with the wall of the

sporogenous cell and the attached sporogenous hypha. The ornamentation on

the inner surface of the outer spore wall differentiates during spore formation.

The inner wall forms de novo during spore formation like an ‘endospore. In

Paradentiscutata species, the outer wall is also continuous with the wall of the

sporogenous cell and the sporogenous hypha attached. Although uncertain

Intraornatosporaceae fam. nov. (Gigasporales) ... 129

Fics 32-35. Dentiscutata biornata (isotype, OSC 49583): Typical germination shields have one

germ pore (gp) 12-24 compartments and 12-24 germ tube initiations (gti); compartments (dense

lobes) separated by large folds; shield periphery is dentate.

whether the middle wall forms completely de novo or differentiates from a thin

layer present on the inner surface of the outer wall, we did observe that during

spore formation MWL1 differentiates first, showing several laminae when

exposed to PVLG and other lactic acid based mountants. Mw12 differentiates

on the inner surface of developed Mw 1. Thereafter, rw is formed de novo, and,

in the final stage of spore development the germination shield differentiates on

the outer IW surface.

Discussion

Distinct spore morphologies were recognized for both Paradentiscutata

and Intraornatospora. Paradentiscutata spp. have three-walled spores with a

diagnostic tuberculate ornamentation on the inner surface of the middle wall

and an openly organized, multiple-lobed, yellow-brown to brown germination

shield on the outer rw surface. Intraornatospora intraornata has a tuberculate

ornamentation of the inner ow surface, and on the outer Iw surface a multiple

lobed, hyaline to light yellow germination shield that regularly appears to be

rudimentary even in older spores, since clearly visible germ tube initiations have

rarely been found (Fics 2-4). Germination itself has yet to be observed in spores

130 ... Goto & al.

of this species, so we do not know whether I. intraornata germinates through

the rudimentary germ shield or directly from the warty inner projections of the

outer wall (as in Gigaspora). The two new genera, morphologically somewhat

dissimilar, might better be separated into two families. However, the database

so far is too small to make a final conclusion on this aspect.

The two Paradentiscutata spp. can easily be distinguished through their

ornamentation on the outer spore surface, a character that clearly separates

species in other glomeromycete families (e.g., Racocetra verrucosa and

R. persica, Dentiscutata nigra and D. reticulata, Acaulospora scrobiculata and

A. cavernata; Oehl et al. 2008, 2011d). Of the Gigasporales species, P. maritima

resembles only Dentiscutata biornata, which differs in a compact germination

shield with multiple (12-24) compartments in mature spores (Fics 32-35)

that are not openly organized or multiple-lobed (Spain et al. 1989) as in

P. maritima, which has 4-8(-10) lobes. Additionally, the double ornamentation

of D. biornata belongs to the outer spore wall according to Spain et al. (1989).

These important morphological differences point out species dissemblance and

support the new speciation. However, further research may consider checking

the phylogenetic relationships between D. biornata and the two new species.

Species of Intraornatosporaceae, Dentiscutataceae, and Gigasporaceae are

predominantly found in warmer climates. The Intraornatosporaceae have so far

been detected only in tropical areas, while Dentiscutataceae spp. are known

also from sub-tropical and Mediterranean regions and Gigasporaceae spp. have

at times been recovered from temperate regions (Jansa et al. 2002, Oehl et al.

2010). It is possible that Dentiscutataceae and Gigasporaceae may have evolved

to cooler periods and climates than Intraornatosporaceae. Further AM fungal

diversity research will also help verify this hypothesis.

Acknowledgments

The authors acknowledge, in special, the valuable comments and revisions of

several experts on the manuscript and appreciate the corrections by Shaun Pennycook,

Nomenclatural Editor, and suggestions by Lorelei L. Norvell, Editor-in-Chief. This work

was supported by Protax (Program of Capacitation in Taxonomy), Conselho Nacional

de Desenvolvimento Cientifico e Tecnolégico (CNPq), which provided a research grant

to Leonor C. Maia and scholarships to Danielle K. Alves da Silva and Daniele M.A.

de Assis; Programa de Biodiversidade do Semi-arido (PPBio); FACEPE (Fundacao de

Amparo a Ciéncia e Tecnologia do Estado de Pernambuco) which provided financial

support to G.A. Silva; Universidade Federal de Pernambuco (UFPE), which provided a

grant to F. Oehl as ‘visiting professor: The authors would like to thank the Millennium

Inorganic Chemicals Mining, a Cristal Company, for logistical support.

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with a unique spore wall structure. Mycotaxon 109: 483-491.

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MYCOTAXON

http://dx.doi.org/10.5248/119.133

Volume 119, pp. 133-136 January-March 2012

Radulomycetopsis (Agaricomycetes),

a new corticioid genus from India

G.S. DHINGRA*, PRIYANKA & JASPREET KAUR

Department of Botany, Punjabi University, Patiala 147 002, India

*CORRESPONDENCE TO: dhingragurpaul@gmail.com

ABSTRACT — A new corticioid genus, Radulomycetopsis, is described for the new species,

R. cystidiata, from West Kameng in Arunachal Pradesh.

Key worps — Eastern Himalaya, Bomdila, angiosperm host

While conducting a fungal foray in Bomdila in West Kameng of Arunachal

Pradesh, India, Dhingra made a collection on decaying angiospermous

branches. The macroscopic and microscopic characters were compared with

similar genera in Corticiaceae (Rattan 1977, Thind & Dhingra 1985, Eriksson et

al. 1981, Hjortstam et al. 1987) but could not be assigned to any already known,

hence the description of a new genus. Morphological traits show similarities

with Radulomyces.

Radulomycetopsis Dhingra, Priyanka & J. Kaur, gen. nov.

MycoBank MB 560517

Differs from Radulomyces in basidiocarps that are dark-colored, lack clamp connections,

and possess cystidia.

TyPE SPECIES: Radulomycetopsis cystidiata Dhingra, Priyanka & J. Kaur

ErymMoLocy: The name of the genus is based on the resemblance with genus

Radulomyces.

Basidiocarp resupinate, adnate, effused, membranous-ceraceous; hymenial

surface smooth to slightly tuberculate, orange to brownish orange to reddish

brown; margins fibrillose, concolorous but paler; hyphal system monomitic;

generative hyphae branched at wide angles, without clamps, basal hyphae

covered with a thick sheath of brownish red matter, which dissolves in 3%

KOH; cystidia projecting, thin- to slightly thick-walled; simple to somewhat

branched hyphoid structures present in the hymenium; basidia clavate to

134 ... Dhingra, Priyanka & Kaur

WRAPS ) I

{ —S

AWG,

PEE 4

Fics 1-6. Radulomycetopsis cystidiata: microscopic structures.

1. Basidiospores; 2. basidia; 3. hyphoid structures in the hymenium; 4. generative hyphae;

5. cystidium; 6. vertical section through basidiocarp.

subclavate, 4-sterigmate, without a basal clamp; basidiospores broadly ellipsoid

to subglobose, smooth, thin- to slightly thick-walled, inamyloid, acyanophilous;

both basidia and basidiospores rich in oil drops.

REMARKS— Radulomycetopsis resembles Radulomyces in producing simple

to somewhat branched hyphoid structures in the hymenium, similar basidia,

Radulomycetopsis cystidiata gen. & sp. nov. (India) ... 135

Fic. 7. Radulomycetopsis cystidiata: basidiocarp showing hymenial surface.

and broadly ellipsoid to subglobose basidiospores that are thin- to slightly

thick-walled, non-amyloid, acyanophilous, and with oil rich protoplasm.

Radulomycetopsis differs in having dark colored fruitbodies, hyphae without

clamps, and presence of cystidia, a combination of features that supports an

independent genus. A sample has been studied by Prof. Kurt Hjortstam, who

supports the concept of a new genus.

Radulomycetopsis cystidiata Dhingra, Priyanka & J. Kaur, sp. nov.

MycoBank MB 560518 FIGS 1-7

Differs from Radulomyces confluens by the absence of clamp connections and the

presence of cystidia.

Type: India, Arunachal Pradesh: West Kameng, Bombila, on decaying angiospermous

branches, 29 August 1981, Dhingra 19767 (PAN, holotype).

Erymo.ocy: The epithet refers to the presence of cystidia.

Basidiocarp resupinate, adnate, effused, up to 430 um thick in section,

membranous-ceraceous; hymenial surface smooth to slightly tuberculate,

orange to brownish orange to reddish brown when fresh, becoming orange

gray to grayish orange to brownish orange on drying; margins thinning,

irregular in outline, fibrillose. Hyphal system monomitic; generative hyphae

up to 3 um wide, thin-walled, branched at wide angles, septate, without clamps;

basal zone composed of sparsely branched hyphae, running almost parallel to

the substrate, covered with some brownish-red matter which dissolves in 3%

136 ... Dhingra, Priyanka & Kaur

KOH, followed by a zone of comparatively more richly branched, loosely and

irregularly interwoven hyphae which merge into the hymenial zone where the

hyphae become vertical and much branched forming a dense texture. Cystidia

70.0-85.0 x 9.0-11.0 um, cylindrical to sub cylindrical, thin- to somewhat

thick-walled, projecting up to 50 um out of the hymenium. Some simple to

slightly branched hyphoid structures, are also observed in the hymenium. These

are thin-walled, with oily contents, and look different from dendrohyphidia.

Basidia 25.0-37.0 x 7.0-8.5 um, clavate to subclavate, often constricted with

oily contents, 4-sterigmate, with no basal clamp; sterigmata up to 7.0 um long.

Basidiospores 6.0-9.0 x 5-7.25 um, broadly ellipsoid to subglobose, thin- to

slightly thick-walled, inamyloid, acyanophilous, uniguttulate or with many oil

drops.

REMARKS—Radulomycetopsis cystidiata is similar to Radulomyces confluens,

which is easily distinguished by the presence of clamp connections and absence

of cystidia.

Acknowledgements

The authors thank Prof. Nils Hallenberg (Gothenburg, Sweden) and Prof. B.M.

Sharma (Plant Pathology, COA, CSKHPAU, Palampur, H.P., India) for peer review and

Prof. Joost Stalpers (CBS Fungal Biodiversity Centre, Netherlands) for suggesting correct

generic and specific names in Latin. Head, Department of Botany, Punjabi University,

Patiala, is thanked for providing research facilities.

Literature cited

Eriksson J, Hjortstam K, Ryvarden L. 1981. The Corticiaceae of North Europe - VI. Fungiflora,

Oslo. pp. 1051-1276.

Hjortstam K, Larsson KH, Ryvarden L. 1987. The Corticiaceae of North Europe - I. Fungiflora,

Oslo. pp. 1-59.

Rattan SS. 1977. The resupinate Aphyllophorales of the North Western Himalayas. Bibliotheca

Mycologica 60: 1-427.

Thind KS, Dhingra GS. 1985. Thelephoroid fungi of the Eastern Himalayas-I. Res. Bull. (Sci.) Pan.

Uni. 36: 165-174.

MY COTAXON

http://dx.doi.org/10.5248/119.137

Volume 119, pp. 137-140 January-March 2012

First record of Dendrothele mangiferae (Agaricales, Basidiomycota)

from the Neotropics

SERGIO P. GORJON

Centro de Investigacion y Extension Forestal Andino Patagénico (CIEFAP),

Area de Proteccién. CC 14, 9200 Esquel, Chubut, Argentina

CORRESPONDENCE TO: spgorjon@hotmail.com

Asstract — Dendrothele mangiferae, previously known only from the Mascarene Islands,

is reported for the first time from Central America. It has been found in Costa Rica growing

on bark of living Terminalia catappa. A full description, line drawing, and colour photos are

provided.

KEY worps — corticioid fungi, Manuel Antonio, ornamented basidiospores

Introduction

A conspicuous corticioid species, Dendrothele mangiferae, has recently

been collected from Costa Rica on bark of living Terminalia catappa L.

(Combretaceae), a large tree naturally widespread in subtropical and tropical

zones of Indian and Pacific Oceans and planted extensively throughout the

tropics. It constitutes the first record from outside the Mascarene Islands of

Réunion and Mauritius (Boidin et al. 1996). Specimens are fully described and

illustrated. Additional comments on the generic placement and related species

are provided.

Materials & methods

For light microscopic studies, samples were mounted in 3% potassium hydroxide

(KOH), Melzer’s reagent (IKI), and 0.1% cotton blue in 60% lactic acid to determine

cyanophily of basidiospore walls. Line drawings were made with a camera lucida

attachment. Specimens are deposited in INB and SALA.

Taxonomy

Dendrothele mangiferae Boidin & Duhem, Bull. Soc. Mycol. France 112(2): 106,

1996. PLATES 1-2

138 ... Gorjén

PiatE 1. Dendrothele mangiferae (S.P. Gorjon 3383).

Hymenial elements: a) basidiospores; b) basidia; c) hyphae; d) dendrohyphidia; e) host cells.

Dendrothele mangiferae, new to Central America ... 139

PLaTE 2. Dendrothele mangiferae (S.P.Gorjon 3383). Basidiome. Scale Bar = 2 cm.

BASIDIOMATA resupinate, orbicular at first, then effuse, adnate, white,

hymenial surface smooth, cracked when mature, margin abrupt. HyPHAL

SYSTEM monomitic, hyphae clamped, 2-4 um wide, thin-walled, obscured by

the abundant crystalline encrustation. DENDROHYPHIDIA filamentous, sinuous,

with few branches, clamped. Basip1a suburniform at first, then subcylindrical,

cyanophilous, 100-150 x 15-25 um, basally clamped, with 4 long sterigmata

about 15-25 um. Cystip1A absent. BAstp1osporEs subglobose, (22-)24-27 x

22-25 um, when immature and still attached to the sterigmata broadly ellipsoid,

seemingly smooth but minutely spinulose in Melzer’s reagent and cotton blue,

walls hyaline, slightly thickened, with a conspicuous blunt apiculus, faintly

cyanophilous, inamyloid, nondextrinoid.

HABITAT AND DISTRIBUTION — Known from the Mascarene Islands growing

on trunks of Mangifera indica L. (Anacardiaceae) and undetermined trees; and

from Costa Rica on bark of living Terminalia catappa on the Pacific coastal

shore (PLATE 3).

SPECIMENS EXAMINED — COSTA RICA. MANUEL ANTONIO, Espadilla beach, 9°23'35"N

84°09'12"W, 2-5 m a.s.l., 23.VI.2011, on bark of living Terminalia catappa, leg. S.P.

Gorjon, coll. 3380, 3381, 3382, 3383, 3384.

ComMENts — Dendrothele mangiferae is easily recognized by the large globose

and minutely ornamented basidiospores. Within Dendrothele Hohn. & Litsch.,

140 ... Gorjén

PiaTE 3. Habitat of Dendrothele mangiferae with Terminalia catappa trees

on the Pacific coast of Costa Rica.

there are two additional species with spinulose basidiospores: Dendrothele

candida (Schwein.) PA. Lemke, with globose basidiospores (13-)15-20 x

(11-)13-16(-20) um and clampless hyphae; and Dendrothele nivosa (Berk.

& M.A. Curtis ex Hohn. & Litsch.) PA. Lemke, with subglobose to ovoid

basidiospores 16-24 x 14-17 um and clamped hyphae. Phylogenetic studies

show that Dendrothele is a polyphyletic genus, and that D. nivosa seems not

to be closely related to other Dendrothele species, including the generic type,

D. papillosa Hohn. & Litsch. [= D. griseocana (Bres.) Bourdot & Galzin] (Karen

K. Nakasone, pers. comm.). Further studies are needed within Dendrothele

species with ornamented basidiospores to elucidate their phylogenetic

relationships.

Acknowledgments

Juliano M. Baltazar (Brazil) and Cristiano Losi (Italy) acted as presubmission

reviewers. The Protection area of CIEFAP is acknowledged for technical support.

Literature cited

Boidin J, Lanquetin P, Duhem B. 1996. Contribution a la connaissance du genre Dendrothele

(Basidiomycotina, Aphyllophorales). Bull. Soc. Mycol. France 112(2): 87-126.

MY COTAXON

http://dx.doi.org/10.5248/119.141

Volume 119, pp. 141-148 January-March 2012

Two new fungi from Mexico: Anaseptoidium gen. nov.

and Cylindrosympodium sosae sp. nov.

RAFAEL FE. CASTANEDA-RuIZ’, GABRIELA HEREDIA”, ROSA M. ARIAS-MOTA?,

MARC STADLER? , MASATOSHI SAIKAWA‘* & ERIC H.C. MCKENZIE}

"Instituto de Investigaciones Fundamentales en Agricultura Tropical ‘Alejandro de Humboldt’

(INIFAT), Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200

?Instituto de Ecologia, A.C., Congregacion El Haya, Xalapa, Veracruz 91070, México

°InterMed Discovery GmbH, Otto-Hahn-str. 15, D-44227 Dortmund, Germany

‘Department of Biology, Tokyo Gakugei University,

Nukuikita-machi, Koganei-shi, Tokyo 184-8501, Japan

°Landcare Research, Private Bag 92170, Auckland, New Zealand

*CORRESPONDENCE TO: Marc.Stadler@t-online.de

ABSTRACT — Two new anamorphic fungi are described and illustrated from a cloud forest in

Mexico. Anaseptoidium mycophilum gen. & sp. nov. is distinguished by monoblastic, integrated,

mostly determinate conidiogenous cells and solitary, 2—3-septate, oblong to cylindrical, pale

brown conidia that usually remain attached after maturity. Cylindrosympodium sosae sp. nov.

is characterized by cylindrical, mostly 4—5-septate, smooth, hyaline to subhyaline, sometimes

guttulate conidia.

Key worps — mycophilous fungi, systematics

During an investigation of microfungi on dead plant material in a cloud forest

located in the private protected area “Agiiita Fria’, Veracruz, Mexico, two

undescribed anamorphic fungi were collected. Individual samples of plant

material were placed in paper and plastic bags taken to the laboratory and

treated according to Castafieda (2005). Mounts were prepared in polyvinyl

alcohol-glycerol (8 gin 100 ml of water, plus 5 ml of glycerol) and measurements

made at a magnification of x1000. Micrographs were obtained with a Nikon

Eclipse E600 light microscope (Japan). One new genus and two new species are

described below.

Taxonomy

Anaseptoidium RF. Castafieda, Heredia & R.M. Arias, anam. gen. nov.

MycoBank MB561820

142 ... Castafieda-Ruiz & al.

Coloniae in substrato naturali effusae, reptantes, funiculosae, brunneae. Stomatopodia

absentia. Conidiophora macronematosa interdum in cellulla conidiogenae reducta. Cellulae

conidiogenae monoblasticae, determinatae. Conidia solitaria, acrogena, cylindrica usque

ad oblonga, fimbriate, septata, pallide brunnea ad brunnea.

TYPE SPECIES: Anaseptoidium mycophilum R.F. Castafieda et al.

EryMoLocy: Greek, Ana-, meaning upwards, back, and again; Latin, septoidium,

referring to an anamorphic genus Septoidium.

CoLoniges on the natural substrate effuse, creeping, funiculose, brown.

Mycelium superficial and immersed. STOMATOPODIA absent. CONIDIOPHORES

macronematous, mononematous, erect, sometimes reduced to conidiogenous

cells. CONIDIOGENOUS CELLS monoblastic, integrated, determinate. Conidial

secession schizolytic. Conrp1A solitary, acrogenous, cylindrical to oblong,

fimbriate, septate, smooth or verruculose, pale brown to brown. Teleomorph

unknown.

Anaseptoidium mycophilum R.F. Castafieda, Heredia & R.M. Arias,

anam. sp. nov. Fies. 1, 2

MycoBank MB561823

Conidiophora macronematosa, 1-2-septata, 7.6-17.3 x 4.5-5.5 um, interdum in cellula

conidiogena reducta. Cellulae conidiogenae monoblasticae, 4-17 x 2-6.5 um. Conidia

acrogena, oblonga vel cylindrica, (1-)2(-3)-septata, plerumque eccentrica fimbriata ad

basim, 12-17 x 4-6 ym.

Type: Mexico, Veracruz: Agiita Fria, 19°31'N 96°59’W, on synnemata of Phaeoisaria

clavulata (Grove) E.W. Mason & S. Hughes, 8 February 2010, G. Heredia & V. Sosa

(Holotype: XAL CB1696).

EryMo_ocy: Greek, myco-, meaning fungus, fungal, Greek, -philum, meaning loving.

CoLoniges on the natural substrate, effuse, creeping, funiculose, brown,

mycophilic. Mycelium mostly superficial. Hyphae septate, branched, 2-4

um diam., smooth, pale brown, ascending on the synnematal conidiomata

of the host. SromatTopopiA absent. CONIDIOPHORES macronematous,

mononematous, erect or prostrate, straight, simple, 1-2-septate, smooth,

7.6-17.3 x 4.5-5.5 um, very pale brown, sometimes reduced to conidiogenous

cells. CONIDIOGENOUS CELLS monoblastic, terminal, determinate, integrated,

pale brown, 4-17 x 2-6.5 um. Conidial secession schizolytic. Conip1a solitary,

acrogenous, oblong to cylindrical, (1-)2(-3)-septate, mostly eccentric and

fimbriate at the base, rounded at the apex, smooth, 12-17 x 4-6 um, very pale

brown, dry, usually remaining attached to the conidiogenous loci after maturity.

Teleomorph unknown.

Note: Anaseptoidium resembles Septoidium G. Arnaud, which differs in

monoblastic, percurrently proliferating, annellate conidiogenous cells and

stomatopodia formed regularly by the assimilative hyphae on the substrate.

Most species are plant pathogenic (Baker 1955, Ellis 1971, 1976). Esquivel

Anaseptoidium gen. nov. & Cylindrosympodium sosae sp. nov. (Mexico) ... 143

Fic. 1. Anaseptoidium mycophilum (ex holotype XAL CB1696, on synnemata

of Phaeoisaria clavulata). a-e. Conidiophores, conidiogenous cell and

conidia. a—b. Scale bars = 20 um, c-e scale bars = 50 um.

(2011), who illustrated several conidiogenous events observed in the Septoidium

anamorph of Parodiopsis hurae (G. Arnaud) R.E.D. Baker & W.T. Dale,

confirmed the enteroblastic percurrent proliferations of the conidiogenous

cells in Septoidium.

In Anaseptoidium, the secession of each conidium is schizolytic, but the outer

and inner wall layers do not separate simultaneously as in most related fungi.

The outer wall layer(s) breaks first, and many conidia are observed attached

144 ... Castafieda-Ruiz & al.

Fic. 2. Anaseptoidium mycophilum (ex holotype XAL CB1696). a-i. Conidio-

genous cells and conidia. Scale bars = 5 um, j-1. Conidiophores, conidiogenous

cells and conidia. Scale bars = 10 um.

only by the inner wall layer(s) as can be confirmed by the pale or colorless

pigmentation of the conidiogenous cell apex just below each attached conidium

(Fic. 2 a-k). The inner wall layer(s) may separate independently of the outer

wall separation(s). As a result, conidia are seen somewhat separated (1—2 um)

from the outer wall layers while still attached on each conidiogenous locus.

Anaseptoidium gen. nov. & Cylindrosympodium sosae sp. nov. (Mexico) ... 145

Fic. 3. Cylindrosympodium sosae (ex holotype XAL CB1699). a-f. Conidia,

g. Conidiophore, conidiogenous cells and conidium. h-l. Conidiogenous

cells and conidia. Scale bars = 10 um.

Cylindrosympodium sosae R.F. Castafieda, Heredia & R.M. Arias,

anam. sp. nov. Figs. 3, 4

MycoBank MB561822

Ad Cylindrosympodii africani differt in conidiophora macronematosa, 6-15-septata,

71-106 x 5-10 um, leprosa vel verruculosa ad basim et conidia 4-5-septata, 25.5-28.5

x 4-4.5 um.

146 ... Castafieda-Ruiz & al.

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10pm

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oak

om be

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Fic. 4. Cylindrosympodium sosae (ex holotype XAL CB1699).

Conidiophores, conidiogenous cells and conidia.

Type: Mexico. Veracruz: Agiiita Fria, 19°31'N 96°59’ W, on decaying leaf of Liquidambar

styraciflua L. (Hamamelidaceae),8 February 2010, V. Sosa (Holotype: XAL CB1699).

ETYMOLOGY: Latin, sosae, in honor to Dr. Vinicio Sosa Fernandez (INECOL, Mexico)

who collected and kindly provided the specimen.

Anaseptoidium gen. nov. & Cylindrosympodium sosae sp. nov. (Mexico) ... 147

CoLonigs on the natural substrate effuse, pilose, yellowish brown or reddish

brown. Mycelium mostly immersed. Hyphae septate, branched, smooth or

verruculose, yellowish brown or brown, 1.5-3 um diam. CONIDIOPHORES

macronematous, mononematous, erect, straight or flexuous, simple, 6-15-

septate, verruculose to leprous near the base and smooth towards the apex,

71-106 x 5-10 um, yellowish brown or reddish brown below, hyaline or

subhyaline towards the apex. CONIDIOGENOUS CELLS polyblastic, integrated,

intercalary and terminal, subhyaline or hyaline, 8-15 x 3-4.5 um, geniculate,

indeterminate, with several holoblastic sympodial proliferations and flattened

conidiogenous loci. Conidial secession schizolytic. Conrp1a solitary,

acropleurogenous, cylindrical, truncate at the base, 4—5-septate, smooth, hyaline

or subhyaline, 25.5-28.5 x 4-4.5 um, sometimes guttulate, dry. Teleomorph

unknown.

Note: Cylindrosympodium is distinguished by “flat-topped denticles” on

conidiogenous cells that form solitary, cylindrical, septate, hyaline conidia

(Castafieda-Ruiz & Kendrick 1990, 1991, Paulus et al. 2003, Marvanova &

Laichmanova 2007, Crous et al. 2007). Cylindrosympodium sosae strongly

resembles C. africanum (Morgan-Jones et al.) Marvanova (Marvanova &

Laichmanova 2007), which is easily differentiated by its brown or pale brown

smooth conidiophores and 3-4-septate 31-33 x 2 um conidia (Morgan-Jones

et al. 1983).

Acknowledgments

The authors express their sincere gratitude to Dr. Lori Carris and Dr. De-Wei Li for

their critical review of the manuscript. The authors are deeply indebted to the IE004

project from CONABIO of Mexico for financial support and INECOL A. C., Xalapa,

Ver., Mexico and Cuban Ministry of Agriculture for facilities. The authors thank Drs.

Andrea Romero, Xiu Guo Zhang, D.J. Bhat, Sanjay K. Singh and Gregorio Delgado for

their generous and valued assistance with literature not otherwise available. We thank

Mirtha Caraballo for technical assistance. We also acknowledge the facilities provided

by Dr. P.M. Kirk and Drs. V. Robert and G. Stegehuis through the IndexFungorum

and Mycobank websites. Dr. Lorelei L. Norvell’s editorial and Dr. Shaun Pennycook’s

nomenclature reviews are greatly appreciated.

Literature cited

Baker RDE. 1955. Species of the genus Parodiopsis Maubl. found in Trinidad. Mycological Papers

58: 1-16.

Castafieda-Ruiz RF. 2005. Metodologia en el estudio de los hongos anamorfos. 182-183, in: Anais

do V Congresso Latino Americano de Micologia. Brasilia.

Castafieda-Ruiz RE, Kendrick B. 1990. Conidial fungi from Cuba: I. University of Waterloo Biology

Series. 32:53: p.

Castafeda-Ruiz RE, Kendrick B. 1991. Ninety-nine conidial fungi from Cuba and three from

Canada. University of Waterloo Biology Series 35: 132 p.

148 ... Castafieda-Ruiz & al.

Crous PW, Braun U, Schubert K, Groenewald JZ 2007. Delimiting Cladosporium from

morphologically similar genera. Stud. in Mycol. 58: 33-56.

http://dx.doi.org/10.3114/sim.2007.58.02

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Surrey, Kew.

Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Surrey,

Kew.

Esquivel REA. 2011. Observaciones sobre la morfologia y conidiogénesis de Septoidium st.

Parodiopsis hurae (Arn) Baker & Dale. (Hifomiceto, Dematiaceo). http://agrociencia-panama.

blogspot.com/2011/03/observaciones-sobre-la-morfologia-y.html.

Marvanova L, Laichmanova M. 2007. Subulispora biappendiculata, anamorph sp. nov. from Borneo

(Malaysia) and a review of the genus. Fungal Diversity 26: 241-256.

Morgan-Jones G, Sinclair RC, Eicker A. 1983. Notes on hyphomycetes. XLIV. New and rare

dematiaceous species from the Transvaal. Mycotaxon 17: 301-316.

Paulus BC, Gadek PA, Hyde, KD. 2003. Cylindrosympodium cryptocaryae sp. nov. (anamorphic

fungi), with keys to the described species and to similar genera. Australian Systematic Botany

16: 577-580. http://dx.doi.org/10.1071/SB03005

MY COTAXON

http://dx.doi.org/10.5248/119.149

Volume 119, pp. 149-156 January-March 2012

Candelariella, Ochrolechia, Physcia, and Xanthoria

species new to Turkey

KENAN YAZICI’’, ANDRE APTROOT? & ALI ASLAN?

‘Biology Department, Science Faculty, Karadeniz Technical University, 61080, Trabzon, Turkey

?ABL Herbarium G.v.d. Veenstraat 107, NL-3762 XK Soest, The Netherlands

*Biology Department, Kazim Karabekir Education Faculty, Atatiirk University, Erzurum, Turkey

*CORRESPONDENCE TO kcagri_1997@yahoo.com

ABsTRACT — Candelariella kuusamoensis, Ochrolechia frigida, Physcia phaea, and Xanthoria

papillifera were determined as new to Turkey and the Middle East following a recent

lichenological foray in Igdir and Van regions (Turkey). Geographic distribution, substrate,

chemistry, and comparisons with morphologically similar taxa are presented.

Key worps — Ascomycetes, biodiversity, lichen

Introduction

Although many lichen taxa have recently been recorded for Turkey (Candan

et al. 2010, Candan & Halici 2009, Halici et al. 2010a,b, Kinalioglu 2009,

Kinalioglu & Aptroot 2010, Kinalioglu 2010a,b,c, Karagéz et al. 2011, Perez-

Ortega et al. 2010, Yazici & Aslan 2009, 2010a,b), the lichen biota of many

regions are still unknown. No lichenized fungi have thus far been reported for

the Igdir region, although some surveys of lichens have been conducted in the

neighbouring Ardahan region (Osyczka et al. 2011, Yazici et al. 2010a,b) and

81 taxa were reported from the Van province (Aslan & Oztiirk 1995, Szatala

1941, 1960, Senkardesler & L6k6s 2010). Ten Candelariella, fifteen Ochrolechia,

sixteen Physcia, and twelve Xanthoria taxa have previously been reported from

Turkey.

The climate of the Igdir region is characterized by hot dry summers and

cold snowy winters. The mean annual temperature is 11.6°C, average humidity

is 63%, and the mean annual rainfall is about 257.6 mm. Summers are

characterized by moderate precipitation and the winters are generally marked

with high precipitation (Akman 1999).

The Igdir region has few forested areas and is dominated by steppe. Mountain

Zordag (Igdir: Center), which is exposed to high light conditions, is a windswept

150 ... Yazici, Aptroot & Aslan

area with gently sloping terrain dominated by grass, rocks, and streams with

scattered coniferous and deciduous trees. Prunus, Pyrus, Populus, Salix, and

Elaeagnus trees are seen occasionally along the stream between Tasképrii and

Karaca villages (Tuzluca) (Baytop & Denizci 1963). There are no trees but many

calcareous and siliceous rocks in the Tasucan and Bahcecik villages, an exposed

windswept area.

The Van region has a continental climate, characterized by long cold snowy

winters and hot dry summers. The mean annual temperature is 9°C, average

humidity is 51.10% , and the mean annual rainfall is about 380 mm, with a

precipitation range of 370-570 mm (Akman 1999). The collection sites are

exposed to high light conditions and windswept with gently sloping terrain

comprising grass and volcanic calcareous rocks. A river valley with trees was

sampled only in Saray town (Van). In the other regions of the Van district there

are no deciduous or coniferous trees (Baytop & Denizci 1963).

Material & methods

During a recent lichenological research foray in Igdir and Van regions (Turkey),

lichen samples were collected from 07 December 2008 to 27 September 2009 and 13

June 2010 to 12 July 2010. Air-dried samples were examined with a Nikon SMZ1500

stereomicroscope and a Nikon Eclipse 80i light microscope. Identifications were

determined by consulting appropriate keys in the literature (Brodo et al. 2001, Dobson

2005, Karnefelt 1990, Kukwa 2009, Moberg 1977, Poelt 1974, Poelt & Vézda 1981,

Redchenko et al. 2010, Smith et al. 2009, Thomson 1997, Wetmore 2005, Wirth 1995).

Secondary metabolites were identified by the usual spot tests.

Vouchers are stored in the herbarium of the Biology Department, Science Faculty,

Karadeniz Technical University, Trabzon, Turkey (KTUB) and the Biology Department,

Kazim Karabekir Education Faculty, Atatiirk University, Erzurum, Turkey (ATA-

KKEF).

Species recorded

Candelariella kuusamoensis Rasanen, Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo,

12(1): 56.1939 Fig 1

Thallus muscicolous, thick, dense, granular-warty, yellow to orange-yellow,

to 3.2 cm in diam., almost cushion or aggregated break parts-like, to deep

areolate cracks, consisting of closely crowded large granules (0.1-0.7 mm),

upper surface uneven, yellow. Medulla green. Apothecia rare, to 0.25-1mm

diam., asci clavate, Candelaria-type, 12-32-spored, ascospores 9-15 x 4.5-5.5

um, non-septate, hyaline. Algae trebouxioid. Hypothecium hyaline; hymenium

hyaline, c. 65-70 um high. Paraphyses 2.5-3um thick. Cortex K -, C -, KC -,

P -. Medulla K-, C-, KC-, P -. Thallus contains pulvinic acid.

Candelariella kuusamoensis grows on moss, plant remains, soil, more rarely

on rocks and wood and wooden fences, and on the top of poles in alpine areas.

Four lichenized fungi new to Turkey... 151

-_* a é ah « :

Fic. 1. Candelariella kuusamoensis, habitus. Scale= 1mm

Previously known from Asia, Europe, and North America. New to Turkey and

the Middle East.

SPECIMEN EXAMINED: TURKEY. IGp1r: CENTER, Zorda& mountain, 39°45'49.05"N

43°53'20.13"E, on mosses, 2400m, 13.06.2010, (leg. K.Yazici,) KTUB-2023.

Ochrolechia frigida (Sw.) Lynge, Lich. Nov. Zemlya: 182.1928 Fig 2

Thallus crustose, muscicolous, to 1.2-2 cm diam., + orbicular, thin crust

or thick granules, effuse, + verrucose, uneven-granulate, occasionally with few

cylindrical-coralloid and branching spinulose extensions, whitish to yellow-

white, pinkish or gray, soralia rare. Apothecia rare, to 1 mm wide, sessile to

adnate; margin thin to slightly thickened, entire or wrinkled; disk flat, red-

brown, slightly roughened, epruinose. Hypothecium dull brown; epihymenium

brown; hymenium 130-180 um, hyaline; paraphyses slender; Asci 8-spored,

ascospores simple, ovoid, 26-43 x 15-26 um. Cortex K-, C+ red, KC+ red, P-;

medulla K-, C+ red, P-; disc K-, C+ pink to red, KC + red, P-. Thallus contains

gyrophoric acid, + variolaric acid.

Ochrolechia frigida is a bipolar lichen, occuring mainly on soil, mosses,

humus, plant remains, and at the base of plants, It is especially found in raised

bogs on mountain tops in arctic-boreal and alpine regions in the northern

152 ... Yazici, Aptroot & Aslan

er. Ts e . ‘

Te Sa

: ht ap

@ s

Fic. 2. Ochrolechia frigida, habitus. Scale= 1cm

hemisphere. Previously known from Antarctica, Asia, Australasia, Europe,

North America, and South America. New to Turkey and the Middle East.

SPECIMEN EXAMINED: TURKEY. IGpir: Tuz.uca, between Task6prii-Karaca villages,

39°52'07.40"N 43°28'57.94"E, 2134m, 14.6.2010, (leg. K.Yazici,) KTUB-2038.

REMARKS— Ochrolechia frigida differs from O. upsaliensis and O. alaskana in

its smaller ascospores (25-40 x 15-26 um) and apothecia (O. upsaliensis has

ascospores 55-75 x 26-38 um). Also, O. alaskana has variolaric acid, which

O. frigida lacks (Kukwa 2009). In addition, the thallus of O. frigida is delicate,

grey, brown-grey, or brown, occasionally with a yellow tinge and distinct,

branched spines, while the thallus in O. alaskana is thick and firm with a yellow

tinge, and spines are lacking or occur occasionally in some specimens.

Physcia phaea (Tuck.) J.W. Thomson, Beih. Nova Hedwigia 7: 54. 1963 Fic 3

Thallus + orbicular, or irregular, to 1.5-2.2 cm in diam., usually closely

appressed to the substratum, steel-gray to dark gray, or brownish gray, white

maculae present, rugose; pruina, soredia, isidia or lobules absent; lobes

radiating, flat to slightly convex, 1-1.5 mm wide, typically crowded and

overlapping, eciliate. Medulla white. Lower surface white to dark gray or brown,

with white to black rhizines. Lower surface pale tan to brownish, with abundant

Four lichenized fungi new to Turkey... 153

Fic. 3. Physcia phaea, habitus. Scale= 1mm

or sparse pale to brown rhizines, 0.3-1mm long. Apothecia numerous, 0.5-1.5

mm diam., margins lecanorine, crenulated when young, perfectly round and

somewhat raised; disc dark brown to black, rarely covered with a white pruina;

ascospores brown, l-septate, Physcia-type, 16-20 x 7-9 um. Cortex K + yellow,

C + slightly yellow, KC + slightly yellow, P -. Medulla K + yellow, C -, KC +

yellow, P -. Thallus contains atranorin and zeorin. Pycnidia numerous; conidia

immersed, subcylindrical, 4-6 x 1 um.

Physcia phaea occurs mainly on granitic (less commonly, calcareous) rocks,

or mosses (mostly in open areas), also on basaltic or gneissic and nutrient-

enriched rocks. Previously known from Asia, Australasia, Europe, and North

America. New to Turkey and the Middle East.

SPECIMENS EXAMINED: TURKEY. I6p1r: Tuziuca, Tasucan village, 39°51'26.61"N

43°35°41.06"E, on calcareous rock, 3168m, 02.07.2010, (leg. K.Yazici), KTUB-2039.

VAN: BASKALE, Baskale-Agagi1 Dikmen Yol Ayrimi, Camlik Képrii Mevkisi: 37°57'09"N

44°05'35"E, on calcareous rock, 1886 m, 29.04.2009, (leg. A.Aslan), ATA-KKEF-1951;

Gevas, Akdamar Adasi Karsisi, Akdamar Camping Yani, 38°18'32"N 43°02'24"E,

on calcareous rock, 1662 m, 05.09.2009, (leg. A.Aslan), ATA-KKEF-1955; CENTER,

Gedikbulak Kéyii, Tuzla Karsis, 38°51'13"N 43°26'32"E, on calcareous rock, 1723 m,

07.09.2009, (leg. A-Aslan), ATA-KKEF-1954, Van Kalesi, 38°30'14"N 43°19'56"E, on

calcareous rock, 1681 m, 07-11.12.2008, (leg. A.Aslan), ATA-KKEF-1953, Erek Dagi

Dogu Yamaci: 38°29'59”N 43°29'22"E, on calcareous rock, 1976 m, 01-05.10.2008, (leg.

154 ... Yazici, Aptroot & Aslan

A.Aslan), ATA-KKEF-1950; CaLprran, Caldiran Crkisi, 39°08'07"N 43°55'45"E, on

calcareous rock, 2053 m, 28.04.2009, (leg. A.Aslan), ATA-KKEF-1952.

REMARKS— Physcia phaea resembles P aipolia and P. cascadensis, but is

distinguished from P. aipolia by occasionally growing on a bark and having

apothecia that lack soralia and the heavy pruina characteristic of P. aipolia.

Physcia cascadensis, a rare and rock-dwelling rosette lichen, like P phaea lacks

soredia and has a K + yellow medulla but differs in its very pale gray cortex,

absence of maculae, and pruinose apothecia.

Be -

Fic. 4. Xanthoria papillifera, habitus. Scale= 1mm

Xanthoria papillifera (Vain.) Poelt, Mitt. Bot. Staatss. Miinchen 2: 29.1954 = Fie 4

Thallus crustose, small, to 5-15 mm diam., forming + appressed, + orbicular,

rosettes, reddish-orange, orange-yellowish; lobes well-developed, inflated

to flattened, or pulvinate, 0.4-0.6 mm wide, 1.5-1.8 mm long, separate or +

overlapped high-arched, , curved, + deeply lobed, mostly clearly separated.

Soredia scattered and isidia-like; isidia dense mostly central regions of the

thallus, coralloid or papillate, finger-like, pruinose, uniformly thick. Apothecia

rare, lecanorine 0.2-0.5 mm diam. Asci 8-spored, ascospore 1-septate,

colourless. Cortex on upper and lower thallus with hyphae + perpendicularly

oriented to the upper surface, attached by rhizines or adhering discs. Cortex

K + purple, C + orange turning orange-brown, K + orange-brown, P -. Medulla

K -, C -, KC -, P-. Apothecia K + purple. Thallus contains anthraquinones.

Four lichenized fungi new to Turkey... 155

Xanthoria papillifera grows on steep surfaces of calcareous and siliceous

rocks, boulders, and mossy soil, often in montane areas. Previously known

from Africa, Asia, Australasia, Europe, and North America. New to Turkey.

SPECIMENS EXAMINED: TURKEY. IGp1r: CENTER, Zorda% mountain, 39°45'49.05"N

43°53'20.13"E, on calcareous rock, 2400m, 13.06.2010, (leg. K.Yazici) KTUB-2022;

Tuztuca, Bah¢ecik village, on roadside, 40°04'05.63"N 43°25'35.47"E, on calcareous

rock, 1754m, 12.07.2010, (leg. K.Yazici), KTUB-2021. VAN: Saray, Saray-Kapikoy Yolu

Kaz Gélii Civari, 38°32'06"N 44°15'21"E, on calcareous rock, 2180 m, 27.09.2009, (leg.

A.Aslan), ATA-KKEF-1957; CALDIRAN, Tendiirek Dai, Uzunyol Kéyii, 39°19'08"N

44°01'56"E, on calcareous rock, 2351 m, 27.04.2009, (leg. A.Aslan), ATA-KKEF-1956.

REMARKS— Xanthoria papillifera is similar to X. sorediata, whichis distinguished

by rounded to flattened lobes and sorediate isidia. This species has also been

called Xanthoria domogledensis Vézda.

Acknowledgements

We are grateful to Dr. Orvo Vitikainen and Dr. Michele D. Piercey-Normore for

revisions and helpful comments on an earlier draft of this manuscript. This study was

supported by TUBITAK (projects 107T035 and 108T566).

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MY COTAXON

http://dx.doi.org/10.5248/119.157

Volume 119, pp. 157-166 January-March 2012

A new species and new combinations and records of

Hypotrachyna and Remototrachyna from Bolivia

ADAM FLAKUS’', PAMELA RODRIGUEZ SAAVEDRA”? & MARTIN KUKWA‘4

‘Laboratory of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences,

Lubicz 46, PL-31-512 Krakow, Poland

*Department of Botany and Molecular Evolution, Senckenberg Forschungsinstitut und

Naturmuseum, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany

*Herbario Nacional de Bolivia, Instituto de Ecologia, Universidad Mayor de San Andrés,

Calle 27, Cota Cota, Casilla 10077, La Paz, Bolivia

‘Department of Plant Taxonomy and Nature Conservation, University of Gdansk,

Al. Legionéw 9, PL-80-441 Gdansk, Poland

CORRESPONDENCE TO: 'a.flakus@botany. pl,

*pamela.rodriguez@senckenberg.de & sdokmak@ug.edu.pl

ABSTRACT — Remototrachyna sipmaniana is described as new to science, and three new

combinations, R. aguirrei, R. consimilis, and R. singularis, are proposed. Ten Hypotrachyna

and two Remototrachyna species are reported as new to Bolivia, including the southernmost

localities of H. halei and H. partita, the first record of H. primitiva from the southern

hemisphere, and the second locality for H. neoscytodes.

Key worps — foliose lichens, Neotropics, Parmeliaceae, South America

Introduction

Parmeliaceae Zenker (Lecanorales, Lecanoromycetes) is a large family of

Ascomycota consisting of foliose, fruticose or rarely crustose lichens, but also

lichenicolous non-lichenized taxa (e.g. Elix 1993, Lumbsch & Huhndorf 2007,

PersSoh & Rambold 2002). It is believed to be one of the richest in species within

the phylum and comprises about 1500 species (Blanco et al. 2006). This group

has been extensively investigated by several researchers for more than 50 years,

so the worldwide distribution of many taxa is well known. However, some areas,

especially in the tropics, need further study; one of these is Bolivia. Data on

Parmeliaceae from this country are not numerous and are mostly included in

monographic treatments (e.g. Hale 1965, 1975, 1976; Culberson & Culberson

1981, Nash et al. 1995, Sipman et al. 2009) or rarely floristic papers (e.g. Herzog

1922, Feuerer & Sipman 2005).

158 ... Flakus, Rodriguez Saavedra & Kukwa

During several lichenological expeditions to Bolivia we have collected more

than 20,000 specimens and, in addition, further samples have been obtained

on loan from several herbaria. Among the macrolichens, a large proportion

were representatives of Parmeliaceae. ‘This is the first paper dealing with

new distributional data for Parmeliaceae in Bolivia. Here we report a new

species and records for two morphologically similar, but phylogenetically

distant genera, Hypotrachyna (Vain.) Hale and Remototrachyna Divakar &

A. Crespo. Additionally, we propose transferring three additional species from

Hypotrachyna to Remototrachyna based on anatomical features.

Material & methods

The material studied is deposited in B, KRAM, LPB, UGDA and herb. Flakus.

Additional specimens from B, TUR-Vain and TNS were investigated for comparison.

Lichen substances were studied by thin-layer chromatography (TLC) using the methods

of Culberson & Kristinsson (1970) and Orange et al. (2001). Spot-test reactions with

C, K and P were applied to determine the location of secondary metabolites in some

specimens.

The genus Hypotrachyna in the Neotropics, including some species of Remototrachyna,

has recently been revised by Sipman et al. (2009), so the descriptions and secondary

chemistry of the species are not repeated here. The characteristics of many taxa can also

be found in Hale (1975).

Results

The genus Hypotrachyna

The diagnostic characters of the genus include the dichotomously or

subdichotomously branched lobes with truncate apices, bifusiform conidia,

sparsely to richly dichotomously branched rhizines, and an outer exciple

consisting of plectenchyma with thin cell walls (Crespo et al. 2010, Divakar et

al. 2010). In its current circumscription the genus is paraphyletic (see Blanco

et al. 2006, Divakar et al. 2006, Crespo et al. 2010) and needs further study

for a better understanding of its relationship to other morphologically similar

genera, e.g., Everniastrum Hale ex Sipman, Parmelinopsis Elix & Hale.

Recently the genus Remototrachyna was segregated from Hypotrachyna

basing mainly on the structure of the cupulate excipulum (plectenchymatous

with very thick-walled cells) and genetic differences (Divakar et al. 2010).

During our study we investigated several fertile specimens of Hypotrachyna

from the Neotropics, and in three species, H. aguirrei, H. consimilis and

H. singularis, the excipulum appeared typical for Remototrachyna; thus those

taxa are transferred to Remototrachyna (see below).

We also observed that the plectenchyma in the cupulate exciple of

Hypotrachyna can have thicker cell walls than presented by Divakar et al.

(2010), but cell lumina are more or less rounded or ellipsoid, never elongated

Remototrachyna sp. & combs. nov. (Bolivia) ... 159

and sigmoid in shape as in Remototrachyna (for iconography see Divakar et al.

2010).

Hypotrachyna aspera C.H. Ribeiro & Marcelli, in Marcelli & Ribeiro, Mitt. Inst.

Allg. Bot. Hamburg 30-32: 133. 2002.

This species has been recently described from Brazil (Marcelli & Ribeiro

2002, Sipman et al. 2009). Here it is reported as new to Bolivia, which represents

the first record outside of Brazil.

SPECIMENS EXAMINED — BOLIVIA. Dept. LA Paz. Prov. FRANZ TAMayo, sendero

Keara-Mojos, bajando por la senda de Tokuage a Fuertecillo, 2250 m, 14°36'S 68°56'W,

2001, Jimenez 5318 (LPB); DEPT. TARIJA. PROV. ANICETO ARCE, Filo de Sidras, 1064 m,

22°14'50"S 64°33'28" W, 2010, Flakus 18431 (LPB, herb. Flakus).

Hypotrachyna dentella (Hale & Kurok.) Hale, Smithsonian Contr. Bot. 25: 33. 1975.

In the Neotropics this species is widespread, but rare; it is also known from

the southeastern United States (Hale 1975, Sipman et al. 2009 and literature

cited therein). Here the first Bolivian records are presented.

SPECIMENS EXAMINED — BOLIVIA. Dept. Taria. Prov. ANICETO ARCE, Filo de

Sidras, near campamento de guardaparques, 2 hours of Tarija, 1065 m, 22°14'50"S

64°33'28" W, 2010, Flakus 18529, 18615 (KRAM, LPB, UGDA, herb. Flakus); Serrania de

Propiedad Arnold, 1309 m, 22°13'19"S 64°33'41" W, 2010, Flakus 18757 (KRAM, LPB).

Hypotrachyna halei Sipman, Elix & T.H. Nash, Fl. Neotrop. Monogr. 104: 76. 2009.

Sipman et al. (2009) reported H. halei from Costa Rica, Venezuela, Colombia,

and Ecuador; we now report it from Bolivia. The new localities below represent

the southernmost known for this species.

SPECIMENS EXAMINED — BOLIVIA. DEPT. COCHABAMBA. PROV. CHAPARE, Corani,

3261 m, 17°13'36"S 65°53'25"W, 2009, Flakus 12921 (KRAM, LPB); Dept. La Paz.

Prov. FRANZ Tamayo, Pifalito, cerro 29 km en linea recta al Este de Apolo, por el

camino a San Jose, 2000-2490 m, 14°29'42"S 68°15'26"W, 2002, Fuentes 5119 (LPB);

Prov. Nor Yuneas, 4 km del camino principal de Chuspipata hacia Coroico, 2750 m,

16°23'S 67°48'W, 1997, Bach et al. 542 (LPB); PN y ANMI Cotapata, 30 minutes of

Unduavi by Sillu Tincara pre-Columbian route, 3437 m, 16°17'38"S 67°53'33" W, 2009,

Flakus 16331 & Rodriguez (KRAM, LPB); 5 hours of Unduavi by Sillu Tincara pre-

Columbian route, 3429 m, 16°16'33"S 67°52'60"W, 2010, Flakus 16897, 16967, 16972,

16997 & Rodriguez (KRAM, LPB, UGDA, herb. Flakus).

Hypotrachyna lividescens (Kurok.) Hale, Phytologia 28: 341. 1974.

In the Neotropics this species has been reported from Colombia, Guatemala,

and Mexico (Sipman et al. 2009), and we now report it from Bolivia. The

species is also known from Africa, Australia, and Europe (see Sipman et al.

2009, and literature cited therein). The following record is the southernmost in

the Neotropics.

SPECIMENS EXAMINED — BOLIVIA. Dept. TaryaA. Prov. ANICETO ARCE, Papachacra,

Tucumano-Boliviano montane forest, 2195 m, 21°41'36"S 64°29'33"W, 2010, Flakus

19847, 19848 & Quisbert (KRAM, LPB, herb. Flakus).

160 ... Flakus, Rodriguez Saavedra & Kukwa

Hypotrachyna neoscytodes Elix, T.H. Nash & Sipman, in Sipman et al., Fl. Neotrop.

Monogr. 104: 106. 2009.

This species is new to Bolivia and previously known only from the type

locality in Colombia (Sipman et al. 2009).

SPECIMEN EXAMINED — BOLIVIA. Dept. La Paz. PRov. Nor YuNGAs, Carretera

Cotapata-Santa Barbara, Paramo yunguefio, 3573 m, 16°19'23"S 67°56'36.8"W, 2007,

Rodriguez 453 (LPB).

Hypotrachyna partita Hale, Smithsonian Contr. Bot. 25: 52. 1975.

This species was previously known from Colombia, Costa Rica, Mexico,

Panama, Venezuela, and Ecuador, usually from subpaéramo dwarf forests

(Sipman et al. 2009). The following Bolivian records extend its distributional

range southward.

SPECIMENS EXAMINED — BOLIVIA. Dept. La Paz. PRov. CAMACHO, Pacoamba

cerca Wila Kala, 4286 m, 15°24'40"S 69°04'24"W, 2010, Flakus 17696/3, 17699, 17755

& Rodriguez (KRAM, LPB, UGDA, herb. Flakus); Prov. Nor Yuncas, PN y ANMI

Cotapata, 5 hours of Unduavi by Sillu Tincara pre-Columbian route, 3429 m, 16°16'33"S

67°52'60"W, 2010, Flakus 16863, 16882 & Rodriguez (KRAM, LPB).

Hypotrachyna primitiva Hale & Lopez-Fig., Bryologist 81: 592. 1979 [“1978”].

H. primitiva was previously known from Colombia and Venezuela (Hale &

Lopez-Figueiras 1979, Sipman et al. 2009). The Bolivian localities presented

here are the southernmost known and the first records for the Southern

Hemisphere.

SPECIMENS EXAMINED — BOLIVIA. Dept. La Paz. Prov. FRANZ Tamayo, laguna

Tolca Cocha, al NE de Keara Nuevo, 3900 m, 14°41'13"S 69°05'18"W, 2006, Fuentes,

Mendoza, Lopez, Madariaga 9927 (LPB, herb. Flakus); Prov. Nor YuNGAS, Cotapata,

3050 m, 16°17'S 67°51'W, 1998, Franken, Meneses, Villavicencio 6 (LPB).

Hypotrachyna protocetrarica Elix, T.H. Nash & Sipman, in Sipman et al., FI.

Neotrop. Monogr. 104: 127. 2009.

This species, previously known from Ecuador and Peru (Sipman et al. 2009),

is reported as new to Bolivia.

SPECIMENS EXAMINED — BOLIVIA. Dept. La Paz. Prov. FRANZ Tamayo, laguna

Tolca Cocha, al NE de Keara Nuevo, 3900 m, 14°41'13”S 69°05'18"W, 2006, Fuentes,

Mendoza, Lopez, Madariaga 9909 (LPB, herb. Flakus); PRov. MuRILLo, Valle de Zongo,

laguna Viscachani, 3840 m, 1989, Coello 6 (LPB); ibid., 1988, Arrazola 29 (LPB).

Hypotrachyna pseudosinuosa (Asahina) Hale, Smithsonian Contr. Bot. 25:58.

192:

The widely distributed species in tropical and subtropical regions of the

world (Hale 1975, Sipman et al. 2009, and literature cited therein) is recorded

for the first time to Bolivia.

SPECIMENS EXAMINED — BOLIVIA. DEPT. COCHABAMBA. PROV. CHAPARE, near

Incachaca village, 2294 m, 17°14'13"S 65°49'02"W, 2006, Flakus 8222, 8256 (KRAM,

Remototrachyna sp. & combs. nov. (Bolivia) ... 161

LPB, herb. Flakus); Dept. La Paz. Prov. IrURRALDE, forest above Tumupasa village,

14°08'51"S 67°53'34"W, 350 m, 2008, Kukwa 7023a (UGDA, LPB).

Hypotrachyna subformosana Hale ex Elix, T.H. Nash & Sipman, in Sipman et al.,

Fl. Neotrop. Monogr. 104: 153. 2009.

This species, widespread but rare in the Neotropics, is reported here as new

to Bolivia. It is also known from Fiji (Hale 1975, Sipman et al. 2009).

SPECIMEN EXAMINED — BOLIVIA. Dept. La Paz. PRov. Nor YuNGAS, Coroico village,

1550 m, 16°11'10"S 67°43'16"W, 2010, Flakus 16414, 16449 & Rodriguez (KRAM, LPB,

herb. Flakus).

The genus Remototrachyna

This genus is morphologically very similar to Hypotrachyna but differs in

having a scleroplectenchymatous cupulate exciple, broad, subirregular lobes

with rounded apices, short, mostly dichotomously branched rhizines, a high

hymenium, and large ellipsoid ascospores; it is distantly related genetically to

Hypotrachyna (Divakar et al. 2010). In the course of this study we observed

larger variability in exciple structure of Hypotrachyna than reported by Divakar

et al. (2010). The cells can have thicker walls than previously reported, but in

Remototrachyna the hyphae are much more thick-walled, with very elongated,

thin, and linear to sigmoid cell lumina, which are usually almost vertically

orientated.

Basing on anatomical and morphological characters, four additional

species were recognized, one new and three transferred from Hypotrachyna

(see below). Probably more Hypotrachyna members should be transferred to

Remototrachyna, but further investigations of fertile material are needed.

According to Divakar et al. (2010), the ancestral range of Remototrachyna

was restricted to India. In the light of the new results the genus has a much wider

distribution and is more diverse in the Neotropics than previously supposed.

Possibly, the ancestral origins of Remototrachyna should be reevaluated.

Remototrachyna aguirrei (Sipman, Elix & T.H. Nash) Flakus, Kukwa & Sipman,

comb. nov.

MycoBank MB 561981

= Hypotrachyna aguirrei Sipman, Elix & T.H. Nash, Fl. Neotrop. Monogr. 104: 25. 2009.

This species has been recently described from Bolivia, Costa Rica, Colombia,

and Peru (Sipman et al. 2009).

Remototrachyna consimilis (Vain.) Flakus, Kukwa & Sipman, comb. noy.

MycoBank MB 561982

= Parmelia consimilis Vain., Acta Soc. Fauna Fl. Fenn. 7(1): 58. 1890.

= Hypotrachyna consimilis (Vain.) Hale, Smithsonian Contr. Bot. 25: 28. 1975.

Type — [Brasil] Minas Minas Geraés, Caraca, 1885. Vainio, Lich. Brasil. Exs. 1295

(TUR-V 3022 - lectotype, selected by Hale 1975: 28).

162 ... Flakus, Rodriguez Saavedra & Kukwa

This species is known from Mexico, the Caribbean, Venezuela, and Brazil in

the Neotropics, and also Taiwan and Papua New Guinea (Hale 1975, Louwhoff

& Elix 2002, Sipman et al. 2009 and literature cited therein). Here it is reported

as new to Bolivia.

SPECIMENS EXAMINED — BOLIVIA. DEPT. COCHABAMBA. PROV. CARRASCO, Carrasco

National Park, near Phaqcha, 20 km of Monte Punku village, 17°27'13"S 65°16'44"W,

2850 m, 2008, Kukwa 6161 (UGDA, LPB); DEPT. La Paz. Prov. B. SAAVEDRA, Cerro

Asunta Pata, 1500 m, 15°05'S 68°29'W, 1997, Bach et al. 204 (LPB).

ADDITIONAL SPECIMEN EXAMINED — Vainio, Lich. Brasil. Exs. 1133 (TUR-V 3021,

fertile).

Remototrachyna singularis (Hale) Flakus, Kukwa & Sipman, comb. nov.

MycoBank MB 561983

= Parmelia singularis Hale, Phytologia 28: 267. 1974.

= Hypotrachyna singularis (Hale) Hale, Smithsonian Contr. Bot. 25: 63. 1975.

This rare Neotropical species previously known from Colombia, Ecuador

and Peru (Hale 1975, Sipman et al. 2009), is reported as new to Bolivia.

SPECIMEN EXAMINED — BOLIVIA. DEPT. COCHABAMBA. PROV. CHAPARE, near

Incachaca village, 2317 m, 17°14'11"S 65°49'02"W, 2006, Flakus 8320 (KRAM, LPB,

herb. Flakus).

Remototrachyna sipmaniana Kukwa & Flakus, sp. nov. PLATE 1

MycoBank MB 561927

Remototrachynae speciebus medulla acidum protocetraricum et isidiis cortice acida

lecanoricum et gyrophoricum continente differt.

Type — Bolivia, Dept. Cochabamba, Prov. Carrasco, Carrasco National Park, near

Phaqcha, 20 km of Monte Punku village, 17°27'13"S 65°16'44"W, 2850 m, Yungas

montane cloud forest, close to the river, on rock, 20 July 2008, M. Kukwa 6166 (UGDA-

L 15224 — holotype; LPB - isotype).

EryMoLocy — ‘The species is named after Harrie J.M. Sipman (Botanisches Museum,

Berlin Dahlem), an eminent lichenologist dealing with tropical lichens, and co-author

of the monograph of the genus Hypotrachyna in the Neotropics.

MORPHOLOGICAL CHARACTERS — THALLUS saxicolous, 10-20 cm wide,

moderately adnate, not coriaceous, irregularly lobate; Loses subirregular,

broad, 5-8(-10) mm wide, usually black-rimmed, with entire, crenate, dissected

or irregularly incised margins and subrotund apices, usually distinctly ciliate,

sometimes with few short and ciliate laciniae; UPPER SURFACE gray to whitish-

gray, sometimes brownish at the tips, smooth or finely rugulose, shiny, in older

parts usually delicately cracked, isidiate, lacking soredia, pustules, dactyls or

lobules; 1stp14 laminal (mostly present in older parts of the thallus), rarely

marginal, scattered to moderately dense, slender, erect, simple to branched,

concolorous with thallus, but apices brown, rarely slightly procumbent, intact,

few with very short cilia; cit1a abundant, black, similar to rhizines, simple

to dichotomously branched; MEDULLA white; LOWER SURFACE black, shiny,

Remototrachyna sp. & combs. nov. (Bolivia) ... 163

PiaTE 1. Morphology of Remototrachyna sipmaniana (holotype). A: lobes; B: thallus; C: apothecia

with isidiate margin; D: branched marginal cilia; E: isidia. Scale bars: A, B = 1 cm; C = 1 mm;

D, E=0.5 mm.

smooth, with a bare brown marginal zone; RHIZINES moderately dense, but

scarce and shorter near the margin, simple (at the margin) or more often up

to 3 times dichotomously branched, black to dark brown, with paler apices;

APOTHECIA laminal, subpedicellate, 2-5 mm wide; disc concave, light brown

to chest-nut brown; THALLINE MARGIN entire or crenate, concolorous with the

thallus, rarely brown near the disc, usually isidiate (isidia shorter and wider

than those present on the thallus); CUPULATE EXCIPLE c. 50 um high, consisting

of plectenchyma with thick cell walls; HyYMENIUM c. 50 um high; suBHYMENIUM

c. 40 um; asci 8-spored; ascospores hyaline, ellipsoid, 9.5-11 x 5.5-6 um;

PYCNIDIA not seen.

CHEMISTRY — Atranorin (major; thalline and apothecial margin cortex),

protocetraric acid (major; medulla of thallus and apothecial margin) and

gyrophoric acid (submajor, with minor amounts of lecanoric acid; cortex

of isidia). Spot test reaction and fluorescence: upper cortex of thallus and

apothecial margin: K+ yellow, C-, P-, UV-; medulla of thallus, isidia and

apothecial margin: P+ red-orange, C-; cortex of isidia: P-, C+ pink red.

164 ... Flakus, Rodriguez Saavedra & Kukwa

DISTRIBUTION AND HABITAT — The new species is known only from two

localities in Carrasco National Park, Bolivia. It grows on boulders in Yungas

mountain cloud forest.

ADDITIONAL SPECIMENS EXAMINED — BOLIVIA. DEPT. COCHABAMBA. PROV.

CHAPARE, near Incachaca village, 17°14'09"S 65°48'51" W, 2198 m, Yungas montane

cloud forest, on rock, 2006, Flakus 7852 (B, KRAM, LPB, herb. Flakus).

REFERENCE MATERIAL EXAMINED — JAPAN. PRov. Kir: Koya. Saxicolous. October 19,

1952. Coll. Y. Asahina 52101 (TNS - isolectotype of Parmelia koyaensis).

COMMENTS — Remototrachyna sipmaniana is distinguished from all other

species of the genus by the abundant, branched cilia. However, the structure

of the cupulate exciple and the general morphology agree with the diagnostic

characters of Remototrachyna.

The species shows considerable variation in the production of isidia. In the

holotype they are very abundant on the thallus as well as on the apothecial

margin, but in the paratypes isidia are less frequent; in some thalli they are

sparse and many apothecia are not isidiate.

Remototrachyna sipmaniana is also distinguished by the production of

protocetraric acid in the medulla of the thallus, isidia and apothecial margin

whereas gyrophoric acid is restricted to the cortex of the isidia. Three other

isidiate species of Remototrachyna, R. incognita (Kurok.) Divakar & A. Crespo,

R. consimilis and R. koyaensis (Asahina) Divakar & A. Crespo are chemically

similar, but differ in other characters. Remototrachyna incognita contains

gyrophoric acid, but together with protolichesterinic acid (Divakar & Upreti

2005), whereas R. sipmaniana produces protocetraric and gyrophoric acids; the

former occurs in Asia (Divakar & Upreti 2005), while the latter occurs in the

Neotropics.

Both R. consimilis and R. koyaensis contain protocetraric acid (e.g., Hale

1975, Divakar & Upreti 2005, Sipman et al. 2009), and although gyrophoric acid

has been detected in some specimens in minor or trace amounts, the reaction

with C has been always reported as negative (Louwhoff & Elix 2002, Sipman et

al. 2009). Remototrachyna consimilis has very similar ascospore dimensions to

R. sipmaniana (see Sipman et al. 2009), but it differs by having much narrower

lobes, (0.5-)1-2(-3) mm wide, and partly yellow-pigmented lower medulla

(Sipman et al. 2009). Remototrachyna koyaensis can be readily separated by

the production of caperatic acid (absent in R. sipmaniana) and the ascospore

size (9.5-11 x 5.5-6 um in R. sipmaniana; versus more than 14 um long and

7 um wide in R. koyaensis) (Louwhoff & Elix 2002, Divakar & Upreti 2005,

Sipman et al. 2009, sub Hypotrachyna). Additionally, they differ in distribution:

R. koyaensis occurs in Asia, Papua New Guinea and in Mexico (Louwhoff &

Elix 2002, Sipman et al. 2009), whereas R. sipmaniana occurs in Bolivia. We

examined the types of Parmelia consimilis and P. koyaensis and did not detect

Remototrachyna sp. & combs. nov. (Bolivia) ... 165

gyrophoric acid by TLC nor did the isidia react with C. Possibly gyrophoric

acid is a relatively rare accessory substance in R. consimilis and R. koyaensis, but

the chemical variability needs to be studied in more collections.

In the Neotropics R. sipmaniana could be also confused with two other

isidiate species, R. costaricensis (Nyl.) Divakar & A. Crespo and R. rhabdiformis

(Kurok.) Divakar & A. Crespo. However, both have alternative chemistry, the

former produces fatty acids and the latter norstictic, salazinic and consalazinic

acids (Sipman et al. 2009).

The new species could also be mistaken for some chemically similar isidiate

species of Hypotrachyna, but they do not contain major quantities of both

protocetraric and gyrophoric acids in the same thallus, nor are they distributed

as in R. sipmaniana.

Acknowledgments

We are very grateful to Dr Harrie J.M. Sipman and Professor J.A. Elix (Canberra)

for reviewing the manuscript, and for valuable and constructive comments on previous

versions of this paper. We are also greatly indebted to Rosa I. Meneses Q., the Director

of Herbario Nacional de Bolivia, Universidad Mayor de San Andrés, La Paz for generous

cooperation, and the Curators of herbaria for the loan of specimens. A. Flakus warmly

thanks Dr Harrie J.M. Sipman (Berlin) for making specimens available for this study,

and for kind hospitality during his visit to the B herbarium. This research received

support from the National Centre for Research and Development (NCBiR) in Poland

under the LIDER Programme for the years 2010-2013 (no. 92/L-1/09).

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39: 52-69. http://dx.doi.org/10.1016/j.ympev.2005.12.015

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MY COTAXON

http://dx.doi.org/10.5248/119.167

Volume 119, pp. 167-173 January-March 2012

Mycena moconensis, a new species in section Polyadelphia

from Argentina

N. Niveiro*’, O. Poport’, D. DESJARDIN? & E. ALBERTO?

‘Instituto de Botanica del Nordeste, IBONE (UNNE-CONICET),

Sargento Cabral 2131, CC 209 Corrientes Capital, CP 3400, Argentina

"Dept. of Biology, San Francisco State University,

1600 Holloway Ave., San Francisco, CA 94132, U.S.A.

°Instituto Tecnologico Chascomus, IIB-INTECH (UNSAM-CONICET),

Cam. Circ. Laguna Km. 6, Chascomus, Buenos Aires, CP 7130 Argentina

* CORRESPONDENCE TO: niconiveiro@hotmail.com

ABSTRACT — A new agaric species, Mycena moconensis, is described from Paranaense

Rain Forest, Argentina. This species is characterized by small bicoloured basidiomata with

yellow to orange yellow or golden pilei and magenta to purple stipes, pip-shaped spores, and

spinulose cheilocystidia and pellicular hyphae.

Key worps — Agaricomycetes, Agaricales, Mycenaceae, new taxa, taxonomy

Introduction

Mycena (Pers.) Roussel comprises approximately 500 species, widely

distributed in the world (Kirk et al. 2008). It is a polyphyletic genus where

comprehensive molecular analyses are needed to clarify the infrageneric

classification and species circumscriptions.

South American species of Mycena have been studied by Spegazzini (1887,

1898), Rick (1938), Dennis (1961), Singer (1969, 1973, 1989), Valenzuela &

Moreno (1995), Raithelhuber (1984a,b, 1985a-f, 1996a,b, 2004) and Maas

Geesteranus & de Meijer (1997, 1998). Raithelhuber (1985f, 1996a) published

monographic studies that included identification keys to Mycena species in the

region. Maas Geesteranus & de Meijer (1997) also provided a key to species

described or mentioned previously by Singer.

There are 114 Mycena species so far recorded in Argentina. Most have been

found in the Andino-Patagonic forest and the Yungas region of northwestern

Argentina, with only six species known from the Paranaense forest (Spegazzini

1926, Wright et al. 2008, Lechner et al. 2006, Wright & Wright 2005, Niveiro

et al. 2010).

168 ... Niveiro & al.

According to Maas Geesteranus, “many more species of Mycena await to be

discovered in Parana State (Brazil) but, however desirable, the search cannot

for various reasons be continued” (Maas Geesteranus & de Meijer 1998).

During a survey of the Agaricomycetes of northern Argentina, we recently

collected some specimens that do not match any other known species of Mycena.

We propose herein M. moconensis, a new species in section Polyadelphia.

Material & methods

Macroscopic description is based in fresh material, according to Largent (1986) and

Lodge et al. (2004). Microscopic features are described from material mounted in KOH

5%, phloxine (1%), and Melzer’s reagent. The following notations are used: L = number

of lamellae reaching the stipe; x = arithmetic mean of the spore length and width;

Q = quotient of length and width indicated as a range of variation; Qm = mean of Q

values; n = number of spores measured. Herbarium specimens are deposited in CTES,

BAFC, and LIL. Herbarium abbreviations follow Holmgren et al. (1990).

Results

Mycena moconensis Niveiro, Alberté & Desjardin, sp. nov. Fics 1-8

MycoBank MB519458

Differing from other species of Mycena sect. Polyadelphya by its yellow to golden pileus

and magenta to purple stipe.

Type: Argentina, Misiones, Parque Provincial Mocona, Sendero dela Gruta, (27°09'13.2"S

53°54'04.7"W), 17.V.2008, leg. N. Niveiro et al. 767, (Holotype, CTES 0591002).

ETYMOLOGy: From Mocona Provincial Park.

BASIDIOMATA scattered to gregarious. PILEUs 1-8 mm in diameter, broadly

parabolical to hemispherical or campanulate, without umbo; margin entire,

decurved, translucent striate to sulcate; surface smooth, slightly pruinose to

silky, dry; yellow to orange yellow or golden. CONTEXT thin. ODor and TASTE

absent. LAMELLAE adnate to slightly decurrent by a tooth, subdistant (L = 11-

13) without lamellulae, up to 0.5 mm wide, white; entire edge, concolorous.

STIPE 30-60(-90) x 0.5-1.5 mm, central, cylindrical, equal, hollow, smooth,

glabrous, apex magenta to dark magenta, purple or Indian red, paler on the

base; attached to the substratum by a mycelial pad of brown radiating hyphae.

SPORE PRINT white.

BASIDIOSPORES 5.7-8.6 x 3-4.5 um, [x = 7.5 x 4 um, Q = 1.59-2.27;

Q_, = 1.87, n = 38), oblong to pip-shaped, smooth, hyaline, amyloid, thin-

walled, without germ pore. Bastp1A 20-23 x 6-7 um, clavate, 4-sterigmate.

PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 10-16 x 7-10 um, hyaline, broadly

clavate to subglobose, densely spinulose, with simple cylindrical excrescences

2-2.6 um long. HYMENOPHORAL TRAMA regular; hyphae 2.5-11 um diam,

dextrinoid, non-gelatinous. PILEIPELLIs a thin cutis of repent, radially aligned

hyphae; hyphae 2.5-5.5 um diam., densely covered with short cylindrical

Mycena moconensis sp. nov. (Argentina) ... 169

Fics 1-3. Mycena moconensis: 1. General aspect; 2. Detail of pileus; 3. Stipe with mycelial

filaments radiating from the base. Scale bars: 1 = 20 mm; 2 = 2 mm; 3 = 1 mm.

170 ... Niveiro & al.

Fics 4-8. Mycena moconensis: 4. Basidia; 5. Basidiospores; 6. Cheilocystidia; 7. Hyphae of the

stipitipellis; 8. Pileipellis and hypoderm. Scale bar = 10 um.

spinulae, non-gelatinous. HyPODERMIUM pseudoparenchymatous, with cells

18-37 um diam., dextrinoid. sTIPITIPELLIS hyphae 3.5-5.5 um diam., densely

covered with short cylindrical spinulae. CAuLocystTip1A absent or rare and

scattered. CLAMP CONNECTIONS present.

Habitat: Gregarious in small clusters, on leafy litter.

ADDITIONAL SPECIMENS EXAMINED — ARGENTINA, Misiones, Parque Provincial

Mocona, Sendero de la Gruta, (27°09'17.1"S 53°54'02.6"W), 25/V/2009, Niveiro &

Michlig 1276 (BAFC); 25/V/2009, Niveiro & Michlig 1296 (LIL).

Mycena moconensis sp. nov. (Argentina) ... 171

Discussion

This beautiful new species is characterized by small bicolored basidiomata

with yellow to orange yellow or golden pilei and magenta to purple stipes.

The following combination of features refer this species to section

Polyadelphia (Maas Geesteranus 1986, Maas Geesteranus & de Meijer 1997):

small basidiomata with few lamellae, thin stipes with radiating basal mycelium,

cheilocystidia densely covered with simple excrescences, pip- shaped

basidiospores, and a non-gelatinous pileipellis composed of hyphae densely

ornamented with small spinulae.

Mycena moconensis could be confused with “M. coprinoides” P. Karst. sensu

Raithelh., a species previously recorded in Argentina (Raithelhuber 2004).

The latter species differs, however, in forming a pale yellow to pale ochraceous

pileus with a weakly striate margin, crowded and adnexed-adnate lamellae, a

pale yellow stipe with a darker base, and fusiform cheilocystidia. It should be

noted that Maas Geesteranus (1981) considers Mycena coprinoides P. Karst. as

representing a species of Coprinus, based on his examination of the holotype

specimen. The taxon reported by Raithelhuber (2004) as “M. coprinoides” may

represent an undescribed species.

The presence ofa citron yellow to barium yellow pileus, sulcate-striate margin,

adnate-decurrent, distant lamellae, and pyriform to clavate cheilocystidia with

excrescences also diagnose Mycena citricolor (Berk. & M.A. Curtis) Sacc., a

pathogen of coffee known from the New World tropics. Mycena citricolor differs

in forming a yellow stipe, slightly smaller basidiospores (4.2-6 x 2.3-3.5 um),

and pileipellis hyphae that are embedded in a gelatinous matrix (Pegler 1983).

It also differs in forming bioluminescent mycelium on which are produced

stilboid asexual propagules with the anamorphic Decapitatus flavidus (Cooke)

Redhead & Seifert.

The similar M. melinocephala Singer, described from Argentina on Alnus

wood, differs in forming a yellowish honey-coloured pileus and stipe and

larger, subglobose to globose basidiospores (8-10.5 x 7.5-9.5 um) (Singer

1973). Mycena ixoxantha Singer, a species described from Colombia with

citrine yellow pileus and amber coloured stipe, differs in forming gelatinous

pileipellis tissue (Singer 1973).

Two species from the northern hemisphere, Mycena albiceps (Peck) Gilliam

and M. capillaris (Schumach.) P. Kumm., share similar micro-morphological

features with M. moconensis but differ in forming white to pale greyish white

pilei and dark greyish brown to black stipes (Maas Geesteranus 1986).

Maas Geesteranus & de Meijer (1997, 1998) described two species of Mycena

sect. Polyadelphia — M. elongata Maas Geest. & de Meijer and M. tuberifera

Maas Geest. & de Meijer — from Parana State (Brazil) near the location where

M. moconensis was found. They are characterized by very dark brown pilei and

172 ... Niveiro & al.

a pale yellow (M. elongata) or white (M. tuberifera) stipe. The authors regarded

both species as unusual for section Polyadelphia by differing from northern

hemisphere representatives in forming smooth stipitipellis hyphae and simple,

non-spinulose caulocystidia, characters not observed in M. moconensis.

Acknowledgments

The authors wish to thank T.J. Baroni and R.H. Petersen for the critical revision of

the manuscript. This research was made possible by the support of Reserva de Biosfera

Yaboty, MERNRyT - Proyecto Regional Araucaria XXI, Bosque Atlantico - AECID;

Myndel Botanical Foundation; SGCyT (UNNE) and the Argentine Nat'l Research

Council (CONICET).

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agaricalium IV). Fieldiana, Bot. 21: 1-133.

Spegazzini C. 1887. Fungi fuegiani. Bol. Acad. Nac. Ci. Republ. Argent.11: 135-308.

Spegazzini C. 1898. Fungi argentini novi vel critici. Anales Mus. Nac. Hist. Nat. “Bernardino

Rivadavia’” 6: 81-365.

Spegazzini C. 1926. Observaciones y adiciones a la micologia argentina. Bol. Acad. Nac. Ci. Republ.

Argent. 28: 276-406.

Valenzuela E, Moreno G. 1995. Contribucidn al estudio del género Mycena (Agaricales,

Basidiomycotina) en la X Region de Chile. Bol. Soc. Micol. Madrid 20: 179-194

Wright JE, Wright AM. 2005. Checklist of the mycobiota of Iguazu National Park (Misiones,

Argentina). Bol. Soc. Argent. Bot. 40(1-2): 23-44.

Wright JE, Lechner BE, Popoff OF. 2008. Atlas pictdérico de los hongos del Parque Nacional Iguazu.

Ed. L.O.L.A. Buenos Aires. Argentina. 228 p.

MY COTAXON

http://dx.doi.org/10.5248/119.175

Volume 119, pp. 175-179 January-March 2012

A new species of Fulvifomes (Hymenochaetaceae)

from Cambodia

L1I-WEI ZHOU! & WEI-MIN ZHANG?

“State Key Laboratory of Forest and Soil Ecology, Institute of Applied Ecology,

Chinese Academy of Sciences, Shenyang 110164, P. R. China

*Guangdong Institute of Microbiology, Guangzhou 510070, P. R. China

* CORRESPONDENCE TO: liwei_zhou1982@163.com

ABSTRACT — Fulvifomes cambodiensis sp. nov. is described and illustrated from Preah Vihear,

Cambodia. It produces perennial, pileate and solitary basidiomes with a concentrically

sulcate zonate pileal surface, homogeneous context, hymenial setae, and yellowish

acyanophilous basidiospores. The new species macroscopically resembles F. rimosus, which

differs in lacking hymenial setae and having cyanophilous generative hyphae and slightly

smaller basidiospores. The macroscopically similar Fomitiporia hartigii is distinguished by

larger hyaline basidiospores and the absence of setae. Fulvifomes johnsonianus has abundant

hymenial setae and quite similar hyphal structure to E cambodiensis, but its much smaller

basidiospores and effused-reflexed to pileate basidiomes distinguish it from the new species

with pileate and ungulate basidiomes with cracked pileal surface.

Key worps — Hymenochaetales, polypore, taxonomy

Introduction

After Murrill (1914) segregated Fulvifomes Murrill (Hymenochaetaceae

Donk), typified by F. robiniae (Murrill) Murrill, from Phellinus Quél., many

mycologists did not accept his genus (Ryvarden & Johansen 1980, Gilbertson

& Ryvarden 1987, Ryvarden & Gilbertson 1994, Nufiez & Ryvarden 2000),

although it was treated as a subgenus of Phellinus by Dai (1999). Recent molecular

phylogenetic analyses clearly support it as an independent genus closely related

to Aurificaria D.A. Reid and Phylloporia Murrill (Wagner & Fischer 2002,

Larsson et al. 2006). Morphologically, while all three genera produce colored

basidiospores, the dimitic hyphal system of Fulvifomes distinguishes it from the

two monomitic genera (Dai 2010). Except for the dimitic hyphal system and

colored basidiospores, Fulvifomes species have few distinctive morphological

characters (Dai 2010).

176 ... Zhou & Zhang

During a study on wood-decaying fungi in Cambodia, an unknown

Fulvifomes species was discovered, which is here described and illustrated as

Fulvifomes cambodiensis. Morphological differences between F. cambodiensis

and other similar species according to Chinese materials are also noted.

Materials & methods

The studied specimen was deposited at the herbaria of Guangdong Institute

of Microbiology (GDGM) and Institute of Applied Ecology, Chinese Academy of

Sciences (IFP). The microscopic procedure follows Dai (2010). Sections were studied at

magnification up to x1000 using a Nikon Eclipse E 80i microscope and phase contrast

illumination. The variation in the size of the spores was presented by excluding 5%

of measurements from each end of the range, which were given in parentheses. The

abbreviations are used as follows: IKI = Melzer’s reagent, IKI- = negative in Melzer’s

reagent, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB- = acyanophilous,

L = mean spore length (arithmetic average of all spores), W = mean spore width

(arithmetic average of all spores), Q = variation in the L/W ratios within the specimen

studied, and n = number of spores, hyphae or setae measured from given number of

specimens. Line drawings were made with the aid of a light tube. Special color terms

follow Anonymous (1969) and Petersen (1996).

Taxonomy

Fulvifomes cambodiensis L.W. Zhou & W.M. Zhang, sp. nov. FIGs 1-2

MycoBank MB 561815

Differs from Fulvifomes rimosus in the presence of hymenial setae, slightly larger

basidiospores, and acyanophilous generative hyphae.

Type: Cambodia, Preah Vihear, on angiosperm wood, 12.V.2006, GDGM 20949

(holotype in GDGM, isotype in IFP).

EryMo.Locy: cambodiensis (Lat.): refers to the country of Cambodia.

BASIDIOMES: Perennial, pileate, ungulate, in section triquetrous, usually solitary,

woody hard and without odour or taste when dry. Pileus projecting up to 6

cm, 15 cm wide and 8 cm thick at base; pileal surface deep reddish brown to

dark clay-buff, glabrous, concentrically sulcate with wide zones, glabrous with

age, cracked to slightly rimose with age, margin obtuse. Pore surface yellowish

brown, sterile margin yellowish brown, up to 10 mm; pores round to angular,

5-6 per mm; dissepiments thin, entire. Context cinnamon, woody hard, up to

13 mm thick. Tubes cinnamon, woody hard, tube layers distinct, annual layer

up to 3 mm long.

HyPHAL STRUCTURE: Dimitic; generative hyphae without clamp connections;

skeletal hyphae IKI-, CB-; tissues darkening but otherwise unchanged in

KOH.

CONTEXT: Generative hyphae scanty, hyaline, thin-walled, occasionally

branched, 2-3 um in diam; skeletal hyphae dominant, yellowish brown, thick-

Fulvifomes cambodiensis sp. nov. (China) ... 177

FiGuRE 1. Fulvifomes cambodiensis basidiocarp (holotype).

walled, flexuous, unbranched, without a simple septum, tightly interwoven,

3-5 um in diam (n = 10/1).

TuBeEs: Generative hyphae frequent, hyaline, thin-walled, occasionally

branched, frequently septate, sometimes diminishing at the end or middle,

1.7-2.5 um in diam; skeletal hyphae dominant, yellowish brown, thick-walled

with a distinct lumen to subsolid, flexuous, unbranched, interwoven, sometimes

agglutinated, 2-3.5 um in diam (n = 10/1). Hymenial setae frequent, dark

brown, subulate to ventricose, thick-walled, 12.7-30.4 x 6.5-17 um (n = 10/1).

Hymenium collapsed, cystidia, cystidioles, basidia and basidioles not seen.

Spores: Basidiospores broadly ellipsoid, yellowish, thick-walled, smooth,

CB-, IKI-, (5.5-)5.8-6.2(-6.3) x (4.5-)4.6-5.2 um, L = 5.98 um, W = 4.95 um,

Or= 1.241 n= 3071).

REMARKS: Fulvifomes cambodiensis is macroscopically characterized by

solitary perennial, pileate, woody hard basidiomes with a crust-covered and

concentrically sulcate pileal surface and homogeneous context; microscopically

it has hyaline thin-walled generative hyphae, yellowish brown thick-walled

skeletal hyphae, hymenial setae, and yellowish acyanophilous basidiospores.

Fulvifomes rimosus (Berk.) Fiasson & Niemela shares with E cambodiensis

a perennial pileate ungulate growth habit, concentrically sulcate zonate pileus,

178 ... Zhou & Zhang

10 ym

FiGuRE 2. Fulvifomes cambodiensis (holotype).

a: Spores. b: Setae. c: Hyphae from trama. d: Hyphae from context.

and obtuse pileal margin. However, the basidiomes of F. rimosus are darker.

Microscopically, E rimosus lacks hymenial setae and has slightly smaller

basidiospores (L = 5.54 um, W = 4.54 um) and cyanophilous generative hyphae

(Dai 2010).

Fulvifomes cambodiensis sp. nov. (China) ... 179

The macroscopically similar Fomitiporia hartigii (Allesch. & Schnabl)

Fiasson & Niemela shares with E cambodiensis the perennial and ungulate

basidiomes with the concentrically sulcate cracked pileal surface, obtuse pileus

margin, and almost identical pore sizes (Dai 2010), making the two species

very hard to differentiate in the field. But F hartigii is distinguished by its wider

hyphae, lack of setae, and larger hyaline basidiospores that are dextrinoid and

strongly cyanophilous (Dai 2010).

Fulvifomes johnsonianus (Murrill) Y.C. Dai has quite similar hyphal structure

to F cambodiensis, and the two species also share abundant hymenial setae and

ellipsoid basidiospores (Q = 1.23, Dai 2010). However, E johnsonianus has

much smaller basidiospores (2.8-3.2 x 2.2-2.6 um; Dai 2010). Macroscopically,

F. johnsonianus has effused-reflexed to pileate basidiomes, which are easily

differentiated from the pileate and ungulate basidiomes with cracked pileal

surface of F cambodiensis (Dai 2010).

Acknowledgements

We express our gratitude to Drs. Bao-Kai Cui (Beijing Forest University, China) and

Rosa Mara Borges da Silveira (Universidade Federal do Rio Grande do Sul, Brazil) who

reviewed the manuscript. The research was financed by the National Natural Science

Foundation of China (Project No. 30910103907).

Literature cited

Anonymous. 1969. Flora of British fungi. Colour identification chart. Her Majesty’s Stationery

Office, London. 1 p.

Dai YC. 1999. Phellinus sensu lato (Aphyllophorales, Hymenochaetaceae) in East Asia. Acta Botanica

Fennica 166: 43-103.

Dai YC. 2010. Hymenochaetaceae (Basidiomycota) in China. Fungal Diversity 45: 131-343.

http://dx.doi.org/10.1007/s13225-010-0066-9

Gilbertson RL, Ryvarden L. 1987. North American polypores 2. Megasporoporia—Wrightoporia.

Fungifora, Oslo. pp. 431-885.

Larsson K-H, Parmasto E, Fischer M, Langer E, Nakasone KK, Redhead SA. 2006. Hymenochaetales:

a molecular phylogeny for the hymenochaetoid clade. Mycologia 98: 926-936.

http://dx.doi.org/10.3852/mycologia.98.6.926

Murrill WA. 1914. Northern polypores. Privately printed, New York. 64 p.

Nujfiez M, Ryvarden L. 2000. East Asian polypores 1. Ganodermataceae and Hymenochaetaceae.

Synopsis Fungorum 13: 1-168.

Petersen JH. 1996. Farvekort. The Danish Mycological Society’s colour-chart. Foreningen til

Svampekundskabens Fremme, Greve. 6 p.

Ryvarden L, Gilbertson RL. 1994. European polypores 2. Synopsis Fungorum 7: 388-743.

Ryvarden L, Johansen I. 1980. A preliminary polypore flora of East Africa. Fungifora, Oslo. 636 p.

Wagner T, Fischer M. 2002. Proceedings towards a natural classification of the worldwide taxa

Phellinus s.l. and Inonotus s.l., and phylogenetic relationships of allied genera. Mycologia 94:

998-1016. http://dx.doi.org/10.2307/3761866

MY COTAXON

http://dx.doi.org/10.5248/119.181

Volume 119, pp. 181-195 January-March 2012

Fungal pathogens of ‘cat’s claws’ from Brazil

for biocontrol of Macfadyena unguis-cati

MEIRIELE DA SILVA, ROBERT WEINGART BARRETO*

& OLINTO LIPARINI PEREIRA

Departamento de Fitopatologia, Universidade Federal de Vicosa

Vicosa, Minas Gerais, 36570-000, Brazil

*CORRESPONDENCE TO: rbarreto@ufv.br

ABsSTRACT—Recent searches in Brazil for plant pathogens with potential for use as classical

biocontrol agents of Macfadyena unguis-cati yielded five fungal species, two previously

undescribed. These are described herein and preliminary evaluations based on field and

laboratory observations of the diseases with which they are associated are given. These are:

Guignardia mangiferae (leaf spots), Meliola herteri (black mildew), Passalora macfadyenae

sp. nov. (leaf spots), Pseudocercospora unguis-cati (leaf spots) and Prospodium macfadyenae

sp. nov. (rust). The two newly described species, Pa. macfadyenae and Pr. macfadyenae, are

regarded as having the greatest potential for use in classical biological control.

KEY woRDs —cat’s claws, new species, plant pathology, taxonomy

Introduction

Invasion of natural ecosystems by exotic species represents one of the biggest

threats to biodiversity, as now acknowledged by scientists and governments

throughout the world (IUCN 2000). Once an alien organism has invaded, the

sole long-term sustainable management alternative available for mitigating it

through human intervention is the introduction of natural enemies collected

in the pest’s centre of origin in a process known as classical biological control.

Macfadyena unguis-cati (L.) A.H. Gentry (Bignoniaceae) — common name

cat's claws (in Brazil: unha-de-gato, unha-de-morcego) — is a woody vine that

has a broad distribution in its native range occurring from southern Brazil

and Argentina to Mexico (Everett 1980). Macfadyena unguis-cati takes its

popular names from the tendrils, which have acute divided ends that appear

and function like claws, allowing the plant to firmly attach to trees or other

supports on which it grows (PLATE 1). At maturity it produces abundant showy

yellow flowers making it a useful ornamental plant for cultivation on fences

182 ... Silva, Barreto & Pereira

and garden walls (Lorenzi & Souza 1995). Because of its value as an ornamental

it was introduced to regions outside its native range, where it escaped from

gardens to become a noxious invader of native forests, particularly in the USA,

Australia, China, and South Africa (Henderson 1995, FLEPPC 2005, Williams

et al. 2008, Osunkoya et al. 2009, Dhileepan et al. 2010). Its vigorous vine habit

allows it to ascend tall trunks reaching the treetops (PLATE 2) branching and

spreading over top of the forest canopy. The combination of the biomass weight

and shading of underlying vegetation can be very harmful and even cause the

death of the largest plants in infested areas and suppress the development of

smaller plants and seed germination of native plants (Neser 1996, Huang et al.

2009, King & Dhileepan 2009). In Florida, M. unguis-cati is listed as a Category

1 weed, a rank reserved for species capable of altering plant communities

by substituting and even permanently changing the natural structure of an

ecosystem and its ecological functions (FLEPPC 2005). Between 2003 and

2005 a search was conducted in native situations in Brazil for fungal pathogens

attacking M. unguis-cati that might have potential as biocontrol agents. ‘This

publication presents the results of this preliminary survey with particular

reference to fungal taxonomy and the potential to use mycobiota in classical

biological control.

Material & methods

Prior to the field survey, herbarium records were completely searched for occurrence

and distribution of M. unguis-cati in the following Brazilian herbaria: Herbario

Fanerogamico e Criptogamico do Instituto Agrondémico (IAC); Herbario Maria Eneyda P.

K. Fidalgo (SP); Instituto de Pesquisas Jardim Botanico do Rio de Janeiro (RB); Herbario

do Museu Nacional do Rio de Janeiro (R); Universidade Federal de Vicosa (VIC); Museu

Botanico Municipal (MBM); and Herbario José Badini (OUPR). Using this information,

we prepared an itinerary for surveying the southern and southeastern Brazilian states

Minas Gerais, Rio de Janeiro, Espirito Santo, SAo Paulo, Parana, Santa Catarina, and Rio

Grande do Sul. Surveys were conducted between January 2003 and December 2005.

Selected sites were visited and carefully scrutinized for the presence of M. unguis-cati.

For details on the general survey and laboratory methodologies see Barreto & Evans

(1994). Freshly collected samples were examined under a stereomicroscope. Slides

containing free-hand sections of colonized tissues or fungal structures scraped from

the surface of infected tissues were mounted in lactophenol or lactofuchsin. Isolations

of non-biotrophic fungi were attempted by directly transferring spores or other fungal

structures to plates containing Vegetable Broth Agar (VBA, Pereira et al. 2003), under a

dissecting microscope with a sterilized fine point needle. Observations, measurements,

photographs and line drawings were prepared using an Olympus BX 50 light microscope

fitted with a camera (Olympus Evolt SC 50) and drawing tube.

For scanning electron microscopy the samples were fixed in 2.5% glutaraldehyde +

sodium cacodylate buffer (0.1mol L’, pH 7.2) (1:1) for 1h at room temperature, washed

six times in 0.1mol L? sodium cacodylate buffer (10 m each washing period), post-fixed

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 183

PLATE 1. Macfadyena unguis-cati: a. typical large, yellow, trumpet-shaped flowers;

B. climbing on tree trunks; c. clawed tendrils which enable plant to climb up against

walls, tree trunks, and other supports.

by immersion in 1% O.O, prepared in cacodylate buffer 0.1 mol L” (1:1), and kept for 4

h at 4-8 °C then washed again six times in buffer, dehydrated by 10-min long successive

transfers in an alcohol solution (30, 50, 60, 70, 80, 95, 100%) gradation series at room

temperature, dried in a critical point dryer (Baltec model 030) with CO, as transition

184 ... Silva, Barreto & Pereira

PLATE 2. Macfadyena unguis-cati climbing trees in the field in Brazil.

fluid, and coated with 20 nm thick gold in a sputter coater (Balzers® model FDU 010)

before examination in a Carl-Zeiss Model LEO VP 1430 electron microscope.

Representative specimens were deposited in the herbarium at the Universidade

Federal de Vicosa (Herbarium VIC).

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 185

Taxonomy

Numerous samples of diseased M. unguis-cati were collected during the

survey. Five different fungal pathogens were recognized in association with

cat's claws. Two fungal species were isolated in pure culture. The five fungal

pathogens were associated with the following diseases: leaf spots (3), black

mildew (1), and rust (1). Descriptions of the identified fungi are given below.

Guignardia mangiferae A.J. Roy, Indian Phytopath. 20: 348. 1968. PLATE 3

Lesions on living leaves, minute dark tar-spots corresponding to fungal

conidiomata surrounded by faint chlorotic areas, occasionally associated to

larger necrotic areas. External mycelium absent. Pseudothecia hypophyllous,

immersed, solitary or in groups, subglobose, 136.5-113 um, wall composed of

texture angularis, 17.5-27.5 um, dark brown to black, smooth. Asci bitunicate,

subclavate to cylindrical, 46.5-82 x 11-14 um, eight-spored. Ascospores

biseriate, fusiform-elliptical, 12-14 x 4.5-5 um, unicellular, guttulate, hyaline,

smooth, surrounded by a thick mucilaginous coat.

MATERIAL EXAMINED: BRAZIL. SAo PauLo, on Macfadyena unguis-cati, 16 April 2005,

OLPereira (VIC 30492); Minas Geralts, on Macfadyena unguis-cati, 25 July 2011,

MSilva & OLPereira (VIC 31759).

PLATE 3. A-B: Guignardia mangiferae on Macfadyena unguis-cati. A. Faint chlorotic symptoms

on infected leaves; B. circular individual pseudothecia on adaxial leaf surface. c-p: Guignardia

mangiferae (VIC 30492). c. Section through a pseudothecium showing the asci; D. bitunicate asci

containing biseriate, hyaline ascospores. Bars: c = 30 um; D = 10 um.

186 ... Silva, Barreto & Pereira

COMMENTS: Guignardia mangiferae was observed causing some damage to few

isolated leaves on infected M. unguis-cati plants (PLATE 3A-B) but is regarded

here as having no potential for classical biocontrol of M. unguis-cati. This is the

first record of G. mangiferae, a polyphagous opportunistic pathogen of numerous

plant families (Rodrigues et al. 2004), on a representative of Bignoniaceae

(M. unguis-cati) worldwide.

PiateE 4. Meliola herteri (VIC 30485): a. external hyphae with alternate appressoria; B. opposite

conidiogenous cells; c. branching-dentate setae; D. ascospores. Bars: A-c = 10 um; D = 20 um.

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 187

Meliola herteri Hansf., Sydowia 9: 65. 1955. PLATE 4

Colonies amphigenous, mostly epiphyllous, 4-50 mm diam, sooty,

scattered, numerous, confluent, black. External mycelium net-forming, 6-9.5

um diam hyphae, slightly undulate, alternately branching at acute angles, dark

brown, septate, producing phialides (mucronate hyphopodia) and appressoria

(capitate hyphopodia), smooth, brown; appressoria alternate apical, stalk-cells

cylindrical 6-16 x 5.5-9.5 um, head-cells oval or elliptical, 14-19 x 9.5-14 um

dark brown; phialides intermixed with appressoria, ampulliform to cylindrical,

14-25.5 x 5.5-9 opposite or alternate, scarce. Perithecia superficial, aggregate,

globose, 112-155 um diam. Mycelial setae scattered, straight to slightly curved,

155-253 x 6-9.5 um, the apex usually 2-4 furcate and secondarily 2-3 branched,

brown. Asci evanescent at maturity. Ascospores cylindric to oblong, 44-50.5 x

13-19 um, constricted at the septa, apex and base rounded, 4-septate, brown,

smooth.

MATERIAL EXAMINED: BRAZIL. R10 GRANDE DO SUL, on Macfadyena unguis-cati, 6

August 2004, OLP (Pereira) (VIC 30485).

ComMENts: Three Meliola species are known to occur on M. unguis-cati:

Meliola thaxteri Hansf., M. bidentata Cooke, and M. herteri (Hansford 1961).

The morphology of the specimen collected on M. unguis-cati during the survey

clearly placed this fungus within M. herteri as described by Hansford (1961).

Meliola herteri has been previously recorded on M. unguis-cati in Paraguay,

Amphilophium vauthieri DC. in Argentina, and Parabignonia maximilianii

(Mart ex DC.) Bureau in Brazil (Hansford 1961). This is the first report of M.

herteri on M. unguis-cati in Brazil. Meliola herteri is, as are the Meliolaceae in

general, a weak obligate biotroph and the black mildew diseases the family

causes do not significantly affect their hosts (Hansford 1961). This fungal group

therefore has little relevance for use in biological control.

Passalora macfadyenae Meir. Silva, O.L. Pereira & R.W. Barreto, sp.nov. PLATE 5

MycoBANnk 561616

Morphologically similar to Passalora adenocalymmatis and Passalora pyrostegiae but

differs in shorter (31.5-114 um) conidia, longer (16-114 tum) conidiophores, and a well-

developed stroma.

Type: Brazil, Minas Gerais, on Macfadyena unguis-cati, 18 May 2004, OLP (Pereira)

(VIC 30480, holotype).

EryMo_oey: from the host genus name.

Lesions on living leaves, similar to those cause by P. unguis-cati but somewhat

darker centrally, 4-55 mm in diam., coalescing to cover large areas of leaf surface

and leading to leaf blight. Internal mycelium indistinct. External mycelium

absent. Stromata erumpent, 63-142 x 35-95 um, dark brown. Conidiophores

hypophyllous arising from stromata, in dense fascicles, cylindrical, straight to

188 ... Silva, Barreto & Pereira

PLaTE 5. A: Passalora macfadyenae on Macfadyena unguis-cati: necrotic symptoms on infected

leaves. B-c: Passalora macfadyenae (VIC 30480): B. conidiophore fascicle on stroma; c. conidia

(straight to slightly curved). Bars = 10 um.

slightly curved, 16-114 x 3-6 um, 0-8 septate, unbranched, brown, smooth.

Conidiogenous cells polyblastic, integrated, mostly cylindrical, 5-17.5 x 3.5-5

um, brown. Conidiogenous loci conspicuous, 1-3 per conidiogenous cell,

1-2.5 um, thickened and darkened. Conidia dry, solitary, cylindrical-obclavate,

straight to slightly sinuous, 31.5-114 x 3-4.5 um, base obconically truncate,

apex rounded, 1-12 septate, hilum thickened and darkened, subhyaline to

olivaceous, smooth.

ADDITIONAL MATERIAL EXAMINED: BRAZIL. SAo PauLo, on Macfadyena unguis-cati,

25 July 2004, OLPereira (VIC 30487).

ComMENTs: No Passalora species are known to associate with Macfadyena taxa.

Ten Passalora species are known to attack Bignoniaceae representatives, but

all are dissimilar to the fungus collected on M. unguis-cati. Conidial sizes are

useful in separating P. macfadyenae from many of the similar Passalora species

of (TABLE 1). Passalora tecomariae Crous & B. Sutton and P. tabebuiae-ochraceae

Inacio & Dianese differ from the new species in wider conidia, P. tabebuiae

(J.J. Muchovej & EA. Ferreira) U. Braun & Crous has narrower conidia, and

P. adenocalymmatis U. Braun & Crous, P. arrabidaeae (Chupp & Viegas) Crous

et al., and P pyrostegiae (Viégas) U. Braun & Crous have longer conidia, smaller

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 189

TABLE 1. Conidial size of Passalora species recorded on members of the Bignoniaceae.

SPECIES ConIDIA (um) REFERENCES

P. adenocalymmatis 35-150 x 3-4.5 Chupp (1954), as “Cercospora adenocalymmae”

P. arrabidaeae 35-150 x 3.5-5 Chupp (1954), as Cercospora arrabidaeae

P. catalpae 40-120 x 2.5-4.5 Chupp (1954), as Cercospora catalpae

P. catalparum 35-125 x 3.5-6 Chupp (1954), as Cercospora catalparum

P leprosa 30-80 x 3.5-6 Chupp (1954), as Cercospora leprosa

P. macfadyenae 31.5-114 x 3-4.5 This publication

P. markhamiae 16-92.5 x 2.5-55.5 Liu & Guo (1982)

P. pyrostegiae 25-170 x 3-4.5 Chupp (1954), as Cercospora pyrostegiae

P. tabebuiae (6—-)8-30(-34) x 2.5-3 | Muchovej & Ferreira (1981)

P. tabebuiae-ochraceae 31-75 x 5-8 Inacio & Dianese (2006)

P. tecomariae 35-90 x 5-6 Crous & Sutton (1997)

or absent stromata, and shorter conidiophores. Passalora caltaparum (Chupp)

U. Braun & Crous has longer conidia, superficial hyphae, and solitary shorter

conidiophores. Passalora markhamiae (X.J. Liu & Y.L. Guo) U. Braun &

Crous has smaller stromata and shorter conidiophores and conidia. Passalora

leprosa (Speg.) U. Braun is easily distinguished by having much larger stromata

(300-500 um diam), longer conidiophores (40-150 um), and shorter conidia

(30-80 um). Additionally, P catalpae (Chupp) U. Braun & Crous has smaller

(<50 um) or no stromata, longer (< 125 um) conidiophores that are geniculate

and rarely septate, and conidia that are acicular-shaped, indistinctly septate,

and longer (40-120 um) than those of P. macfadyenae. Passalora macfadyenae

causes severe necrosis and significant defoliation of its host in the field (PLATE

5A) and appears to have good potential for use in classical biological control of

M. unguis-cati.

Pseudocercospora unguis-cati (Speg.) U. Braun, Mycotaxon 51: 49.1994. PLATE 6

Lesions on living leaves, amphigenous, subcircular, well delimited, infected

tissue initially dark brown surrounded by a pale brown halo, becoming grayish

centrally with a narrow dark brown outer rim at the periphery, 3-35 mm

diam., coalescing and leading to necrosis of extensive areas of leaves. Internal

mycelium indistinct. External mycelium absent. Stromata absent or small and

composed of only a few substomatal swollen cells, pale brown. Conidiophores

hypophyllous emerging through stomata, solitary or forming loose fascicles

of few conidiophores, straight to slightly curved, sometimes restricted to the

conidiogenous cell, cylindrical, 11-50 x 2.5-5 um, 0-2 septate, unbranched,

light brown, smooth, with inconspicuous scars. Conidiogenous cells terminal,

holoblastic, mostly cylindrical, 10-17.5 x 2.5-5, pale brown. Conidiogenous

loci inconspicuous, flat, up to 2.5 um, unthickened, not darkened. Conidia dry,

190 ... Silva, Barreto & Pereira

PLATE 6. A. Pseudocercospora unguis-cati on Macfadyena unguis-cati: necrotic symptoms on infected

leaves. B-c. Pseudocercospora unguis-cati (VIC 26575): B. conidiophores emerging through stoma;

c. conidia. Bars = 10 um.

solitary, acicular-obclavate, straight to slightly curved, 48-119 x 3.5-7 um, base

truncate, apex rounded, 0-6 septate, hilum unthickened, not darkened, pale

brown, smooth.

MATERIAL EXAMINED: BRAZIL, Minas Gerats, on M. unguis-cati, 18 July 2004,

OLPereira (VIC 26575); 18 May 2004, OLPereira (VIC 30479, 30481, 30482); 15

February 2005, OLPereira (VIC 30488); 2 February 2005, OLPereira (VIC 30489); 3

February 2005, OLPereira (VIC 30490, 30491); 25 July 2011, MSilva & OLPereira (VIC

31758). PARANA, on M. unguis-cati, 18 July 2004, OLPereira (VIC 26576). SAo PAULO,

on M. unguis-cati, 26 July 2004, OLPereira (VIC 30486).

CoMMENTs: Only one cercosporoid species was previously known to occur

on Macfadyena — Pseudocercospora unguis-cati, which was described from

a specimen collected on M. unguis-cati in Argentina (Braun 1994). There are

numerous Pseudocercospora species on hosts belonging to the Bignoniaceae

(Crous & Braun 2003, Inacio & Dianese 2006). The survey in Brazil yielded two

cercosporoid fungi that clearly belonged to the genera Passalora (see above)

and Pseudocercospora. These were compared with the species described on

Bignoniaceae. The Pseudocercospora specimen has morphological features that

place it well within P. unguis-cati as described by Braun (1994). This is the first

report of P. unguis-cati, originally described from Argentina, on M. unguis-cati

in Brazil. Pseudocercospora unguis-cati causes necrotic spots on M. unguis-cati

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 191

(PLATE 6A) that may be severe and should be further evaluated as a potential

biocontrol agent.

Prospodium macfadyenae Meir. Silva, O.L. Pereira & R.W. Barreto, sp. nov.

MycoBAnkK 561617 PLATE 7-8

Differs from Prospodium stizophylli in larger (31.25-37.5 um) teliospores, basket-like

sori, and no mesospores and from P. perornatum in longer (47.5-55 um) teliospores,

larger (22.5-30 um) urediniospores, and paraphyses within the sori.

Type: Brazil. Minas Gerais, on Macfadyena unguis-cati, 18 May 2004, OLPereira (VIC

30478, holotype).

ErymMo_oey: from the host genus name.

Lesions on living leaves, mainly on leaf veins but occasionally also widely

distributed over the lamina, starting as chlorotic dots becoming pale brown

centrally with well defined brown borders that later become dark brown to black

and necrotic adaxially, appearing as black dots distributed within chlorotic areas,

abaxially, of variable shape, 0.5-1.5 mm, leading to premature leaf fall. Internal

mycelium intercellular 2.5-5 um diam., branched, septate. Aecia unknown.

Spermogonia epiphyllous, type 7, subepidermal, erumpent, in groups, conical,

16-38 x 6-19 um, reddish brown. Uredinia and telia hypophyllous, in small

scattered groups, basket-like sori, emerging through stomata, 152-221 umdiam.,

peripheral paraphyses 22.5-100 um long, 5-12.5 um wide, curved, cylindrical,

somewhat inflated at base, tapering towards the acuminate apex, aseptate,

chestnut brown, smooth; inner paraphyses, numerous, filiform to cylindrical,

straight, 25-47.5 x 2.5-5 um, aseptate, hyaline, smooth. Urediniospores

globose to broadly ellipsoid, 30-37.5 x 22.5-30 um, aseptate, wall bilaminate,

the outer layer 2-2.5 um thick, pale yellow, echinulate, inner wall up to 2.5

um thick, yellow, germ pores, two, equatorial. Teliospores pedicellate, ellipsoid,

47.5-55 x 31.25-37.5 um, rounded at both ends, one-septate, not constricted at

the septum, wall 2.5-4.5 um thick, chestnut brown, echinulate, two-pored, one

apical pore in upper cell and one pore next to septum in the lower cell; pedicels

attached basally or slightly obliquely, cylindrical, 50-62.5 x 5-7.5 um, hyaline,

persistent, with four or five conspicuous branched lateral appendages formed

submedianly.

ADDITIONAL MATERIAL EXAMINED: BRAZIL. Minas GERAIS, 25 Jul 2011, MSilva &

OLPereira (VIC 31757).

CoMMENTs: Two rust fungi have previously been described in association

with M. unguis-cati: Uropyxis rickiana Magnus [=U. reticulata Cummins]

and Prospodium stizophylli H.S. Jacks. & Holw. (Cummins 1939; Carvalho Jr.

et al. 2002, Hernandez & Hennen 2003). Uropyxis rickiana is known only on

M. unguis-cati from Argentina and Brazil (Hernandez & Hennen 2003, Hennen

et al. 2005) and P. stizophylli on M. unguis-cati and Stizophyllum perforatum

192 ... Silva, Barreto & Pereira

PLATE 7. Prospodium macfadyenae on Macfadyena unguis-cati: A. Rust symptoms on infected plant;

B. detail on adaxial leaf surface; c. detail on abaxial leaf surfaces.

(Cham.) Miers from Brazil (Carvalho Jr. et al. 2002). Uropyxis species differ

from Prospodium species by having two pores on each teliospore cell instead

of only one. The species collected during the present study clearly belongs to

Prospodium. Prospodium macfadyenae differs from P. stizophylli by having

Fungal pathogens of Macfadyena unguis-cati (Brazil) ... 193

PLATE 8. Prospodium macfadyenae (VIC 30478): a. type 7 spermogonium; B. basket-like sori

(squash-mount); c. echinulate urediniospores intermixed with pedicellate 2-celled teliospores.

Scanning electron micrograph (SEM) of spores: D. surface view of echinulate urediniospores;

E. mature teliospores (note branched appendages on pedicels - arrowed). Bars: a = 10 um; B = 50

um; C = 30 um; D = 10 um; E = 20 um.

longer, broader teliospores and the absence of mesospores. Prospodium

stizophylli also does not have basket-like sori, does not produce uredinia,

and has a pseudoreticulate ornamentation on its teliospores. Prospodium

macfadyenae is morphologically most similar to P perornatum Syd. (Cummins

1940, Hennen et al 2005), known only on Tabebuia spp. from Argentina and

Mexico (Hernandez & Hennen 2003), but differs from the latter by having larger

urediniospores, longer teliospores that are not constricted at the septum, and

the presence of paraphyses within the sori in addition to thick walled incurved

peripheral paraphyses.

Prospodium macfadyenae causes significant damage to the foliage of

M. unguis-cati (PLATE 7), often leading to significant defoliation of its host,

and belongs to a group of fungi that includes several highly successful classical

biological control agents of weeds. Rust fungi generally have a high level of host-

194 ... Silva, Barreto & Pereira

specificity (and thus fit well into the safety requirements needed for classical

introductions). Therefore, we regard the prospects for its use in biocontrol of

M. unguis-cati as very good. Although U. rickiana has not been collected during

the surveys in southern and southeastern Brazil, it is likely that this species also

has potential as a biological control agent in warmer equatorial areas where

M. unguis-cati occurs as an invasive plant. It has the same favorable features

described for P. macfadyenae and causes a severe disease on its host producing

conspicuous galls on stems as illustrated in Hernandez & Hennen (2003).

Ideally, further investigations involving additional surveys and a more detailed

evaluation of the potential biocontrol agents already uncovered are needed.

These could yield additional candidates for use in classical biological control

of M. unguis-cati and help pave the way towards introducing Pseudocercospora

unguis-cati, Passalora macfadyenae and Prospodium macfadyenae into exotic

situations where invasions by cat’s claws are under way, contributing to the

mitigation of the environmental damage caused by this important invasive

weed.

Acknowledgments

The authors wish to thank Alan Wood (ARC Plant Protect Res Inst, Stellenbosch,

South Africa) and Harry Evans (CABI Europe - UK) for reviewing the manuscript.

This study was performed with financial support from CNPq (Conselho Nacional

do Desenvolvimento Cientifico e Tecnolégico), FAPEMIG (Fundacao de Amparo a

pesquisa do Estado de Minas Gerais) and CAPES (Coordenacao de Aperfeigoamento

de Pessoal de Nivel Superior). The authors acknowledge the Nucleo de Microscopia e

Microanalise of the Universidade de Federal Vicosa for the use of its facilities and wish

to thank C.A. Vanetti for technical support.

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MY COTAXON

http://dx.doi.org/10.5248/119.197

Volume 119, pp. 197-200 January-March 2012

Typification of Lecanora expallens and

L. expallens var. conizaea (Lecanorales, Ascomycota)

MARTIN KUKWA

Department of Plant Taxonomy and Nature Conservation, University of Gdansk,

Al. Legionéw 9, PL-80-441 Gdansk, Poland

CORRESPONDENCE TO: dokmak@ug.edu.pl

ABSTRACT — Lectotypes are selected for Lecanora expallens and L. expallens var. conizaea

from the collection in H-ACH. Characteristic of the species based on the type collection is

provided.

Key worps — Lecanoraceae, lichenized fungi, lichen metabolites

Introduction

Lecanora expallens is a well known, common and widely distributed

sorediate lichen species, reported from many regions of the world (e.g. Tonsberg

1992, Pisut et al. 1996, Motiejainaité 1999, Diederich & Sérusiaux 2000, Sliwa

& Wetmore 2000, Hafellner & Turk 2001, Coppins 2002, Faltynowicz 2003,

Kurokawa 2003, Aptroot et al. 2004, Ryan et al. 2004, Santesson et al. 2004,

Galloway 2007). It was included in the taxonomical revision of the Lecanora

varia group in North America (Sliwa & Wetmore 2000) and in the recent lichen

flora of the Greater Sonoran Desert Region (Ryan et al. 2004) as well. The

species was also discussed by Printzen (2001) in relation to L. confusa Almb.,

a taxon which is supposed to be an esorediate counterpart of L. expallens and

considered by some authors as doubtfully distinct from the latter (see Sliwa &

Wetmore 2000).

During my research on sorediate crustose lichens, several sterile and fertile

specimens of L. expallens were investigated. After the examination of all

available literature it became clear that although L. expallens was a common

subject of several studies, no type specimen of the species has ever been cited

and actually the identity of the species has remained obscure. That prompted

the examination of the type collections of L. expallens, including L. expallens

var. conizaea. The aim of this paper is to present the typification of L. expallens

and L. expallens var. conizaea.

198 ... Kukwa

Material & methods

Type material from BM-ACH and H-ACH was studied, with additional specimens

from BM, KRAM and UGDA examined for comparison. Lichen substances were studied

by thin-layer chromatography (TLC) in solvents A and C according to the methods of

Culberson & Kristinsson (1970) and Orange et al. (2001).

Taxonomy

Lecanora expallens Ach., Lichenogr. Univer.: 374. 1810.

TyPE: Gallia, (on wood) Du Four 99 (H-ACH 1267, upper left hand specimen — lectotype

designated here).

= Lecanora expallens var. conizaea Ach., Lichenogr. Univer.: 374. 1810.

= Lecanora conizaea (Ach.) Nyl. ex Cromb., J. Linn. Soc., Bot. 15: 440. 1876.

Type: Anglia (on bark of pine) Turner (H-ACH 1267, right hand specimen -

lectotype designated here).

THALLUS crustose, grey green, episubstratal, entirely sorediate, without

any delimited soralia (marginal part not preserved in the collected specimen);

SOREDIA farinose; APOTHECIA lecanorine, sessile, flat to slightly convex,

numerous; Disc brown, delicately pruinose, pruina farinose; THALLINE MARGIN

well developed, persistent, raised above the disc, sorediate, concolorous

with the thallus, cortex not seen; AMPHITHECIUM with two different types of

crystals, one present in the outer part, yellowish in polarized light, K soluble

and N insoluble, the other in the inner part, more whitish in polarized light,

K insoluble and N soluble; PARATHECIUM well visible, up to 40 um wide,

with substance (amorphic?) yellow in polarized light, K soluble, N insoluble;

EPITHECIUM brownish, granulose, granules coarse, yellow in polarized light,

K soluble and N mostly insoluble (or slowly soluble); PARAPHYSEs simple, up to

2 um wide, not capitate; ASCOSPORES 8 in ascus, non-septate, hyaline, narrowly

(9-14 x 4-5 um) or broadly (11 x 6-6.5 um) ellipsoid.

CHEMISTRY — Usnic acid (+), zeorin (+), thiophanic acid (+), xanthone

similar to arthothelin (+), ‘expallens unknown’ sensu Tonsberg (1992) and

trace of unidentified xanthone (+, both only in the lectotype of L. expallens

var. conizaea) were detected in the studied type specimens. That agrees with

the chemistry reported by Tonsberg (1992), Sliwa & Wetmore (2000), Printzen

(2001) and Ryan et al. (2004).

ComMENTs — According to Karnefelt & Thell (2007), the main part of Acharius

collection is placed at the University of Helsinki as H-ACH. Less comprehensive

(but still large) collections are also present in the Museum of Evolution in

Uppsala (UPS-ACH), the Natural History Museum in Stockholm (S), and the

Natural History Museum, London (BM-ACH) (see also Vondrak & Vitikainen

2008). Specimens of L. expallens were found only in H and BM.

The collection in H-ACH contains five specimens, all associated at least

with the country name and some also with further information. Two of them

Lecanora species lectotypified ... 199

correspond to the protologue in Acharius (1810), therefore the collection in

H-ACH is suitable for the typification. In BM-Ach the specimen consists of

several pieces of wood covered with thallus of L. expallens, but no details of

locality and collector are provided.

Acharius (1810) cited “Habitat ad truncos Salicinos Galliae. Du Four.

Persoon. Both those collections are present in H-ACH; however, only Du

Four’s specimen belongs to L. expallens in the currently accepted sense, and

that one is selected in this paper as a lectotype to preserve the current concept

of the species. The second specimen represents Pertusaria coccodes (Ach.) Nyl.

(norstictic and connorstictic acids were detected by TLC). Both specimens

have also been examined and annotated by O. Almborn in 1963, but his results

have never been published (O. Vitikainen, pers. information).

Acharius (1810) cited three specimens of L. expallens var. conizaea: “Habitat

in cortice Pini ac ligno Quercus vetusto Angliae, Turner; Helvetiae, Schleicher.’

In H-ACH only one specimen annotated as L. expallens var. conizaea exists; it

is on piece of bark of pine from ‘Anglia and is here selected as lectotype. This

name (as L. conizaea) was often misapplied for L. conizaeoides Nyl. ex Cromb.

and L. strobilina (Spreng.) Kieff. (Laundon 1976, 2003). However, as the type

corresponds morphologically and chemically with the lectotype of L. expallens,

L. expallens var. conizaea is confirmed as a synonym of L. expallens, which was

already mentioned by Laundon (2003).

Acknowledgments

Dr Christian Printzen (Frankfurt am Main) and Dr Lucyna Sliwa (Krakéw) are

warmly thanked for peer-reviewing the paper. I am also very grateful to Dr Orvo

Vitikainen (Helsinki) for making the type collection available for the study, suggestions

how to present the results and the discussion on the typification, Dr Scott LaGreca

(formerly London) and Dr Anders Nordin (Uppsala) for the hospitality during the visits

to BM and UPS. This work was supported by the SYNTHESYS grant GB-TAF-1013.

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200 ... Kukwa

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5: 201-269,

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MY COTAXON

http://dx.doi.org/10.5248/119.201

Volume 119, pp. 201-216 January-March 2012

The morphology of Ganoderma species

with a laccate surface

MABEL GISELA TORRES-TORRES?”* & LAURA GUZMAN-DAVALOS?

‘Universidad Tecnologica del Chocd, Ciudadela Medrano, Quibd6, Chocd, Colombia

?Departamento de Botanica y Zoologia, Universidad de Guadalajara

Apartado postal 1-139, Zapopan, Jal., 45101, Mexico

*CORRESPONDENCE TO: magitoto@yahoo.com

ABSTRACT — A discussion on the importance of morphological features in the taxonomy of

Ganoderma subg. Ganoderma is presented. More than 500 specimens of 37 species, including

33 types, were checked: G. argillaceum, G. atkinsonii, G. boninense, G. capense, G. chonoides,

G. colossus, G. concinnum, G. corrugatum, G. curtisii, G. dorsale, G. elegantum, G. longistipitatum,

G. mexicanum, G. multicornum, G. multiplicatum, G. nevadense, G. nitidum, G. oerstedii,

G. orbiforme, G. oregonense, G. parvulum, G. perturbatum, G. perzonatum, G. praelongum,

G. pulverulentum, G. ravenelii, G. resinaceum, G. sequoiae, G. sessile, G. sessiliforme,

G. simulans, G. stipitatum, G. subfornicatum, G. subincrustatum, G. vivianimercedianum,

G. weberianum, and G. zonatum. Color of the context, resinous deposits, structure of the

basidiospores, and protuberances of the pileipellis cells are among the most important

features for characterization of the species.

Key worps — Basidiomycota, Ganodermataceae

Introduction

Ganoderma P. Karst. is characterized by its double-walled, colored

basidiospores with a truncate or subacute apex and ornamented exosporium.

The genus includes species with both dull [subg. Elfvingia (P. Karst.) Imazekil]

and laccate pilei (subg. Ganoderma). The pileipellis has been differently

characterized by different authors: (1) Furtado (1965a) refers to a derm of the

palisadoderm type in Elfvingia and layer of cells ina hymeniderm in Ganoderma,

while Steyaert (1980) refers to a trichoderm in Elfvingia and “hymenioderm”

in subg. Ganoderma. Clémengon (2004) used “crustohymeniderm” for

subg. Ganoderma, so following his definition, the Elfvingia pileipellis is a

crustotrichoderm, because it is also incrusted with a resinous matter. Although

phylogenetic analyses (Moncalvo 2000, Hong & Jung 2004) support the

202 ... Torres-Torres & Guzman-Davalos

monophyly of Ganoderma and its classification in two subgenera, there are

many problems in the resolution below subgeneric level.

The micromorphological structures of some species or groups in Ganoderma

have been studied by, among others, Patouillard (1889), Murrill (1902), Haddow

(1931), Heim (1962), Furtado (1965a,b, 1967), Steyaert (1967a,b, 1972, 1975,

1977), Pegler & Young (1973), Corner (1983), and Adaskaveg & Gilbertson

(1988). More recently, Gottlieb & Wright (1999) made a detailed study of the

basidiospores under the Scanning Electron Microscope (SEM) in an attempt

to solve some of the inter-specific problems. However, these studies have been

restricted to a rather limited number of species and few used the pileipellis cells

as a critical character in their species concepts.

Through the study of more than 500 specimens (including 33 types)

mainly from Africa, Brazil, Costa Rica, Mexico, and USA, we analyzed the

morphological variation of structures in Ganoderma subg. Ganoderma with

the following aims: 1) to study in detail structures currently and potentially

useful in taxonomy and phylogenetic analyses and 2) to probe the utility of

light microscopy in defining basidiospore pillars.

Materials & methods

Specimens examined

More than 500 specimens from the herbaria BPI, BR, ENCB, FH, H, IBUG (including

collections by the first author), INB, K, NY, O, PC, SP, UPS, and XAL were examined

(acronyms follow Holmgren et al. 1990). These specimens comprised 37 species, which

are listed in TaBLE 1, which includes one representative specimen for each species.

Macro and micromorphological observations and identification

The following macromorphological characters were described for each basidiome:

length, width, thickness, adhesion to the substrate, general shape, consistency, weight;

pileus shape, surface consistency, color; context thickness, stratification, color, resinous

deposits; pore form, number per mm, consistency, color; tube length, stratification, color.

Macrochemical reactions used 10% KOH. In addition to the traditional microcharacters,

descriptions include: basidiospore apex, disposition and size of basidiospore pillars,

and pileipellis cell size, shape, and incrustations. Color references follow Kornerup

& Wanscher (1963). Microscopical observations were made from basidioma tissues

mounted in 10% KOH and Melzer’s reagent, with Congo red, phloxine, and cotton blue

additionally used. Basidiospore shape is determined from the Q ratio (1/b; Bas 1969:

320-321) of 20 randomly selected basidiospores. Only mature pileipellis cells were

described. Microscopical observations were made using a 100x oil-immersion objective

in Zeiss K7 and Zeiss Axioskop 40 microscopes. The inter-walled basidiospore pillars

were measured using Axio Vision 4 software with the Zeiss Axioskop 40 microscope.

Most morphological terms follow Vellinga (1998), Furtado (1965a), and Clémencgon

(2004). The pileipellis description follows the concepts of Clemencon (2004).

Morphology of laccate ganodermas ... 203

TABLE 1. Species of Ganoderma subg. Ganoderma with representative collections.

SPECIES

G. argillaceum Murrill

G. atkinsonii H. Jahn et al.

G. boninense Pat.

G. capense (Lloyd) Teng

G. chonoides Steyaert

G. colossus (Fr.) C.F. Baker

G. concinnum Ryvarden

G. corrugatum Steyaert

G. curtisii (Berk.) Murrill

G. dorsale (Lloyd) Torrend

G. elegantum Ryvarden

G. longistipitatum Ryvarden

G. mexicanum Pat.

G. multicornum Ryvarden

G. multiplicatum (Mont.) Pat.

G. nevadense Murrill

G. nitidum Murrill

G. oerstedii (Fr.) Murrill

G. orbiforme (Fr.) Ryvarden

G. oregonense Murrill

G. parvulum Murrill

G. perturbatum (Lloyd) Torrend

G. perzonatum Murrill

G. praelongum Murrill

G. pulverulentum Murrill

G. ravenelii Steyaert

G. resinaceum Boud.

G. sequoiae Murrill

G. sessile Murrill

G. sessiliforme Murrill

G. simulans Wakef.

G. stipitatum (Murrill) Murrill

G. subfornicatum Murrill

G. subincrustatum Murrill

G. vivianimercedianum M.G. Torres

SPECIMEN

ES. Earle 658 (lectotype, NY)

without data (O)

C. Wright s.n. (lectotype, K)

P. van der Bijl s.n. (lectotype, NY)

without data (holotype, BR)

A.S. Oersted s.n. (lectotype, UPS)

L. Ryvarden 16840 (holotype, O)

R.L. Steyaert 45061 (holotype, BR)

P.O. Schallert s.n. (F1119720)

RJ. Rick s.n. (holotype, BPI)

Ryvarden 44573 (holotype, O)

L. Ryvarden 40588 (holotype, O)

without data (lectotype, FH)

G.J. Samuels s.n. (holotype, O)

R. Maziero s.n. (SP250605)

C.E Baker 1489 (lectotype, NY)

P. Wilson 607 (lectotype, NY)

A.S. Oersted s.n. (neotype, UPS)

A. Afzelius s.n. (holotype, UPS)

J.E. Kirkwood s.n. (lectotype, NY)

C.L. Smith s.n. (Lectotype, NY)

R. Rick s.n. (holotype, BPI)

ES. Earle 309 (lectotype, UPS)

ES. Earle & W.A. Murrill 536 (lectotype, NY)

W.E. Broadway s.n. (lectotype, NY)

H.W. Ravenel 2936 (holotype, K)

J.L. Boudier s.n. (holotype, PC)

A. Eastwood 40 (lectotype, NY)

without data (lectotype, NY)

E. & WA. Murrill 392 (lectotype, NY)

T.D. Maitland 556 (isotype, O)

C.L. Smith s.n. (Lectotype, NY)

MLE. Peck s.n. (lectotype, NY)

ES. Earle 176 (lectotype, NY)

E. Bastidas-Varela s.n. (holotype, ENCB)

G. weberianum (Bres. & Henn. ex Sacc.) Steyaert A. Milanez & D. Altimari s.n. (SP61099)

G. zonatum Murrill L.M. Underwood s.n. (lectotype, NY)

Specimens were generally determined after comparing with the type or types of

related species, but the keys of Bazzalo & Wright (1982) and Ryvarden (2004) and the

descriptions of Steyaert (1972) and Corner (1983) were also used.

Results

The features proposed here are discussed below and are being used to

describe and key Ganoderma species from Brazil and Mexico (Torres- Torres et

al., papers in preparation).

204 ... Torres-Torres & Guzman-Davalos

Macromorphological features

Basidiome shape and substrate attachment are of rather restricted value for

taxon delimitation; however, basidiome consistency, weight, and thickness as

well as the presence and attachment of the stipe proved, in some cases, to be

valuable for identification. Ganoderma atkinsonii, G. colossus, G. nevadense,

and G. oregonense have light, spongy basidiomata, while G. zonatum and

G. pulverulentum have light, woody-corky basidiomata. Ganoderma mexicanum,

G. sessile, G. sessiliforme, and G. subincrustatum have relatively light, woody

basidiomata, while Ganoderma oerstedii has heavy, woody basidiomata.

Ganoderma colossus, G. oregonense, G. oerstedii, G. sessile, and G. zonatum

are generally robust, while G. mexicanum, G. parvulum, G. sessiliforme, and

G. weberianum have slender basidiomata.

The more important features of the pileus context for diagnosing species

are: (1) consistency, (2) structure, and (3) presence of resinous deposits. Only

three types of context consistency are recognized. Two — soft-spongy and

fibrous — are easy to distinguish, but there is an intermediate state between soft

and fibrous that is difficult to identify. In this study, the third type was termed

“fibrous-spongy. Ganoderma colossus and G. oregonense have soft-spongy

context, G. sessiliforme has fibrous context, and G. resinaceum has fibrous-

spongy context.

The context structure is classified as (1) duplex, (2) not fully homogeneous,

or (3) homogeneous. In the duplex context there is an abrupt change, with

two separate, contrasting colors, the upper generally light-colored and the

lower generally darker close to the tubes, although shades may vary to some

extent. This context type is present in G. concinnum, G. curtisii, G. dorsale, and

G. sessile (Fic. 1). Not fully homogeneous is used when there is an evident color

difference between the upper and lower parts but without abrupt color changes,

like in G. oerstedii (Fic. 2), G. perturbatum, G. perzonatum, G. weberianum

(Fic. 3), while a homogeneous context has only one color, as in G. parvulum

and G. resinaceum (Fics. 4-5). Contexts with an occasional very thin darker

line just above the tubes that is absent in some specimens are considered as

homogeneous. The context can contain resin-like deposits (hard and brittle,

dull or shiny) that often they form continuous bands (here “resinous bands,”

see G. oerstedii and G. parvulum in Fics. 2, 4) or discrete bodies (here “resinous

incrustations,” see G. weberianum, Fic. 3).

Some authors (Patouillard 1889, Moncalvo 2000, Seo & Kirk 2000) have

dismissed the stipe as a character of no taxonomic value, because some sessile

species can develop a stipe in the laboratory resulting from varying oxygen

conditions. Nevertheless, the stipe is a useful character, with a “true” stipe

defined as one > 5 cm and with tissues differentiated from the pileus context.

Some species develop a generally short (<5 cm) pseudostipe that is merely a

Morphology of laccate ganodermas ... 205

Fics. 1-3. Ganoderma macromorphology. 1. Duplex context (G. sessile). 2. Not completely

homogeneous context (G. oerstedii); continuous bands of the resin-like deposits (arrowed).

3. Not completely homogeneous context (G. weberianum); discrete bodies of the resin-like deposits

(arrowed). Bar = 2 cm.

continuation of the pileus and which Steyaert (1980) termed a “pedicel” and

we call a “substipe.” Species such as G. curtisii, G. concinnum, G. dorsale,

and G. perturbatum always present a true stipe, while G. pulverulentum and

G. weberianum can develop a substipe dependent upon environmental and

nutritional conditions.

206 ... Torres-Torres & Guzman-Davalos

Fics. 4-5. Ganoderma macromorphology. 4. Homogenous context (G. parvulum); continuous

bands of the resin-like deposits (arrowed). 5. Homogeneous context (G. resinaceum); note the

absence of resin-like deposits. Bar = 2 cm.

Micromorphological features

Basipiospores. Following Clémencon’s (2004) terminology, all Ganoderma

basidiospores have a thin, smooth or rugose, hyaline to reddish-yellow

perisporium; an ornamented colored exosporium with pillars, and a thick

reddish-brown or yellowish-brown endosporium. Using the Q ratio categories

(Bas 1969), most species have ellipsoid to elongate (= oblong) basidiospores,

while G. concinnum, G. dorsale and G. perturbatum have distinctively widely

ellipsoid to ellipsoid basidiospores and G. ravenelii and G. zonatum have oblong

to cylindrical basidiospores. Ganoderma basidiospores have a cap in the apex

(Fics. 6-7) that is obtuse and hyaline; after drying, the cap generally collapses

and breaks, making the basidiospore apex truncate (Fics. 7-8). However, in

some species where the cap is absent or very small, the basidiospores have

a subacute apex, as in G. atkinsonii, G. colossus, G. concinnum, G. dorsale,

G. longistipitatum, G. oregonense, and G. perturbatum (Fics. 9-11).

The basidiospore inter-walled pillars are thin to thick; thin (<0.3 um)

in G. mexicanum, G. perzonatum, G. praelongum, and G. zonatum (Fics.

Morphology of laccate ganodermas ... 207

Figs. 6-11. Ganoderma micromorphology. 6. Basidiospores with cap (G. multiplicatum).

7. Basidiospore (right) with cap and (left) with truncate apex (G. sessiliforme). 8. Basidiospores

with truncate apex (G. chonoides). 9-11. Basidiospores with subacute apex (9. G. atkinsonii;

10. G. concinnum; 11. G. oregonense).

12-14), intermediately thick (0.3-0.5 um) in G. corrugatum, G. curtisii,

G. oerstedii, G. pulverulentum, G. resinaceum, G. sessiliforme, G. subincrustatum,

and G. weberianum (Fics. 15-17), and thick (0.5-0.7 um) in G. colossus,

G. longistipitatum, G. oregonense, and G. perturbatum (Fics. 18-20). The pillars

can be free, sub-free, partially anastomosed, and reticulate (= anastomosed).

“Free” pillars appear as dots on the basidiospore surface (e.g. G. perzonatum,

FIG. 21), while “sub-free” are free dots mixed with short anastomosed to shortly

elongated structures (e.g. G. elegantum, FIG. 22). “Partially anastomosed”

is used when more than two pillars are grown together to form an irregular

surface (e.g. G. concinnum, FIG. 23), and “reticulate” is when the ornamentation

forms an almost complete net (e.g. G. colossus, FIG. 24).

STRUCTURE OF THE PILEAR COVER. The pilear surface or pileipellis is formed by

a cell layer similar to a hymeniderm, although in the majority of the species the

cells do not originate from the same level as in the true hymeniderm. Clémen¢gon

(2004) defined the Ganoderma type of hymeniderm as a crustohymeniderm,

which is characterized by thick-walled cells apically covered by a resinous

208 ... Torres-Torres & Guzman-Davalos

12 a 14

Fics. 12-24. Ganoderma micromorphology. 12-14. Basidiospores with thin inter-walled pillars

(12. G. perzonatum; 13. G. praelongum; 14. G. zonatum). 15-17. Basidiospores with intermediate

thick inter-walled pillars (15. G. pulverulentum; 16. G. weberianum; 17. G. corrugatum).

18-20. Basidiospores with thick inter-walled pillars (18. G. colossus; 19. G. longistipitatum;

20. G. perturbatum). 21-25. Disposition of the pillars in the basidiospores: 21. Free (G. perzonatum);

22. Subfree (G. elegantum); 23. Partially anastomosed (G. concinnum); 24. Reticulate (G. colossus).

Morphology of laccate ganodermas ... 209

matter. Cell sizes must be interpreted with care because they vary not only

between species but also in the same specimen according to where the section

has been made. Most species have club-like apical cells although there are

species with branched or knobby cells besides the club shaped cells (Fic. 25).

The shape of the crustohymeniderm cells can be: (1) cylindrical (G. per-

zonatum), (2) almost cylindrical to narrowly clavate (G. weberianum), (3) widely

clavate (G. perturbatum), and (4) clavate (most species). The cell wall is thick

for all except G. colossus, where it is thin. A thick wall may be multi-stratified,

related to the maturity state of the basidiomata; however, some species, like

G. perturbatum, have an evidently multi-stratified wall. Some young pileipellis

cells present secondary septa, more evident in the cells of the pileus periphery;

furthermore, these peripheral cells tend to be lighter and shorter. Incrustations

or granulations can also be present in the pileipellis. The incrustations are easily

dissolved in KOH [e.g. G. stipitatum, Fic. 27], while the granulations remain,

because they are not alkali-soluble (e.g. G. perzonatum, FIG. 26).

In some species the pileipellis cells have a transition from smooth club-like

to antler-like cells. Protuberances and branches (as used in the descriptions

below) differ in that “protuberances” are short projections or prolongations of

the hyphal wall (generally< 8 um long, located laterally or at the apex) that lack

a lumen whereas “branches” may be short or large, apical or lateral but always

possess a lumen. There are shape series in which the cell changes progressively,

but a predominant shape is always observed (Fic. 25). Three large groups of

crustohymeniderm cells might be found: entire (Fics. 25a, 26-28), lobulate-

branched (Fias. 25b, 29-30), and branched (Fic. 31).

1) “Entire” crustohymeniderm cells are cylindrical, cylindrical with slightly

wide apex, cylindrical to narrowly clavate, cylindrical with subcapitate apex, or

narrowly clavate to widely clavate, generally lacking protuberances or branches,

very occasionally with at most two, generally lateral protuberances, e.g. in

G. perturbatum, G. perzonatum, G. stipitatum, and G. sessiliforme (Fics. 25a,

26-28).

2) “Lobulate-branched” crustohymeniderm cells may be subdivided into:

a) clavate cells, with 1-3 round short thick protuberances and/or branches,

but preserving the clavate shape (e.g., G. nitidum; G. zonatum, Fic. 25b); b)

cells with no or 1-2 thick long branches and < 7 short thick protuberances,

where the clavate shape is generally lost (e.g., G. oerstedii; G. subincrustatum,

Fics. 25c, 31; G. multiplicatum, Fics. 29-30).

3) “Branched” crustohymeniderm cells, with two types that generally lose

the clavate shape: a) with 2-3 thin long branches with small protuberances (e.g.,

G. orbiforme, Fic. 25d) and b) the most striking, cells with many antler-like

branches and small protuberances (e.g., G. multicornum, Fic. 25e).

210 ... Torres-Torres & Guzman-Davalos

Fic. 25. Ganoderma crustohymeniderm cells. a. Entire (G. perturbatum, G. sessile). b. Lobulate-

branched (G. nitidum, G. zonatum). c. Lobulate-branched with up to seven short and thick

protuberances and 0 or 1-2 thick and long branches (G. oerstedii, G. subincrustatum). d. Branched

(G. orbiforme). e. Antler-like (G. multicornum).

Morphology of laccate ganodermas ... 211

HYPHAL sysTEM. In all species the hyphal system is di-trimitic, where generative

and binding hyphae are generally difficult to observe. All generative hyphae have

clamped septa and are hyaline to yellowish and (in almost all species) branched.

Skeletal hyphae are of the arboriform type, thick-walled to solid, yellowish to

yellowish-brown, apically branched; thin in the crustohymeniderm, wider in

the context. Binding hyphae are generally thinner than skeletal hyphae, hyaline

to yellowish-brown.

Cystip1A. Vellinga (1998) defined them as sterile, differentiated, terminal

elements in the hymenium. According with this definition G. colossus,

G. oregonense, G. perzonatum, G. pulverulentum, G. sequoiae, and G. stipitatum

have cystidia (Fics. 32-34), but they are difficult to observe.

Discussion

It is clear that species concepts vary in Ganoderma according to the author.

Historically, Ganoderma taxa have been treated as species with great plasticity

(e.g. G. resinaceum by Ryvarden, 2004) or (contrariwise) very narrowly

(Steyaert 1972, 1980). Bazzalo & Wright (1982) and other authors, especially

when synonymizing various different species under the same name, considered

context color of little value. On the other hand, Steyaert (1967a) regarded the

resin-like deposits as unimportant, mainly in the species of subg. Elfvingia.

However, we found that these deposits together with context color are important

in separating taxa. The resin-like deposit (termed “melanoid deposits” by

Gottlieb & Wright, 1999 and melanoid substance by Steyaert, 1980) was called

by Ryvarden (2000) “resinous bands,” the term adopted here but more narrowly

so as to differentiate actual resinous bands from more generalized resinous

incrustations (small, fragmented deposits that do not form bands).

Ganoderma basidiospores have been traditionally described as double

walled with a thin exosporium and thick endosporium, echinulate or with

inter-walled pillars, or struts, or sinuous ridges, with truncate apex (Patouillard

1889, Murrill 1902, Haddow 1931, Overholts 1953, Corner 1983, Tham 1998,

Ryvarden 2000, 2004 among others). The terms exosporium, episporium, and

endosporium have been differently applied for the authors mentioned above

if Cleémencon (2004) is followed. Steyaert (1977) described the basidiospores

with epi-endosporium and echinulate ecto-perisporium, sometimes the ecto-

perisporium crestate besides echinulate. Bazzalo & Wright (1982), who used

“perisporium” and “endosporium,’ considered the ornamentation as derived

from the endosporium, in contrast to Clémenc¢on (2004), who determined that

the exosporium forms the ornamentation. As defined by Kirk et al. (2008),

echinulate (diminutive of echinate: “having sharply pointed spines”) does

not apply to the inner ornamentation seen in Ganoderma, so we prefer to

describe it as pillars. Bazzalo & Wright (1982) and Gottlieb & Wright (1999),

212 ... Torres-Torres & Guzman-Davalos

+” 7

: 4} 4 .

‘ } ry

é { - ; ba : |

e ‘ ’ : ag

aw | . ay »

, ra 7

| 7 qe - 4 5 '

Fics. 26-34. Ganoderma crustohymeniderm cells (26-31): 26. Cylindrical cells with granulations

(arrow) of G. perzonatum. 27. Cylindrical cells with incrustation (arrow) of G. stipitatum. 28.

Clavi-form cells without granulations of G. sessiliforme. 29. Irregular cell with apical branches

and protuberance of G. multiplicatum. 30. Lobulate-branched cells with many protuberances of

G. oerstedii. 31. Lobulate-branched cell with a large lateral branch and a smaller apical branch of

G. multiplicatum. 32-34, Hymenial cystidia (32. G. sequoiae; 33. G. perzonatum; 34. G. stipitatum).

Morphology of laccate ganodermas ... 213

who observed the basidiospore pillars under SEM, proposed them as useful in

determining species. We emphasize their usefulness by noting that not only are

the pillars seen through SEM, pillar characters like size and disposition are also

visible through light microscopy.

The basidiospore cap has been variously named: “bouchon perisporique”

(Heim 1962), apical papilla (Furtado 1962), umbo (Corner 1983), and cap

(Tham 1998). As mentioned above, when the cap collapses and breaks the

basidiospore apex looks truncate; in few cases appears subacute, because the

cap is very small or even absent. The term subacute, described by Steyaert

(1972) for basidiospores with unchanged apex or apex unaltered at maturity,

was adopted by Corner (1983), although this character has not been used

systematically. We consider that it is an important feature in diagnosing taxa;

for example, it supports G. perturbatum as independent from G. resinaceum and

related to G. dorsale. Ganoderma concinnum, G. dorsale, G. longistipitatum, and

G. perturbatum form a distinctive group of species by their stipitate basidiomata

and subacute basidiospore apices.

Basidiospore shape is also an important diagnostic character. Oblong to

cylindrical spores differentiate G. ravenelii and G. zonatum from morphologi-

cally similar species such as the ellipsoid spored G. curtisii and G. orbiforme.

Pileus surface is a complex structure in which the crustohymeniderm cells,

hyphae and resinous-like substances are immersed. Many discussions on its

taxonomic importance in Ganoderma have been made (e.g. Furtado 1965a,

Gottlieb & Wright 1999, Steyaert 1980); however, its current use is limited,

perhaps caused by a lack in precise terminology. Much of the complexity is

caused by the great variability in the branching pattern of the cells in the same

basidioma, creating confusion and difficulty to find a collective term. Torres-

Torres & Guzman-Davalos (2005) realized that pileipellis cells vary greatly

in size and shape depending on the site sampled and on the basidiome age.

For this reason, tissues should be cut away from the basidiome edge, and only

mature cells must be compared.

Imazeki (1939) and Steyaert (1980) classified genera based on pileipellis

(dermis) type. Bazzalo & Wright (1982) had regarded pileipellis types described

by preceding authors as valuable for identifying large groups but not useful for

delimiting species. Subsequently, Gottlieb & Wright (1999) considered three

general dermis types based partly on Steyaert (1980): a) “hymeniodermis vera,”

b) “hymeniodermis formed by diverticulate hyphae... with similar endings,

though wider through the apex, and with lateral branches, acanthophyses-

like,” and c) “hymeniodermis spheroid-pedunculate... composed of claviform

filaments, with or without lateral diverticules, but always with emerging

spheroid-pedunculate or capitate knobs at the apex.’ Their acceptance of these

dermis types and basidiospore ornamentation as useful for delimiting taxa has

214 ... Torres-Torres & Guzman-Davalos

been confirmed in the present investigation. However, we found it difficult

to assign species to one of these pileipellis types, we prefer to describe cells

according to shape and the number of branches and protuberances. ‘There is

one species complex — G. boninense, G. multiplicatum, G. orbiforme, G. sub-

fornicatum and G. oerstedii, all with yellowish-brown to reddish-brown context

and pileipellis cells with protuberances and/or branches — in which species

are differentiated base on basidiospore features and pileipellis cell shape and

branch numbers.

Karsten (1881) did not mention cystidia in his original description of

Ganoderma, and many mycologists accepted his non-cystidiate concept

(Haddow 1931, Heim 1962, Furtado 1965b, 1967, Steyaert 1967b, 1972, Bazzalo

& Wright 1982, Gilbertson & Ryvarden 1986, Ryvarden 1991, 2000). Although

Corner (1983) noted that Demelius (1919) referred to acute ‘cystidioles’ in

G. applanatum (Pers.) Pat. (subg. Elfvingia) and G. lucidum (Curtis) P. Karst.,

we follow Vellinga (1998) and regard them as true cystidia that are present in

at least five species.

In conclusion, we consider that additional important features for

characterizing Ganoderma species that can be directly observed or under light

microscopy should include context color, structure, and resinous deposits;

basidiospore pillar disposition and size; and pileipellis cell shape, protuberances,

branches, and granulations.

Acknowledgments

We are grateful to the curators of BPI, COL, ENCB, FH, H, INBIO, K, NY, O, PC,

SP, UPS, and XAL for the specimen loans that made possible this study. Thanks are

due to Universidad de Guadalajara (projects P3E 50052, 108721, PROCOFIN 7388401),

CONACYT (project 42957), and PROMEP (103.5/03/2580). The first author thanks

Oslo University for a grant to visit O herbarium, Red Latinoamericana de Botanica

for a grant to visit SP herbarium (RLB-05-P5), COLCIENCIAS and Universidad

Tecnoldgica del Chocé for economic help for her Doctoral studies in the Universidad

de Guadalajara, Mexico. The authors thank Tatiana Gibertoni (Universidad Federal de

Pernambuco, Brazil) and Adriana M. Gugliotta (Instituto de Botanica, Sao Paulo, Brazil)

for the critical reviews of the manuscript.

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MY COTAXON

http://dx.doi.org/10.5248/119.217

Volume 119, pp. 217-231 January-March 2012

Four new polypore species from the western United States

JOSEF VLASAK*” , JOSEF VLASAK JR. & LEIF RYVARDEN?

‘Biol. Centre of the Academy of Sciences of the Czech Republic &

*University of South Bohemia, Faculty of Science,

Branisovskd 31, CZ-370 05 Ceské Budéjovice, Czech Republic

*Biological Institute, PO. Box 1045, Blindern, N-0316 Oslo, Norway

*CORRESPONDENCE TO: vlasak@umbr.cas.cz

ABSTRACT — Four new species of polypores are described from California, Oregon, and

Washington, U.S.A. Antrodia madronae, seemingly confined to dead stems and branches

of Arbutus menziesii, is closely related to European Antrodia sandaliae growing on Arbutus

unedo. The generic position is evaluated for Ceriporiopsis pseudoplacenta, a strikingly colored

species with many unique features that seems surprisingly common in the northwestern

states. Fuscoporia palomari is distinguished from the similar EF. viticola by much larger pores

and spores. Skeletocutis subodora differs from S. odora by having thick subiculum, differently

shaped basidiospores, abundant cystidioles, and no skeletal hyphae in the context. The

nrDNA ITS region of all new species gave unique sequences in GenBank.

Key worps — Hymenochaetaceae, Phellinus, Phlebia, ribosomal DNA, taxonomy

Introduction

The forested west coast of the United States is extremely biodiverse due

to its varied topography, climate zones, and geology. The number of vascular

plant species found in California exceeds the total number from the central

and northeastern United States and adjacent Canada, an area ten times larger

(CI 2007). The high degree of plant endemism is also striking; more than 61%

of Californian plant species are found nowhere else in the world. ‘This creates

favorable conditions also for high diversity and endemism of wood-bound

poroid fungi. Nevertheless, in the North American polypore monograph by

Gilbertson & Ryvarden (1986, 1987), only 60 (of 406) species are specified

as occurring in the western states of California, Oregon, and Washington.

This indicates that the region still has a great potential for field research in

polyporology.

Few substantial contributions have been published on USA west coast

polypores since Murrill’s early papers (Murrill 1912, 1915). Overholts, who

218 ... Vlasak, Vlasak & Ryvarden

described several new species sent to him from the Pacific coast states, never

collected in the far west (Overholts 1939). The same is essentially true for

Lowe and other distinguished USA polyporologists. Gilbertson (1976, 1979),

who published some new species and important distribution data, focused his

research primarily on the dry hot regions of Arizona.

We visited various localities on the USA west coast and report here on

several polypore collections. Although the collected material was quite scarce,

we describe four new species supported by distinctive macro- and microscopic

features and unique ITS rRNA sequences.

Materials & methods

Polypores were collected in August (2001, 2003), September (2005, 2009), and April

(2010) during one-week trips to various localities in California, Oregon, and Washington

(Vlasak 2008). The specimens were dried and microscopically inspected in Melzer’s

reagent (IKI) and 5 % KOH. Similar fungi from other parts of USA and Europe were also

collected or loaned from herbaria and studied in detail as noted in ‘Specimens studied’

All collected specimens are deposited in private herbarium of the first author, with types

and some paratypes also in PRM. DNA isolation from critical samples, nr DNA ITS

region sequencing, and comparative phylogenetic analyses methods follow Vlasak &

Kout (2011). The evolutionary history was inferred using the Neighbor-Joining method

(Saitou & Nei 1987) and phylogenetic analyses were conducted in MEGA4 (Tamura et

al. 2007).

Taxonomy

Fuscoporia palomari Vlasak & Ryvarden, sp. nov. PLATES 1-2

MycoBank MB 563062

Basidioma resupinatus vel effuso-reflexus, pori facies brunnea, rotundi vel elongati, 2 pro

mm. Setae subulatae, brunnei, 40-75 x 5-8 um, basidiosporae hyalinae, leves, cylindricae,

8-11 x 2.7-3.5 um, inamyloideae, indextrinoideae.

Type: USA. California: Palomar Mt. State Park, 1500 m above see level, on oak log,

16 Apr 2010, J. Vlasak Jr. JV1004/5-J (Holotype, PRM 915982; isotype, JV 1004/5a-J,

GenBank JN592493).

ErymMo_oey: Referring to the type locality on Mt. Palomar, CA.

BASIDIOME about 7 x 5 cm, resupinate to effused-reflexed, with sharply

delimited sterile margin 0.5-1 mm wide, pilei indistinct, projecting only 2-3

mm and < 1 cm wide, upper surface tomentose, pore surface yellowish brown

with a shiny luster when viewed obliquely. Porss large, circular to angular, in

places somewhat labyrinthine, 2 per mm, with thin, soft dissepiments that are,

though, often thickened on pore edges. CONTEXT brown, fibrous, rather soft,

very thin in resupinate part but up to 2 mm thick in developing pilei; tube layer

concolorous and continuous with the context, up to 3 mm thick. CONTEXTUAL

HYPHAE of two types, some brown in KOH, thick-walled, 2-4 um in diam.,

New western polypores (U.S.A.) ... 219

~esig age”

PLATE 2: Fuscoporia palomari microscopic structures.

a: Basidiospores, b: Hymenial seta, c: Hymenium with basidia and cystidiole.

others hyaline in KOH, thin-walled, with occasional branching, simple-septate,

2-3 um in diam.; tramal hyphae similar. Seraz abundant, narrowly subulate,

220 ... Vlasdk, Vlasak & Ryvarden

thick-walled, very long, 40-75 x 5-8 um, most of them around the upper

length limit, protruding perpendicularly from the hymenium to more than a

half of their length. Basip1a clavate, simple-septate at the base, 15-25 x 5-8

um. CysTIDIOLeEs hyphoid, rarely present, with up to 25 um long and about

1.5 um broad terminal part. BAstp1osporss cylindric, straight, hyaline, thin-

walled, smooth, IKI-, 8-10(-11) x 2.7-3.5 um, few turning brown in KOH.

Causing a white rot.

COMMENTS — Only one juvenile basidiome with rudimentary developing pilei

was collected. However, it shows many unique features: very large, hyaline

cylindric spores, long setae, and large, shining pores. The hyaline basidiospores

do not change in KOH except for a small fraction where the inner contents (not

walls) turn deep brown. These usually have rather uneven walls and distorted

shapes (PLATE 2) that we consider atypical. The ITS nrRNA sequence is unique,

showing significant similarity (max 93%) in GenBank Blast search only with

Fuscoporia viticola (Schwein.) Murrill, a species with similar long setae and

cylindric (but much narrower) basidiospores. Fuscoporia sentifera (T. Hatt.)

Y.C. Dai, originally described from Japan (Hattori 1999) and recently reported

in China (Dai 2010), is similar to F palomari by resupinate to effused-reflexed

basidiocarps, long hymenial setae, and cylindric spores but differs by a hispid

pileal surface and smaller spores (5.8-7 x 2-2.5 um).

Antrodia madronae Vlasak & Ryvarden, sp. nov. PLATES 3-4

MycoBank MB 563081

Basidiomata annua, resupinata, nodulosa vel aliquando pileata, colore ligneo-pallens, pori

rotundi vel elongati et sinuosi, 2-3 pro mm. Basidiosporae hyalinae, leves, tenuitunicatae,

cylindraceae, 10-12 x 2.8-3.8 um, inamyloideae, indextrinoideae. Cariem brunneam in

ligno Arbuti menziesie producet.

Type: USA. Oregon: Oregon Caves entrance, on dead branch of pacific madrone tree, 14

Sep 2007, J. Vlasak Jr. JV0709/117-J (Holotype, PRM 899296; GenBank JN592494).

Erymotoey: Referring to the only known substrate tree, pacific madrone, Arbutus

menziesii Purrs (Ericaceae).

BASIDIOMATA annual, up to 30 x 10 cm, resupinate to effused-reflexed,

nodulose or with narrow sloping pilei developed on a decurrent pore surface,

individual pilei up to 4 cm wide, soft when fresh, upper surface ochraceous to

light brown colored, coarsely rugose, azonate, margin sharp, tomentose, pore

surface concolorous. Pores circular when immature, (1-)2(-3) per mm, on

sloping parts often sinuous and elongated; dissepiments thick. CONTEXT thin

and about the same color as the pore surface, tube layer concolorous, up to

7 mm thick. HyPHAL sysTEM dimitic, generative hyphae clamped, branched,

thin-walled or somewhat thick-walled, 2-4 mm in diam., dominating in all

parts of basidiocarp, thickened walls gelatinizing and dissolving in KOH,

New western polypores (U.S.A.) ... 221

PLATE 3: Antrodia madronae microscopic structures.

a: Basidiospores, b: Basidia, c: Skeletal hyphae.

PLATE 4: Antrodia madronae. Basidiome (JV0709/90).

skeletal hyphae thick-walled but never solid, 3-5 um in diam., dissolving in

KOH. Cystip1a none. BaAsrip1A narrowly clavate, with four sterigmata and a

basal clamp, 25-40 x 6-8 um. Basip1osporEs hyaline, thin-walled, smooth,

IKI-, cylindrical or somewhat fusiform, 10-12 x 2.8-3.8 um. Causing a brown

rot on dead branches and stems of pacific madrone.

ADDITIONAL SPECIMENS EXAMINED: Antrodia madronae - USA. CALIFORNIA:

Crescent City, Myrtle Creek, on dead stem of Arbutus menziesii, 12 Sep 2007, J. Vlasak

(JV0709/90). Antrodia sandaliae - ITALY. Sarpinta: Montarbu Forest, Sarolegne,

222 ... Vlasdk, Vlasdk & Ryvarden

Arbutus unedo, 29 Nov 2003, A. Bernicchia (HUBO 7803, GenBank JN592495); 29 Nov

2003, A. Bernicchia (HUBO 7784, ITS sequence is identical with that of HUBO 7803);

24 Nov 1999, A. Bernicchia (HUBO 7340).

COMMENTS — More material is needed to describe the extent of macroscopic

variation of this species. The description is based on only two specimens; the

type is a small, resupinate basidiocarp in the wood-crevice of a dead, broken

branch of a living tree, while the other, growing on a dead standing tree, is a

large, partly pileate basidiocarp with a strikingly rugose pileus surface but old

and heavily damaged by insects (PLATE 4). Antrodia madronae is characterized

by large cylindrical basidiospores, rather soft fruitbodies dominated by

generative hyphae, extremely soft trama that almost dissolved during the

microscopic preparation in Melzer’s reagent, large pores, and causing a brown

rot. Undoubtedly, the fungus is closely related to the Mediterranean A. sandaliae

Bernicchia & Ryvarden, also growing on an arbutus (Arbutus unedo L.) and

characterized by a white pulvinate basidiome, similar soft trama and hyphal

system, and similarly shaped but smaller basidiospores (7.5-10.0 x 1.8-3.0

um; Bernicchia & Ryvarden 2001, Gorjon & Bernicchia 2009). Although both

species have KOH-soluble thick-walled hyphae, the pores, basidia and spores of

A. madronae are distinctly larger and the basidiome is varicolored.

SEQUENCE ANALYSIS — The amplified ITS1-5.8S-ITS2 rDNA region of A. madronae

was 565b long between ITS1 and ITS4 primer sequences (White et al. 1990) and

the sequence was rather similar to that of A. sandaliae HUBO 7803 and HUBO

7784. The sequence variation between A. madronae and A. sandaliae of 2.0%

(4/199) in ITS1 and 2.4% (5/212) in ITS2 suggests a very close relationship

regardless of the pronounced macroscopical differences. Other species in

GenBank Blast Search are only distantly related, with A. albobrunnea (Romell)

Ryvarden (max. 83% sequence similarity) seemingly the closest.

Ceriporiopsis pseudoplacenta Vlasdk & Ryvarden, sp. nov. PLATES 5-8

MycoBank MB 563082

Basidiomata annua, resupinata, salmon pink pallide colore sed carnosa et cartilaginea post

siccatione, deinde rubro-brunnei colore. Pori rotundi vel angulati, 3-4 pro mm. Systema

hypharum monomiticum, hyphae hyalinae, fibulatae, ramosae, 2.0-3.0 um latae, quae

materiam oleaceam continent. Basidiosporae hyalinae, leves, ellipsoideae, 3.5-4.5(-5) x

2.2-3 WM.

Type: USA. Washington: Forks, Bogachiel State Park, 6 Aug 2003, Picea sitchensis

(Bong.) Carriére. log, J. Vlasak JV0308/68 (Holotype, PRM 899297; isotype, JV0308/

68a; GenBank JN592496, JN592497, JN592504).

ErymMo_oey: from the Latin for false placenta, based on the superficial similarity with

Postia placenta.

BASIDIOMATA annual, resupinate, up to 20 x 10 cm, soft when fresh, tough and

fleshy when dry, and soaked with resinous substance, not readily separable;

New western polypores (U.S.A.) ... 223

PLATE 5: Ceriporiopsis pseudoplacenta. Basidiome in situ (JV0709/54).

pore surface pale apricot to salmon-pink when fresh, exuding white drops,

turning up dark reddish brown on drying. Porzs circular to angular, 3-4 per

mm, with thick, entire dissepiments that finally become thin and lacerate;

margin narrowly sterile but sometimes up to 15 mm broad in actively growing

stage, whitish at first, later pale apricot, on drying dark reddish brown to black

and often peeling off. CONTEXT pale orange-pinkish when fresh, blackish when

dry, less than 1 mm thick; tube layer pale salmon-pink, up to 5 mm thick.

HyPHAL SYSTEM monomitic, hyphae hyaline, thin-walled, often branched,

2-4 um in diam., with abundant, often closely spaced clamps. In Melzer's

reagent, the hyphae are covered by small oil drops that often fuse into large,

brownish, oily blotches; in KOH the hyphae are stripped of the oily matter

that forms irregular, brown aggregates in the tissue. CysTIpIA none. BASIDIA

inconspicuous, not much broader than hyphal ends, thin-walled, narrowly

clavate, 4 -sterigmate, with a basal clamp, 15-25 x 4-5 um,. BASIDIOSPORES

broadly ellipsoid, distinctly narrowing to the apex in most cases, hyaline, thin-

walled, smooth, IKI-, 3.5-4.5(-5) x 2.2-3 um. On rather rotten logs of various

coniferous trees. White rot.

ADDITIONAL SPECIMENS EXAMINED: Ceriporiopsis pseudoplacenta - USA.

WASHINGTON: Hoh River, conifer log, 6 Aug 2003, J. Vlasak (JV0308/61, PRM 899298,

ITS sequence identical with JN592497); CALIFORNIA: Kings Canyon Nat. Park, 12

224 ... Vlasdk, Vlasdk & Ryvarden

PLATE 6: Ceriporiopsis pseudoplacenta. Basidiome in situ (JV0709/53).

Aug 2001, conifer log, J. Vlasak (JV0108/82A, PRM 899299); Crescent City, Jed Smith

State Park, Simpson Grove, hemlock, 8 Sep 2007, J. Vlasak (JV0709/53, PRM 899300,

GenBank JN592498, JN592505); 8 Sep 2007, J. Vlasak (JV0709/54); TENNESSEE: Great

Smoky Mt., Gregory Ridge Trail, hemlock, Sep 2005, J. Vlasak (JV0509/52, GenBank

JN592499, JN592506). Auriporia aurea - USA. CALIFORNIA: Sequoia Nat. Park,

Wolverton Area, Abies magnifica A. Murray log, 14 Sep 2001, J. Vlasak (JV0109/92,

PRM 915966, GenBank GU595152). Auriporia aurulenta - CZECH REPUBLIC. Red

Marsh Nat. Preserve, Pinus sylvestris L., 25 Oct 2009, J. Vlasak (JV0910/12, PRM 915967,

GenBank GU595153). Ceriporiopsis sp. - USA. FLoripa: Everglades Nat. Park, Long

Pine Key, 20 Apr 2009, Pinus elliottii Engelm., J. Vlasak (JV0904/46, GenBank JN592500,

JN592507). Postia placenta - USA. NEw York: Upper Jay, Adirondacks Park, Pinus sp.,

20 Sep 2005, J. Vlasak (JV0509/174, GenBank JN592502); CALIFORNIA: Yosemite Nat.

Park, Tuolumne Grove, Pinus lambertiana Douglas, 17 Aug 2001, J. Vlasak (JV0108/98,

GenBank JN592501). SLOVAK REPUBLIC: Dobroc Virgin Forest, 21 Sep 2009, Picea

abies L. (H.Karst.), J. Vlasak (JV0909/16, GenBank JN592503).

COMMENTS — Ceriporiopsis pseudoplacenta is probably widely distributed

in the western USA, as we found it in various locations during every trip to

California and Washington (Vlasak 2008). One specimen comes also from the

Great Smoky Mt., Tennessee. It is difficult to understand why this strikingly

colored fungus has not previously been described, yet we found nothing similar

in the literature. In the field, C. pseudoplacenta can be mistaken for Auriporia

aurea (Peck) Ryvarden or Postia placenta (Fr.) M.J. Larsen & Lombard because

New western polypores (U.S.A.) ... 225

PLATE 7: Ceriporiopsis pseudoplacenta. Basidiome in situ (JV0509/52).

of similar colors and soft, fleshy consistency. Auriporia aurea (and the similar

European A. aurulenta A. David, Torti¢ & Jeli¢c) differ by having skeletal hyphae

and thick-walled cystidia. Postia placenta is difficult to separate by microscopy

alone because both species have similar hyphal system with no special

structures and similar spores that are, however, smaller in C. pseudoplacenta.

Macroscopical differences are more distinctive, especially when monitoring

basidiome changes during development and drying. Postia placenta starts

growing usually as a deep-red basidiome, changing to rosy-pink and, when

old or dry, to whitish-pink or light ochre. C. pseudoplacenta follows just the

opposite coloration steps; it starts as whitish or light salmon pink, becoming

increasingly more deeply red until maturing to red brownish or black in the

subiculum and margin. Also, the consistency of dried fruitbodies is quite

different: soft and brittle in P placenta but tough and rubber-like (reminiscent of

beef jerky) in C. pseudoplacenta. Auriporia aurea, A. aurulenta, and P. placenta

cause a dry brown rot. The wood under C. pseudoplacenta basidiomes is usually

also quite dark as the fungus causes some red-brownish wood coloration, but

its stringy fibrous structure with small pockets corresponds more to white

rot. Unfortunately, we did not perform oxidase tests on fresh fruitbodies.

Ceriporiopsis gilvescens (Bres.) Domanski, a white rot polypore, also sometimes

resembles C. pseudoplacenta, but its salmon-pinkish colours are usually

226 ... Vlasdk, Vlasak & Ryvarden

PLATE 8: Ceriporiopsis pseudoplacenta microscopic structures.

a: Basidiospores, b: Basidia, c: Hyphae.

paler and the spores are narrower (1.5 - 2 um). Antrodiella albocinnamomea

Y.C. Dai & Niemela, widely distributed in Asia (Dai & Niemela 1997; Dai 2012),

is macroscopically similar to C. pseudoplacenta, but differs in its dimitic hyphal

structure and oblong-ellipsoid spores (3.7-5 x 2.1-2.9 um).

The taxonomic position of C. pseudoplacenta is a mystery, as the sequence

analysis (see below) revealed very low homology with known fungi. So we chose

for it the congregate genus Ceriporiopsis Domanski that encompasses white rot

species that are poroid, resupinate, and monomitic with clamp connections.

SEQUENCE ANALYSIS — Successful amplification and sequencing of the

C. pseudoplacenta ITS (nrDNA) region from four of six collected specimens

showed only minor differences between individual specimens (PLATE 9).

5.88 rDNA is highly homologous with many other polypores, but for the

very long ITS1 region (280-283b) the GenBank MegaBlast search found

“no significant similarity,’ which we never experienced before for polypore

species. ITS2 analyses indicate very low similarity with seemingly distant

fungi, with Ceriporiopsis species showing the closest support among the

polypores. Sequence analyses weakly support clustering of C. gilvescens with

C. pseudoplacenta, and apart from the brown-rot fungi Postia Fr. Auriporia

Ryvarden and Antrodia P. Karst. in a separate clade (PLATE 9). Some Phlebia

Fr. species, macroscopically similar except for the hymenophore form, cluster

even more closely with C. pseudoplacenta, as also shown in the nrDNA LSU

sequence analyses (PLATE 10). For the time being, the generic position of the

new species must be regarded as provisional.

New western polypores (U.S.A.) ... 227

JV030868 C. pseudoplacenta n.sp. hapl(1) WA

gg, | JV030868 C. pseudoplacenta n.sp. hapl(2) WA

JV070953 C. pseudoplacenta n.sp. CA

JV030861 C. pseudoplacenta n.sp. WA

JV050952 C. pseudoplacenta n.sp. TN

a4 *DQ056859 Phlebia radiata

45 *AB084616 Phlebia brevispora

*FJ496686 C. gilvescens

JV0904 46 Ceriporiopsis sp. Florida

100

oF *F J496683 C.aneirina

*DQ491414 Antrodia albida

100 *GU594152 Auriporia aurea

100 *GU594153 Auriporia aurulenta

77 *EF524035 P. placenta

JV0108 98 P. placenta Tuluome Grove CA

JV0509 174 P. placenta Adirondacks NY

JV0909 16 P. placenta Dobroc Europe

sae

0.02

PrateE 9. Evolutionary relationships using the Neighbor-Joining method of 16 Antrodia, Auriporia,

Ceriporiopsis, Phlebia, and Postia specimens based on nr DNA ITS region sequence. The bootstrap

consensus tree inferred from 1000 replicates is taken to represent the evolutionary history of the

taxa analyzed. The percentage of replicate trees in which the associated taxa clustered together

in the bootstrap test (1000 replicates) is shown next to the branches. The tree is drawn to scale,

with branch lengths in the same units as those of the evolutionary distances used to infer the

phylogenetic tree. The evolutionary distances are in the units of the number of base substitutions

per site. Accession numbers with * represent sequences retrieved from GenBank. All other

GenBank accession numbers refer to new sequences listed in this paper.

JV0709 53 C. pseudoplacenta n.sp. CA

JV0309 68 C. pseudoplacenta n.sp. WA

JV0509 52 C. pseudoplacenta n.sp. TN

“AF 287885 Phlebia radiata

JV0904 46 Ceriporiopsis sp. FL

*DQ144618 C. gilvescens

“F J496704 C. aneirina

“AF515355 P. placenta

0.01

PLaTE 10. Evolutionary relationships of 8 Ceriporiopsis, Phlebia and Postia specimens based on

nrDNA LSU region sequence. Phylogenetic analysis as referred in PL. 9.

228 ... Vlasdk, Vlasak & Ryvarden

Another Postia placenta-like specimen (JV0904/46, PLATE 9) we have

collected in early spring from the tropical region of southern Florida from

slough pine (Pinus elliottii) is morphologically very similar to P. placenta or

C. pseudoplacenta except for slightly shorter spores, but its sequence is more

similar to Ceriporiopsis aneirina (Sommerf.) Domanski. We need additional

material before resolving the identity of this specimen.

Skeletocutis subodora Vlasak & Ryvarden, sp. nov. PLATES 11-13

MycoBank MB 563083

Basidiomata annua, resupinata, usque ad 10 cm lata et 1 cm spissa, colore albicante, pori

rotundi vel angulati, 3-5 pro mm, contextus albus, 1-4 mm crassus. Systema hypharum

monomiticum, hyphae fibulatae, cystidioli hyalinni, fusiformes, basidiosporaes hyalinae,

leves, tenuitunicatae, non amyloideae, non dextrinoideae, 5-6 x 2.0-2.3 um.

Type: USA. Oregon: Crater Lake visitors center, Pseudotsuga menziesii (Mirb.) Franco

log, 15 Sep 2007, J. Vlasak JV0709/134 (Holotype, PRM 899301; isotype, JV0709/134a,

GenBank JN592509).

EryMo_oey: From the Latin for odora-like, referring to the similarity to S. odora.

Cre | es,

vO2

SHAD)

PLaTE 11. Skeletocutis subodora microscopic structures.

a: Basidiospores, b: Hymenium with basidia and cystidioles.

New western polypores (U.S.A.) ... 229

PLATE 12. Skeletocutis subodora. Basidiome (PRM 899301, holotype)

BASIDIOMATA annual, resupinate, found in two patches effused to 5 x 5 and

10 x 10 cm, up to 1 cm thick, soft when fresh, no odor noted, drying brittle

and shrinking; sterile margin 1-3 wide, well demarcated, whitish, tawny on

drying; pore surface cream-colored, drying pale beige. Pores at first regular,

3-5 per mm, with very thin dissepiments, on drying becoming more dented

and more irregular. SuBIcULUM white to pale cream, 1-4 mm thick; tube layer

concolorous with pore surface, resinous and compressed when dry, distinctly

ochre-colored compared with the white subiculum, up to 8 mm thick. HyPHAL

SYSTEM monomitic, hyphae hyaline, thick to thin-walled, often branched, with

clamps, 2-4 um in diam., their ends on dissepiments edges sparingly incrusted

by sharp crystals. CysTIDIAL ELEMENTS as fusoid cystidioles abundant and

conspicuous, thin-walled, with a basal clamp, 15-25 x 2-6 um. Basrp1a clavate,

4-sterigmate, with a basal clamp, 15-20 x 5-6 um. BASIDIOSPORES narrowly

ellipsoid to cylindrical, hyaline, thin-walled, smooth, IKI-, 5-6 x 2-2.3 um. On

decorticated log of Douglas fir, causing a white pocket rot.

ADDITIONAL SPECIMENS EXAMINED: Ceriporiopsis guidella - ITALY: Ris. Di Sasso

Frantino, Rio Cullacce (Forli), Acer sp., 22 Sep 2001, A-R. Bernicchia (coll. 7731).

Skeletocutis diluta - USA. FLoripa: Everglades Nat. Park, Long Pine Key, Pinus elliottii,

29 Aug 2010, J. Vlasak (collection JV1008/61, PRM 899302, GenBank JF692198).

Skeletocutis odora - CLECH REPUBLIC. Zofin Nat. Preserve, Picea abies log, 22 Jul

2010, J. Vlasak (JV 1007/7, GenBank JN592509).

ComMENTs. The fungus was collected as a probable Skeletocutis odora (Sacc.)

Ginns, which is actually very similar macroscopically and with a sequence

that shows a high degree of homology (see below). The most important

macroscopical diagnostic feature is the thick (< 2-4 mm) subiculum, barely

1 mm thick in S. odora. The distinctive (relatively broad non-allantoid)

basidiospores, conspicuous large cystidioles, and monomitic context also

distinguish S. subodora. The macroscopically similar Ceriporiopsis guidella

Bernicchia & Ryvarden (Bernicchia & Ryvarden 2003) also produces similar

230 ... Vlasdk, Vlasdk & Ryvarden

cystidioles and spores but differs in an extremely thin subiculum, growth on

hardwoods, and different ITS nrRNA sequence (see below).

Thus far the new species is known only from the type locality, although

the occurrence of the similar Skeletocutis odora in the USA warrants mention.

Lowe (1966) stated that S. odora is known with certainty only from Peck’s

type and one other report from Pennsylvania, while Gilbertson & Ryvarden

(1987) do not cite the species at all from the USA. We also never found S. odora

during numerous excursions in either the east or west USA, although it is not

uncommon in old-growth conifer forests in Europe.

SEQUENCE ANALYSIS — Theamplified ITS1-5.8S-ITS2 sequence from Skeletocutis

subodora (603b long between ITS1 and ITS4 primers) was somewhat similar

(variations of 3.0% (6/195) for ITS1 and 4% (8/205) for ITS2) to that of S. odora

(PLATE 13), suggesting that the two species are related to but different from

each other. Surprisingly, Tyromyces chioneus (Fr.) P. Karst., with a still unsettled

taxonomic position, shows surprisingly more homology than other Skeletocutis

species.

109 ; JV1007 7 Skeletocutis odora

FJ903307 Skeletocutis odora

JV0709 134 Skeletocutis subodora n.sp.

*AJO06676 Tyromyces chioneus

JF692198 Skeletocutis diluta

*AJ006678 Skeletocutis kuehneri

*F J496687 Ceriporiopsis guidella

*F J496683 Ceriporiopsis aneirina

44 *F J496686 Ceriporiopsis gilvescens

0.02

PLaTE 13. Evolutionary relationships of 9 Skeletocutis and Ceriporiopsis specimens based on nr

DNA ITS region sequence. Phylogenetic analysis as referred in PL. 9.

Acknowledgements

Sergio Pérez Gorjon and Yu-Cheng Dai have kindly acted as presubmission reviewers

and their help with text improvement is acknowledged. We are grateful also to Petr

Vampola for checking the microscopic inspection results and valuable comments. ‘This

research was supported by CEZ:AV0Z50510513 fund.

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79: 57-66.

Bernicchia A, Ryvarden L. 2003. A new white-rot polypore from Italy. Mycotaxon 88: 219-224.

CI [Conservation International] 2007. Biodiversity hotspots. (accessed online 01.Sep.2011).

http://www. biodiversityhotspots.org/xp/hotspots/mediterranean/Pages/default.aspx

New western polypores (U.S.A.) ... 231

Dai YC. 2010. Hymenochaetaceae (Basidiomycota) in China. Fungal Diversity 45: 131-343.

http://dx.doi.org/10.1007/s13225-010-0066-9

Dai YC. 2012. Polypore diversity in China with an annotated checklist of Chinese polypores.

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Dai YC, Niemela T. 1997. Changbai wood-rotting fungi 6. Study on Antrodiella, two new species

and notes on some other species. Mycotaxon 64: 67-81.

Gilbertson RL. 1976. The genus Inonotus (Aphyllophorales, Hymenochaetaceae) in Arizona. Mem.

New York Bot. Gard. 28: 67-85

Gilbertson RL. 1979. The genus Phellinus (Aphyllophorales, Hymenochaetaceae) in western North

America. Mycotaxon 9: 51-89.

Gilbertson RL, Ryvarden L. 1986. North American polypores, Vol. 1. Fungiflora. Oslo. pp. 1-433.

Gilbertson RL, Ryvarden L. 1987. North American polypores, Vol. 2. Fungiflora. Oslo. pp.

434-885.

Gorjén SP, Bernicchia A. 2009. Antrodia sandaliae (Polyporales, Basidiomycota), an interesting

polypore collected in the Iberian Peninsula. Cryptog. Mycol. 30(1): 53-56.

Hattori T. 1999. Phellinus setifer sp. nov. and P acontextus, two noteworthy polypores from

temperate areas of Japan, with notes on their allies. Mycoscience 40: 483-490.

Lowe JL. 1966. Polyporaceae of North America. The genus Poria. State Univ. Coll. Forestry Syracuse

Univ. Techn. Publ. 90: 1-183.

Murrill WA. 1912. Polyporaceae and Boletaceae of the pacific coast. Mycologia 4(2): 91-100.

http://dx.doi.org/10.2307/3753546

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Overholts LO. 1939. Geographical distribution of some American polypores. Mycologia 31(6):

629-652. http://dx.doi.org/10.2307/3754336

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phylogenetic trees. Mol Biol Evol 4: 406-425.PMid:3447015

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(MEGA) software version 4.0. Mol Biol Evol 24: 1596-1599. http://dx.doi.org/10.1093/molbev/

msm092

Vlasak J. 2008. Polypores in national parks in the USA North-West (in Czech). Mykologické listy

104: 30-34.

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White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal

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methods and applications. San Diego, Academic Press.

MY COTAXON

http://dx.doi.org/10.5248/119.233

Volume 119, pp. 233-239 January-March 2012

Studies on Croatian Basidiomycota 2: Marasmiellus milicae sp. nov.

ARMIN MESIC', ZDENKO TKALCEC! & VLADIM{R ANTONIN?

'Ruder Boskovic¢ Institute, Bijenicka 54, HR-10000 Zagreb, Croatia

*Moravian Museum, Dept. of Botany, Zelny trh 6, CZ-659 37 Brno, Czech Republic

CORRESPONDENCE TO *: amesic@irb.hr, *ztkalcec@irb.hr, vantonin@mzm.cz

ABSTRACT — A new species, Marasmiellus milicae, is described from the Mediterranean

part of Croatia and compared with similar species. It is characterized by having rather

small, white pileus, distinctly to strongly eccentric, curved, short stipe, white hymenophore

becoming dark brown when dry, (sub)amygdaloid basidiospores, mostly diverticulate

cheilocystidia, pileipellis and stipitipellis with strongly developed Rameales-structure, and

gelatinized trama. Color photographs and line drawings of microscopic elements and

basidiomata accompany the description.

Key worps — biodiversity, mycobiota, Omphalotaceae, taxonomy

Introduction

During the last decade of intensive research of Croatian Basidiomycota,

a significant number of rare and new species has been found. Some of these

have been published in the last few years (Hausknecht et al. 2007; Mesic &

Tkaléec 2008, 2009; Tkaléec & MeSi¢ 2008; Tkaléec et al. 2009) and the

remaining will be published in a series of papers under the title “Studies on

Croatian Basidiomycota,” of which this is the second. In the present paper we

describe a new species of Marasmiellus (Omphalotaceae, Agaricales) found in

the Mediterranean part of Croatia.

Materials & methods

The description of Marasmiellus milicae is based on two collections consisting of

27 basidiomata in total. The macroscopic description is based on fresh basidiomata.

Microscopic features are described from dried material mounted in potassium hydroxide

(KOH) solution and Melzer’s reagent, and observed with a light microscope (brightfield

and phase contrast - PhC) under magnification up to 1500x. Spore measurements do

not include the apiculus. Arithmetic means of spores length and width are shown in

italic font in the center of the measurements. The length/width ratio of all measured

spores is given as the “Q” value (min. - av. - max.), and the range of arithmetic averages

234 ... Mesié, Tkaléec & Antonin

of “Q” value (Q__) of both collections is also given. The holotype and additional collection

are deposited in the Croatian National Fungarium, Zagreb, Croatia (CNF), while an

isotype is deposited in the Herbarium of the Moravian Museum, Brno, Czech Republic

(BRNM).

Taxonomy

Marasmiellus milicae Me&i¢, Tkaléec & Antonin, sp. nov. FIGs 1-10

MycoBANnkK MB 561574

Differs from the most similar species by its white pileus; short, eccentric, dark stipe; white

hymenophore becoming dark brown when dry; (sub)amygdaloid spores on average 9.1

x 5.4 um; mostly diverticulate cheilocystidia; and strongly developed pileipellis and

stipitipellis Rameales-structure.

TYPE: Croatia, island of Mljet (near Dubrovnik), near Prozurska Luka village, 42°43'51"N

17°38'44"E, alt. 60 m, 9 November 2009, leg. M. Cerkez & Z. Tkaléec (Holotype CNF

1/5692, Isotype BRNM 736106).

Erymo.oecy: Named after Croatian mycologist Dr. Milica Torti¢ (1920-2008).

PILEus 6-18 mm broad, hemispherical to convex, margin involute at first, then

inflexed to deflexed, slightly crenulate when young, surface smooth to entirely

rugose, finely tomentose, white, sometimes with grayish to greenish-gray

tinge. LAMELLAE adnate, distant, L = 8-12, | = 0-2(-3), strongly intervenose,

especially in larger specimens, sometimes furcate, often + sinuous, white with

concolorous edge, whole hymenophore becoming dark brown when dry. STIPE

1-3 x 0.5-1.2 mm, distinctly to strongly eccentric, curved in horizontal direction

(basidiomata laterally attached to the substratum), cylindrical with slightly

broadened apex, insititious, pubescent, white when young, then becoming

dark gray to gray-black, sometimes with greenish tinge at the apex. CONTEXT

concolorous with outer surface. SMELL very weak, fungoid when cut.

BASIDIOSPORES 8-9.1-10 xX 4.5—-5.4-6.0 um, average = 9.0-9.3 x 5.3-5.7

um, Q = 1.5-1.70-1.98, Q., = 1.63-1.73, mostly (sub)amygdaloid in side

view, ellipsoid to fusoid in frontal view, thin-walled, hyaline, apiculus ca. 1.0

x 1.3 um. BasipIA 28-47 x 6.0-10 um, 4-spored, clavate. BAsIDIOLES 15-45

x 3.0-10 um, clavate, cylindrical, subfusoid. LAMELLAR EDGE heterogeneous.

CHEILOCYSTIDIA (10-) 13-60 x 5.0-14 um, clavate, subcapitate, (sub)cylindrical,

fusoid, mostly with simple, branched to coralloid apical projections (digitate,

conical, or obtuse) up to 30 x 5.0 um; thin-walled, hyaline. PLEUROCYSTIDIA

absent. PILEAL AND HYMENOPHORAL TRAMA gelatinized, hyphae cylindrical,

thin-walled, sometimes branched, up to 10 um wide. PILEIPELLIs an ixocutis

composed of + cylindrical, thin- to slightly thick-walled, smooth or minutely

incrusted, diverticulate or smooth, up to 7.0 um wide hyphae; terminal cells

and lateral projections up to ca. 20 x 3.0 um, digitate, branched to coralloid,

obtuse (well-developed Rameales-structure). STIPITIPELLIS a cutis composed

Marasmiellus milicae sp. nov. (Croatia) ... 235

Fics 1-3. Marasmiellus milicae. 1. Fresh basidiomata (holotype). 2. Fresh basidiomata (CNF 1/5774).

3. Dried basidiomata (holotype). Bars = 5 mm.

236 ... Mesié, Tkaléec & Antonin

of cylindrical or rarely subinflated, mostly thick-walled [walls up to 1.0(-2.0)

um], smooth or diverticulate, up to 6.0(-15) um wide hyphae; terminal cells

cylindrical to clavate, branched to coralloid, thin- or slightly thick-walled, up

to 10 um wide, some of them with apical, + clavate or subcapitate outgrowth.

CLAMP CONNECTIONS present and abundant in all tissues. CHEMICAL

REACTIONS: all parts of basidioma non-amyloid and non-dextrinoid.

Hasitat — On the type locality in the Mediterranean maquis with

Arbutus unedo, Erica arborea, Phillyrea latifolia, Pistacia lentiscus, and Myrtus

communis; on hanging, dead stem of Lonicera implexa. On additional locality

in evergreen park-forest with Quercus ilex and Laurus nobilis; on dead stem of

Hedera helix.

DISTRIBUTION — Known only from two localities in the Mediterranean part

of Croatia.

ADDITIONAL SPECIMEN EXAMINED: CROATIA: DUBROVNIK-NERETVA County, 10 km

NW from Dubrovnik, Trsteno arboretum in Trsteno village, 42°42’43” N 17°58’35" E, alt.

55 m, 13 November 2009; leg. M. Cerkez & A. Me’ié (CNE 1/5774, BRNM 736107).

REMARKS — Marasmiellus milicae is characterized by having a rather small,

white pileus, distinctly to strongly eccentric, curved, short, dark stipe, white

hymenophore becoming dark brown when dry, (sub)amygdaloid basidiospores,

mostly diverticulate cheilocystidia, pileipellis and stipitipellis with strongly

developed Rameales-structure, and gelatinized trama. Having those characters,

it belongs to sect. Distantifolii Singer (Singer 1973). So far, it has been collected

only in Croatia.

Several marasmielloid species with similar shape of basidiomata are

known in Europe. Among the white colored species, Gymnopus sphaerosporus

M. Villarreal et al. and Marasmiellus margaritifer (Maire) Singer have a

white stipe, globose or subglobose basidiospores, and different cheilocystidia

(Antonin & Noordeloos 2010). From the white colored species of section

Distantifolii, Marasmiellus distantifolius (Murrill) Singer, known from the USA,

Central and South America, especially differs by a pure white stipe, distinctly

longer basidiospores ((8.5—)10.2-16 x 3-5.5(-6) um according to Singer 1973;

11.5-14.5 x 4—5.5 um according to Pegler 1977, 1997), and smaller cheilocystidia

(20-25 x 4.5 um); M. corticum Singer, described from the USA, differs by a

shortly radially ferrugineous-brown or ochraceous-brown striped pileus

on drying, and smaller ((7-)14-21.5 x (5-)7-9.5 um), differently shaped

cheilocystidia (Singer 1973). Marasmiellus afer Pegler, collected in tropical

Africa, has a lateral, rarely absent stipe and narrower (8.2-10.8 x 3-4 um),

lacrymoid basidiospores (Pegler 1967, 1977). Marasmiellus tristis (G. Stev.)

E. Horak (= Campanella tristis (G. Stev.) Segedin) from New Zealand differs

by its pileus turning glaucous or greenish-gray with age, a dirty yellowish

hymenophore when dry, spores with lower Q value (1.4), and the absence of

Marasmiellus milicae sp. nov. (Croatia) ... 237

Fics 4-9. Marasmiellus milicae (holotype). 4. Spores. 5, 6. Cheilocystidia. 7. Stipitipellis. 8. Basidium.

9. Pileipellis. All under PhC microscope. Bars: 4 = 5 um; 5-9 = 10 um.

238 ... Mesié, Tkaléec & Antonin

ans

Fic. 10. Marasmiellus milicae.

a. Elements of pileipellis. b. Spores. c. Cheilocystidia. d. Elements of stipitipellis. Bar = 20 um.

Marasmiellus milicae sp. nov. (Croatia) ... 239

branched or coralloid apical projections on cheilocystidia (Segedin 1993).

Marasmius exillimus Corner, described from Solomon Islands, has a white, pale

fuscous striate pileus, a whitish or pallid fuscous stipe, smaller basidiospores

(5.5-7.5 x 3.5-4 um), and a different pileipellis and stipitipellis (Corner 1996).

Acknowledgments

We are very grateful to Dr. Machiel E. Noordeloos (Netherlands Centre for

Biodiversity Naturalis, Leiden, The Netherlands) and Dr. Alfredo Vizzini (Universita

degli Studi di Torino, Torino, Italy) for their critical review of the manuscript.

Literature cited

Antonin V, Noordeloos ME. 2010. A monograph of marasmioid and collybioid fungi in Europe.

IHW- Verlag: Eching (Germany).

Corner EJH. 1996. The agaric genera Marasmius, Chaetocalathus, Crinipellis, Heimiomyces,

Resupinatus, Xerula and Xerulina in Malesia. Nova Hedwigia Beih. 111: 1-175.

Hausknecht A, Mesi¢ A, Tkaléec Z. 2007. Two remarkable species of Bolbitiaceae (Agaricales) from

Croatia. Osterr. Z. Pilzk. 16: 281-286.

MeSi¢ A, Tkaléec Z. 2008. Entoloma reinwaldii, a rare species new to Croatia. Mycotaxon 105:

295-300.

MeSi¢ A, Tkaléec Z. 2009. Studies on Croatian Basidiomycota 1: Gerhardtia piperata (Agaricales).

Mycotaxon 110: 413-421. http://dx.doi.org/10.5248/110.413

Pegler DN. 1967. Studies on African Agaricales: I. Kew Bull. 21(3): 499-533.

http://dx.doi.org/10.2307/4107943

Pegler DN. 1977. Preliminary agaric flora of East Africa. Kew Bull. Addit. Ser. VI. Royal Botanic

Gardens: Kew (UK).

Pegler DN. 1997. The agarics of Sao Paulo. An account of the agaricoid fungi (Holobasidiomycetes)

of Sao Paulo State, Brazil. Royal Botanic Gardens: Kew (UK).

Segedin BP. 1993. Studies in the Agaricales of New Zealand: some new and revised species of

Campanella (Tricholomataceae: Collybieae). New Zealand J. Bot. 31: 375-384.

Singer R. 1973. The genera Marasmiellus, Crepidotus and Simocybe in the Neotropics. Nova

Hedwigia Beih. 44: 1-517.

Tkaléec Z, Mesi¢ A. 2008. Gloiocephala cerkezii, a new species from Croatia. Mycologia 100(2):

320-324. http://dx.doi.org/10.3852/mycologia.100.2.320

Tkaléec Z, Mesi¢ A, Hausknecht A. 2009. Two new taxa of Bolbitiaceae from Croatia. Mycotaxon

107: 249-258. http://dx.doi.org/10.5248/107.249

MYCOTAXON

http://dx.doi.org/10.5248/119.241

Volume 119, pp. 241-248 January-March 2012

Zelodactylaria, an interesting new genus

from semi-arid northeast Brazil

ALISSON CARDOSO RODRIGUES DA CRUZ’,

Luis FERNANDO PASCHOLATI GUSMAO’, RAFAEL FE. CASTANEDA-RUIZ’,

Marc STADLER? , & DAviID W. MINTER‘*

"Departamento de Ciéncias Biolégicas, Laboratorio de Micologia Universidade Estadual

de Feira de Santana, BR116 KMO3, 44031-460, Feira de Santana

?Instituto de Investigaciones Fundamentales en Agricultura Tropical ‘Alejandro de Humboldt’

(INIFAT), Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200

*InterMed Discovery GmbH, Otto-Hahn-Str. 15, D-44227 Dortmund, Germany

*CABI,Bakeham Lane, Egham, Surrey, TW20 9TY, United Kingdom

*CORRESPONDENCE TO: Marc.Stadler@t-online.de

ABSTRACT — During investigation of microfungi on dead plant material in a semiarid region

of northeast Brazil, an interesting fungus was collected that is described and illustrated here.

Zelodactylaria verticillata anam. gen. & sp. nov. is distinguished by macronematous verticillate

brown conidiophores with marked sympodially proliferating conidiogenous cells with long

cylindrical denticles and solitary obovoid to clavate hyaline 0-1-septate conidia.

KEY worpDs — anamorphic fungi, systematic, bark

Introduction

The Brazilian semi-arid zone, called the Caatinga biome, is located almost

exclusively in the northeast of the country. This is an expanse of dry land that

stretches between 3-17°S and 35-45°W, covers almost 8% of Brazil, and occupies

an approximate 900,000 km? area. The climate of northeast Brazil is one of

the most complex in the world, with an annual rainfall up to 2000 mm along

the coast and only 300-500 mm in the semi-arid zone (Giulietti et al. 2006).

During a mycological survey of anamorphic fungi from the semi-arid region of

Bahia state, a conspicuous fungus was collected that differed morphologically

from any previously described anamorphic fungus. It is described here as a new

genus.

242 ... Cruz & al.

Materials & methods

Samples of plant material were collected during a mycological survey from the semi-

arid region of Bahia, Brazil. Individual collections were placed in paper and plastic

bags, taken to the laboratory and treated according to Castafieda (2005). Mounts were

prepared in polyvinyl alcohol-glycerol (8 g in 100 ml of water, plus 5 ml of glycerol) and

measurements made at a magnification of x1000. Micrographs were obtained with a

Zeiss Axioskop 40.

Taxonomy

Zelodactylaria A.C. Cruz, Gusmao & R.F. Castafieda, anam. gen. nov.

MycoBank MB561559

Conidiophora macronematosa, mononematosa, erecta, ramosa, septata, brunnea Cellulae

conidiogenae polyblasticae, discretae, sympodialiter prolificantes, denticulatae. Conidia

solitaria, clavata, obovoidea usque globosa, septata, hyalina. Teleomorphosis ignota.

Type SpEcIEs: Zelodactylaria verticillata A.C. Cruz et al.

ErymMoLoecy: Greek, Zelo- meaning emulation; -dactylaria, referring to a genus of

anamorphic fungi.

CoLonigs on the natural substratum effuse, hairy, brown to olivaceous or

black. Mycelium superficial and immersed. CONIDIOPHORES macronematous,

mononematous, erect, branched, septate, brown or black, smooth orverruculose.

CONIDIOGENOUS CELLS polyblastic, discrete, sympodially proliferating,

denticulate. Conidial secession schizolytic. Conrp1a solitary, clavate, obovoid

to globose, septate, smooth or verruculose, hyaline. Teleomorph unknown.

Zelodactylaria verticillata A.C. Cruz, Gusmao & R.E. Castafieda,

anam. sp. nov. Figs. 1 -4

MycoBank MB561560

Conidiophora macronematosa, mononematosa, cum ramis verticillatis (ramis 20-50 x 2.5-

4.0 um), 7- ad 12-septata, brunnea ad basim, 150-220 x 4-10 ym. Cellulae conidiogenae

polyblasticae, 10-30 x 2-3 um denticulatae cum denticulis conspicuis, 1.0-2.5 um longis .

Conidia solitaria, clavata vel obovoidea, 0- ad 1-septata, hyalina, 11-17 x 3-5 um.

Type: BRAZIL. Bahia: Senhor do Bonfim, “Serra da Maravilha’, 12°23’S 40°12'W,

on decaying bark of unidentified plant, 22.[X.2006, coll. A.C.R. da Cruz (Holotype:

HUEFS169132).

Erymo_oey: Latin, verticillata, referring to the whorled branches of the conidiophores.

CoLoniEs on the natural substratum, effuse, hairy, brown. Mycelium superficial

and immersed. Hyphae septate, branched, smooth, brown. CONIDIOPHORES

macronematous, mononematous, 7- to 12-septate, erect, straight or flexuous,

smooth, 150-220 x 4-10 um, brown at the base, subhyaline towards the apex;

producing verticillate, 20-50 x 2.5-4.0 um, septate, pale brown to subhyaline

branches. CONIDIOGENOUS CELLS polyblastic, discrete, terminal and intercalary,

10-30 x 2-3 um, subhyaline, with marked sympodial proliferations, with

Zelodactylaria veticillata gen. & sp. nov. (Brazil) ... 243

Fic. 1. Zelodactylaria verticillata.

A-F. Conidiophores. Scale bar = 50 um.

244 ... Cruz & al.

Fic. 2. Zelodactylaria verticillata.

A. Conidiophore and conidiogenous cells. B, D. Conidiogenous cells and conidia.

C. Conidiogenous cell. Scale bars = 10 um.

Zelodactylaria veticillata gen. & sp. nov. (Brazil) ... 245

Fic. 3. Zelodactylaria verticillata.

A-H. Conidia. Scale bar = 10 um.

246 ... Cruz & al.

: S$ F,.

wb! SETS A) toe

pe See:

Fic. 4. Zelodactylaria verticillata.

Conidiophores, conidiogenous cells and conidia. Scale bars = 10 um.

Zelodactylaria veticillata gen. & sp. nov. (Brazil) ... 247

conspicuous, shortly cylindrical, 1.0-2.5 um long, flat-topped denticles.

Conlipia solitary, clavate to obovoid, truncate at the base, (0—) 1-septate, hyaline,

guttulate, smooth, dry, 11-17 x 3-5 um. Teleomorph unknown.

NoTE: Conidial ontogeny in Dactylaria Sacc. (De Hoog 1985, Goh & Hyde 1997,

Paulus et al. 2003), Calcarisporium Preuss (De Hoog 1974) Pleurophragmium

Costantin (Matsushima 1985, 1987, 1993, 1995, 1996; Heredia et al. 2007),

Selenosporella G. Arnaud ex MacGarvie (Castaneda et al. 2009), Selenosporopsis

R.E Castafieda & W.B. Kendr. (Castafieda & Kendrick 1991), Tritirachium Limber

(De Hoog 1972), and Umbellidion B. Sutton & Hodges (Sutton & Hodges 1975)

can be compared with Zelodactylaria, particularly in terms of the sympodial

proliferation with denticulate conidiogenous cells. There are, however,

clear differences from those genera in respect of the verticillate ramification

present in Zelodactylaria. The conidiophores in some Selenosporella species

and the monotypic Selenosporopsis are verticillate, but conidia are unicellular,

filiform to falcate, produced in short and long denticulate conidiogenous loci

respectively. In Calcarisporium and Tritirachium conidia form on sympodially

elongating conidiogenous cells arranged in verticils, but the conidiogenous loci

in Calcarisporium develop at the top of each branch and in Tritirachium are

inconspicuous scars in a long geniculate rachis (Kendrick 2003), which also

differentiate both genera from Zelodactylaria. In Umbellidion the conidiogenous

cells are arranged in umbels, with several short unthickened, inconspicuous

denticles restricted to the apices and the conidia are aseptate.

Acknowledgments

The authors express their sincere gratitude to Drs. Lori Carris and De-Wei Li for

their critical review of the manuscript. The authors are grateful to the Cuban Ministry

of Agriculture for facilities. The authors thank to Drs Uwe Braun, Lori Carris, De-

Wei Li, and Gregorio Delgado for their generous and valued assistance with literature

not otherwise available. We thank Beatriz Ramos and Mirtha Caraballo for technical

assistance. We also acknowledge the facility provided by Dr P.M. Kirk and Drs. V. Robert

and G. Stegehuis through the Index Fungorum and Mycobank websites. Dr Lorelei L.

Norvell’s editorial review and Dr Shaun Pennycook’s nomenclature review are greatly

appreciated.

Literature cited

Castafieda Ruiz RF. 2005. Metodologia en el estudio de los hongos anamorfos. Anais do V

Congresso Latino Americano de Micologia. Brasilia: 182-183.

Castafeda-Ruiz RE, Kendrick B. 1991. Ninety-nine conidial fungi from Cuba and three from

Canada. University of Waterloo Biology Series 35: 132 p.

Castafieda-Ruiz RE, Guerrero B, Adamo GM, Morillo O, Minter DW, Stadler M Gené J, Guarro

J. 2009. A new species of Selenosporella and two microfungi recorded from a cloud forest in

Mérida, Venezuela. Mycotaxon 109: 63-74. http://dx.doi.org/10.5248/109.63

248 ... Cruz & al.

De Hoog GS. 1972. The genera Beauveria, Isaria, Tritirachium and Acrodontium gen.nov. Stud.

Mycol. 1: 1-41. http://www.cbs.knaw.nl/publications/1001/content_files/content.htm

De Hoog GS. 1974. The genera Blastobotrys, Sporothrix, Calcarisporium and Calcarisporiella gen.

nov. Stud. Mycol. 7: 1-88. http://www.cbs.knaw.nl/publications/1007/sim7_files/sim7.htm

De Hoog GS. 1985. Taxonomy of the Dactylaria complex, IV. Dactylaria, Neta, Subulispora and

Scolecobasidium. Stud. Mycol. 26: 1-60.

http://www.cbs.knaw.nl/publications/1026/full%20text.htm

Giulietti AM, Harley RM, Queiroz LP, Rapini A. 2006. To set the scene. 15-19, in: LP Queiroz et

al. (Eds). Towards greater knowledge of the Brazilian semi-arid biodiversity. Ministério da

Ciéncia e Tecnologia. Brasilia.

Goh TK, Hyde KD. 1997. A revision of Dactylaria, with description of D. tunicata sp. nov. from

submerged wood in Australia. Mycol. Res. 101: 1265-1272.

http://dx.doi.org/10.1017/S0953756297004000

Heredia Abarca G, Castafieda-Ruiz RF, Arias RM, Saikawa M, Stadler M. 2007. Anamorphic fungi

from submerged plant material: Acumispora verruculosa sp. nov., Pleurophragmium aquaticum

sp. nov and Pleurophragmium miniumbonato comb. nov. Mycotaxon 101: 89-97.

Kendrick B. 2003. Analysis of morphogenesis in hyphomycetes: new characters derived from

considering some conidiophores and conidia as condensed hyphal systems. Can. J. Bot. 81(2):

75-100. http://dx.doi.org/10.1139/b03-008

Matsushima T. 1985. Matsushima mycological memoirs no. 4. Matsushima Fungus Collection,

Kobe, Japan.

Matsushima T. 1987. Matsushima mycological memoirs no. 5. Matsushima Fungus Collection,

Kobe, Japan.

Matsushima T. 1993. Matsushima mycological memoirs no. 7. Matsushima Fungus Collection,

Kobe, Japan.

Matsushima T. 1995. Matsushima mycological memoirs no. 8. Matsushima Fungus Collection,

Kobe, Japan.

Matsushima T. 1996. Matsushima mycological memoirs no. 9. Matsushima Fungus Collection,

Kobe, Japan.

Paulus B, Gadek P, Hyde KD. 2003. Two new species of Dactylaria (anamorphic fungi) from

Australian rainforests and update of species in Dactylaria sensu lato. Fungal Diversity 14:

143-156. http://www.fungaldiversity.org/fdp/sfdp/14-11.pdf

Sutton BC, Hodges CS. 1975. Eucalyptus microfungi. Two new hyphomycetes genera from Brazil.

Nova Hedw. 26: 527-533.

MY COTAXON

http://dx.doi.org/10.5248/119.249

Volume 119, pp. 249-253 January-March 2012

A new species of [juhya, |. oenanthicola

CHRISTIAN LECHAT? & MICHEL HAIRAUD?

"Ascofrance, 64 route de Chizé, F-79360 Villiers en Bois, France

"Impasse des Marroniers F-79360 Poivendre de Marigny, France

* CORRESPONDENCE TO : lechat@ascofrance.fr

ABSTRACT — Ijuhya oenanthicola sp. nov. (Bionectriaceae) is described and illustrated based

on a collection from Oenanthe crocata (Apiaceae) in France. The Acremonium-like asexual

state was obtained in culture.

KEY worps — Ascomycota, Hypocreales

Introduction

The situation of Penvenan on the North Coast of Brittany is a privileged one

for both its marvellous landscapes and temperate climate. Frost is very rare

in winter and summer heat is never overwhelming. Wet zones and marshes

have been preserved around the mouths of rivulets, surrounded by rows of

old Salix trees. Located between Queffioec and the “Anse de Pellinec,’ such a

marsh has proved to be a paradise for fungal diversity. Looking for Helotiales

on previous year plants or herbaceous stems, our attention was drawn to a tiny,

hairy hypocrealean fungus on dead Oenanthe remnants. This collection was

determined to represent a previously undescribed species. Our specimen was

cultured from a single ascospore that produced an Acremonium-like asexual

state.

Materials & methods

Specimens were examined using the methods described by Rossman et al. (1999).

Microscopic observations and measurements were made in water, and the ascospore

ornamentation was observed in cotton blue in lactic acid.

Taxonomy

Tjuhya oenanthicola Lechat & Hairaud, sp. nov. PLATE 1

MycoBank MB 561714

250 ... Lechat & Hairaud

Perithecia subglobosa, apice applanata, ca. 160-235 um diam, subhyalina vel aurantia,

corona subapicalis pilis agglutinatis aurantia, crasse-tunicatis, flexuosis composita, colore

in KOH non mutanda. Asci ca. 51 x 9.5 um, octospori, unitunicati. Ascosporae fusiformes,

ca. 15.4 x 3.2 um uniseptatae, striatae.

Type: France. Brittany: Cétes-d’Armor, Penvenan, Marais de Pellinec, 48°49'53.36"N

3°17'27.23" W, on dead stem of Oenanthe crocata L. (Apiaceae), 12 Mar 2011, leg. Michel

Hairaud, CLL11046 (Holotype, LIP; ex-type culture CBS 129747).

ErymMo.ocy: The epithet refers to the substratum on which this species was collected.

PERITHECIA Scattered on stem of Oenanthe, superficial, subglobose, (160-)

200-220(-235) um diam, at first white, becoming pale orange to dark orange,

collapsing laterally, not changing color in 3% KOH or lactic acid, with orange,

fasciculate, thick-walled hairs, agglutinated to form teeth arranged in a stellate

fringe around upper margin of perithecia, teeth up to 60 um wide at base. Hairs

flexuous, thick-walled, 20-150(-165) um long, 2-3 um wide, rounded at tip,

hyaline, smooth, cylindrical with wall 1-1.7 um thick, aseptate. PERITHECIAL

WALL 18-23 um thick, composed of two regions: outer region 10-14 um wide,

of globose to ellipsoidal, 3.5-6 x 2-3.5 um cells, with yellow to orange wall

1.5-2.2 um thick; inner region 5-8 um wide, of elongate, flattened cells 5-11

x 1.5-2 um, with hyaline wall 0.5-1 um thick. Asc (40-)45-55(-60) x (6-)

8-10(-12) um ("= 51 x 9.5 um, n = 20), clavate, apices flat to rounded, without

ring, containing irregularly biseriate ascospores, completely filling each ascus

when 8-spored, numerous asci in which 1-3 or even 4 of 8 ascospores aborted.

ASCOSPORES (12-)14-16.5(-18) x (2.5-)3-3.5(3.8) wm (™ = 15.4 x 3.2 um, n = 30),

fusiform, rounded at ends, 1-septate, hyaline, striate, constricted at septum.

Anamorph: Acremonium-like

In Cutture: After two weeks at 25°C on Difco PDA containing 5mg/L

streptomycin, colony 3-3.5 cm diam, spreading a reddish brown coloration

in medium, mycelium white to pale yellow, producing an Acremonium-like

culture at margin of colony, composed of monophialidic conidiophores, 42-55

um long, 3-4 um diam with 1-septum at base, arising from smooth hyphae

2.5-3.8 um diam, producing cylindrical to widely ellipsoidal conidia (3.5-)

4-6.5(-7) x 2-4 um (™ = 6.1 x 3.2 um, n = 30), hyaline, smooth, non-septate,

with a basal abscission scar.

Key to /juhya species with fasciculate hairs

(modified from Lechat & Courtecuisse 2010)

1. Hairs 200-300 um long; ascospores (24—)30-60(-110) x 4-7(-8) um, striate;

ascomata pale yellow ... I. peristomialis (Berk. & Broome) Rossman & Samuels

As Rains AVerAS IO <2 OOO NG hax siete te ccd gta ld x semaine ost Neral na! ty vated) dw EU 2

Zi NSCOSPOLES AV EL ACTG al IMU ON Ges es iu met ah Al Ali Bley Ale UM hela Raiceleca Ye 3

JASCOSPOLeS:AVeTagMore TZ MIT ONG: ah seta h c.tlen eet eutatlon aeeetien adie evkae ate ener a’ 4

Tjuhya oenanthicola sp. nov. (France) ... 251

NA wg

— \

ESS

PLATE 1. Ijuhya oenanthicola (based on holotype material). a. Perithecia. b. Fasciculate hairs.

c. Median section of perithecial wall. d. Asci. e. Ascospores. Scale bars: a = 200 um; b = 50 um;

c= 15 um; d-e = 10 um. Additional photos at http://www.ascofrance.fr.

252 ... Lechat & Hairaud

3. Ascospores (8.5-)9.5-11.5(-12.5) x 2.8-3.2(-3.5) um, striate;

ascomata pale yellow to brownish-orange,

hairs 28-80 x 2-2.5(-3) um ............0-. I. equiseti-hyemalis Lechat & Baral

3. Ascospores 6-8(-9) x 3-4 tum, spinulose; ascomata orange-yellow,

hairs 150-200 x 3-4 um ........... I. dentifera (Samuels) Rossman & Samuels

A. ASCOSPORES SETI ALE sc. sgAticca stills: Soste lege ersten lin: osteo sua eae Arses Meee ay we Asan Salata gee uate ae 5

4, Ascospores spinulose 14.5-20 x (2.5-)3-5(-5.4) um, ascomata brownish-orange,

HairS BOSOM LOMB acetyl ee, seoosta 5 ott I. parilis (Syd.) Rossman & Samuels

5. Asci (4—6—)8-spored, ascospores (12—)14—-16.5(-18) x (2.5-)3-3.5(3.8) tum;

hairs 20-140 um x 2.5-3 um, non-septate, ascomata dark orange I. oenanthicola

SF ASCILCONSIANtly S2SPOLEM. lok. ivan. astdirls wiikte y plete e dette 5 stllte ¥ agtlbe wht Aa dabe ts 6

6. Ascospores (10.5—)11-13(-14) x 2.5-3.5 um; ascomata dark orange,

haiesup:torkoO: pina lOnor oo ioe ee hts oe I. antillana Lechat & Courtec.

6. Ascospores (19-)21-28 x 3.5-4.5um; ascomata dull orange,

hairs up to 100 um long............... I. chilensis (Speg.) Rossman & Samuels

Discussion

In the genus [juhya Starback (Bionectriaceae) based on the type I. vitrea

Starback, ten species were recognized by Rossman & Samuels (1999). Since

then, Ijuhya hongkongensis J. Frohl. & K.D. Hyde (Frohlich & Hyde 2000),

I. hubeiensis Y. Nong & W.Y. Zhuang (Zhuang et al. 2007), I. equiseti-hyemalis

(Lechat & Baral 2008), and I. antillana (Lechat & Courtecuisse 2010) were

added.

Tjuhya oenanthicola differs from other Ijuhya species bearing fasciculate hairs

in having < 200 um long non-septate hairs, > 14 um long striate ascospores,

numerous asci with 1-3 (or even 4) out of 8 ascospores aborted, and its

occurrence on Oenanthe crocata.

Acknowledgments

We thank Dr. Amy Rossman, Systematic Mycology & Microbiology Laboratory

USDA-ARS, Beltsville, MD, USA, and Jacques Fournier, Las Muros, 09420 Rimont,

France, for their reviews of this manuscript. We hereby express our gratitude to Mrs

Brigitte Capoen for her kindness in providing further collections from the same locality.

Finally, we thank the Fondation Langlois for its assistance.

Literature cited

Frohlich J, Hyde KD. 2000. Palm microfungi. Fungal Diversity Research Series 3: 1-393.

Lechat C, Baral HO. 2008. A new species of Ijuhya on Equisetum hyemale and its Acremonium anamorph,

with notes on Hydropisphaera arenula. Ost. Z. Pilzk. 17: 15-24.

Lechat C, Courtecuisse R. 2010. A new species of [juhya, I. antillana, from the French West Indies.

Mycotaxon 113: 443-447. http://dx.doi.org/10.5248/113.443

Rossman AY, Samuels GJ, Rogerson CT, Lowen R. 1999. Genera of Bionectriaceae, Hypocreaceae

and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1-248.

Tjuhya oenanthicola sp. nov. (France) ... 253

Samuels GJ. 1976. Perfect states of Acremonium. The genera Nectria, Actiniopsis, Ijuhya,

Neohenningsia, Ophiodictyon, and Peristomialis. New Zealand J. Bot. 14: 231-260.

Zhuang WY, Nong Y, Luo J. 2007. New species and new Chinese records of Bionectriaceae and

Nectriaceae (Hypocreales, Ascomycetes) from Hubei, China. Fungal Diversity 24: 347-357.

MYCOTAXON

http://dx.doi.org/10.5248/119.255

Volume 119, pp. 255-260 January-March 2012

Two new species of Hyphoderma (Agaricomycetes) from India

PRIYANKA* & G.S. DHINGRA

Department of Botany, Punjabi University, Patiala 147 002, India

*CORRESPONDENCE TO: bansalpriyanka.6@gmail.com

ABSTRACT - Two new corticioid species, Hyphoderma bicystidiatum and H. subglobosum, are

described from Kangra Hills in Himachal Pradesh.

KEY worpbs - northwestern Himalaya, angiosperm host

While conducting fungal forays in district Kangra of Himachal Pradesh,

India, Priyanka collected two interesting corticioid fungi that were referred

to Hyphoderma based on detailed macroscopic and microscopic comparisons

(Eriksson & Ryvarden 1975, 1976; Martin & Gilbertson 1977; Rattan 1977;

Burdsall & Nakasone 1983; Dhingra 1989, Dhingra & Singla 1993, Natarajan

& Kolandavelu 1998, Dhingra & Singh 2009, Dhingra et al. 2009, Singh et al.

2010). The specimens could not be assigned to any known taxa and are here

described as two new species. Samples of both collections were sent to Prof.

Nils Hallenberg, who also supported the concept of referring the new species

to Hyphoderma.

Hyphoderma bicystidiatum Priyanka & Dhingra, sp. nov. Figs 1-7

MycoBank MB 560555

Differs from Hyphoderma pallidum in the presence of thick-walled clavate encrusted

cystidia.

Type: India, Himachal Pradesh: Kangra, Shiv Nagar, on decaying angiospermous

branches, 25 August 2009, Priyanka 4298 (PUN, holotype).

Erymo oey: The epithet refers to the presence of two types of cystidia.

Basidiocarp resupinate, effused, adnate, up to 250 um thick in section; hymenial

surface smooth to somewhat tuberculate under the lens, yellowish white to pale

yellow; margins thinning out, concolorous but paler. Hyphal system monomitic;

generative hyphae up to 5.5 um wide, ordinarily branched, with clamps at all

256 ... Priyanka & Dhingra

RRR

are

rr SeaeF

RY re

are

Fics 1-6. Hyphoderma bicystidiatum: microscopic structures.

1. vertical section through basidiocarp; 2. basidiospores; 3. basidia; 4. generative hyphae;

5. smooth cystidium; 6. thick-walled encrusted cystidium.

septa, thin-walled; hyphae loosely arranged and parallel to the substrate in the

basal region and gradually becoming vertically oriented towards the hymenium.

Cystidia of two kinds: (i) 82.0-122.0 x 12.3-13.8 um, tubular, basally widened

and somewhat thick-walled, apically thin-walled, smooth; (ii) 69.0-110.0 x

11.0-13.8 um, clavate to subclavate, thick-walled, encrusted in the apical third

part. Basidia 13.6-22.3 x 6.2-7.5 um, clavate to subclavate, 4-sterigmate, with

oily contents and basal clamp; sterigmata up to 4.8 um long. Basidiospores

6.2-8.8 x 3.0-4.1 um, subellipsoid to suballantoid, thin-walled, smooth,

inamyloid, acyanophilous, with oily contents.

REMARKS—Hyphoderma bicystidiatum resembles H. argillaceum and H. pallidum

in having basally widened cystidial elements and resinous matter in the

Hyphoderma spp. nov. (India) ... 257

Fic. 7. Hyphoderma bicystidiatum: basidiocarp showing hymenial surface.

context. The new species is also similar to H. pallidum in the shape of

basidiospores. It differs from both, however, in having clavate, thick-walled,

encrusted, cystidia. The new combination of features suggests a species of its

own.

Hyphoderma subglobosum Priyanka & Dhingra, sp. nov. Fics 8-13

MycoBank MB 560581

Differs from Hyphoderma clavigerum in subglobose basidiospsore shape.

Type: India, Himachal Pradesh: Kangra, about 3 km from Nurpur towards Sulyali, on

decaying angiospermous branches, 26 August 2009, Priyanka 4299 (PUN, holotype).

Erymo toy: The epithet refers to the shape of basidiospores.

Basidiocarp resupinate, effused, adnate, up to 170 um thick in section; hymenial

surface smooth to somewhat tuberculate under the lens, grayish yellow to

grayish orange; margins thinning out, indeterminate, concolorous but paler.

Hyphal system monomitic; generative hyphae up to 5.6 um wide, ordinarily

branched, with clamps at all septa, thin-walled; hyphae almost parallel to the

substrate in the basal zone, gradually becoming vertical and densely intertwined

in the subhymenial and hymenial zones. Cystidia 50.0-69.0 x 8.0--11.8 um,

clavate, smooth, thin-walled. Basidia 22.0-34.0 x 6.8-10.0 um, clavate to

subclavate, mostly constricted at the middle, with oily contents, 4-sterigmate,

258 ... Priyanka & Dhingra

Raed jy @9C

VAS) i WZ

Lx) 3 4 i J

YNOGALE:

rae See, << L—

— \VE Fes

- 10 um

Fics 8-12. Hyphoderma subglobosum: microscopic structures.

8. vertical section through basidiocarp; 9. basidiospores; 10. basidia;

11. cystidia; 12. generative hyphae.

with basal clamp; sterigmata up to 6.8 um long. Basidiospores 11.8-13.1 x

8.0-10.0 um, subglobose to broadly ellipsoid, thin-walled, smooth, inamyloid,

acyanophilous, with grainy contents.

REMARKS— The subglobose spores and the clavate cystidia with light refracting

content are diagnostic characters. Hyphoderma subglobosum is close to

H. clavigerum in having similarly shaped cystidial elements but differs

considerably in the shape of basidiospores i.e. subglobose in comparison to

Hyphoderma spp. nov. (India) ... 259

pe) Levante, Dal

EP YEE SOS

¥,

Fic. 13. Hyphoderma subglobosum: basidiocarp showing hymenial surface.

suballantoid. The combination of subglobose spores and clavate cystidia with

light refracting content suggests that the collection represents a new species.

Acknowledgements

The authors thank Prof. Nils Hallenberg (Gothenburg, Sweden) for valuable

suggestions and peer review; Prof. B.M. Sharma, Department of Plant Pathology, COA,

CSKHPAU, Palampur, H.P., India for peer review; Head, Department of Botany, Punjabi

University, Patiala, is thanked for providing research facilities.

Literature cited

Burdsall HH Jr, Nakasone KK. 1983. Species of effused Aphyllophorales (Basidiomycotina) from the

Southeastern United States. Mycotaxon 17: 253-268.

Dhingra GS. 1989. Genus Hyphoderma Wallr. em Donk in the Eastern Himalayas. 197-212, in: ML

Trivedi et al. (eds). Plant Science Research in India. Today & Tomorrow’s Printers & Publishers,

New Delhi.

Dhingra GS, Singh Avneet P. 2009. Diversity of Family Hyphodermataceae (Polyporales,

Agaricomycetidae) in Himachal Pradesh. 171-189, in: NS Atri et al. (eds). Germplasm Diversity

& Evaluation - Algae, Fungi & Lichens. Bishen Singh Mahendra Pal Singh, Dehradun, U.K.

(India).

Dhingra GS, Singla Nishi. 1993. Studies in North-West Himalayan Corticiaceae (Basidiomycetes)

~ I. Some interesting species from Dalhousie Hills, J. Ind. Bot. Soc. 72: 29-33.

Dhingra GS, Singh Avneet P, Singla Nishi. 2009. A new species of Hyphoderma (Basidiomycetes)

from India. Mycotaxon 108: 197-199. http://dx.doi.org/10.5248/108.197

Eriksson J, Ryvarden L. 1975. The Corticiaceae of north Europe - III. Fungiflora, Oslo pp.

287-546.

260 ... Priyanka & Dhingra

Eriksson J, Ryvarden L. 1976. The Corticiaceae of north Europe - IV. Fungiflora, Oslo pp.

549-886.

Martin KJ, Gilbertson RL. 1977. Synopsis of wood-rotting fungi on spruce in North-America - I.

Mycotaxon 6: 43-77.

Natarajan K, Kolandavelu K. 1998. Resupinate Aphyllophorales of Tamil Nadu, India. Centre for

Advance study in Botany, University of Madras, Chennai: 133 p.

Rattan SS. 1977. The resupinate Aphyllophorales of the northwestern Himalayas. Bibliotheca

Mycologica 60: 1-427.

Singh Avneet P, Priyanka, Dhingra GS, Singla Nishi. 2010. A new species of Hyphoderma

(Basidiomycetes) from India. Mycotaxon 111: 71-74. http://dx.doi.org/10.5248/111.71

MY COTAXON

http://dx.doi.org/10.5248/119.261

Volume 119, pp. 261-267 January-March 2012

New records of noteworthy gasteroid fungi from Pakistan

N. YousaAk”*, A.R. NIAZI & A.N. KHALID

Department of Botany, University of the Punjab, Quaid-e-Azam Campus, Lahore, 54590, Pakistan

* CORRESPONDENCE TO: nousheenyousaf@gmail.com

ABSTRACT — Gasteroid fungi were collected during the course of fieldwork in Pakistan.

Here, we report on two species, Bovistella japonica and Lycoperdon excipuliforme, from the

Himalayan moist temperate forest of the Khyber Pakhtunkhwa Province, each a new record

for the country. Phellorinia herculeana represents a new record from Cholistan Desert of the

Punjab Province. For each taxon reported, detailed macro- and microscopic descriptions are

presented.

Key worps — Biomes, Nathia gali, taxonomy

Introduction

The wide range of habitats in Pakistan that extends from coastal regions

along the Arabian Sea to arid scrubland in the Punjab region and to the high

altitude forests and tundra of the Himalayas supports a rich diversity of fungi.

One group, the gasteroid fungi, has been documented from the nation in

several publications (Long & Ahmad 1947, Ahmad 1952, Yoshimi & Hagiwara

1992, Khalid & Iqbal 1996, Ahmad et al. 1997, Sultan et al. 2001, Razzaq &

Shahzad 2004, 2007, Iqbal et al. 2006, Sultana et al. 2007, Moreno et al. 2009).

In total, 112 species representing approximately 30 genera have been reported

from Pakistan. Although a considerable number of taxa are known, it is likely

that the present accounting of Pakistani gasteroid fungi is far from complete,

as a large portion of the country has been sampled only in a cursory manner,

if at all.

In this study, we collected gasteroid fungi in two different biomes of Pakistan

— the moist temperate forest of the Himalayas and scrubland of the Cholistan

Desert. The Himalayan moist temperate forest extends from the Murree Hills of

the Ayubia National Park to the Swat Valley (Khyber Pakhtunkhwa province),

along the outer range of the Himalayas at an altitude of around 1850-2500 m

(Ahmad et al. 2006). The Cholistan Desert, which covers a 26,000 km? area

262 ... Yousaf, Niazi & Khalid

in southwest Punjab province, is among the hottest and driest regions in the

country (Akbar et al. 1996).

Here we report on three gasteroid species from Pakistan: Bovistella japonica

and Lycoperdon excipuliforme were collected in the Himalayan moist temperate

forest and represent new records for the country, while Phellorinia herculeana

(previously reported from Pakistan as P. inquinans) represents a new record for

the Cholistan Desert.

Materials & methods

Observations of microscopic characters (e.g., spore dimensions, eucapillitium,

peridial hyphae) were made under a light microscope (at 1000x magnification) on

material mounted in lactophenol and 5% KOH medium. Glebal color and macroscopic

peridial features were observed using Meiji stereomicroscope, and illustrations were

made with the help of a camera lucida.

The specimens have been deposited in gasteroid fungal collection of LAH Herbarium,

Department of Botany, University of the Punjab, Lahore, Pakistan.

Taxonomy

Bovistella japonica Lloyd, Mycol. Writ. 2: 281 (1906) Fic. 1

GASTEROCARP 55-85 mm diam. x 85 mm high, globose to subglobose, white

when young, turning reddish brown with age. STERILE BAsE well developed, up

to 30 mm high, spongy, chambered, basally attached to the substratum by well

developed, thick mycelial strand, having white, branched rhizomorphs heavily

encrusted with soil particles; ostiole lacking; dehiscence irregular, fruiting

body breaking apart apically. PERIpIuM < 1mm thick. ExOPpERIDIUM white

when young, becoming dull brown with age, thin, formed of granular matter

which covers the endoperidium, persistent. ENDOPERIDIUM entirely covered by

exoperidium even in older specimen, light brown, thin, papery. GLEBA brown,

cottony, compact.

BASIDIOSPORES subglobose to ovoid, 4.0-5.5 x 3.8-4.7 um, hyaline to greenish

in KOH, pedicellate, pedicels short to very long, up to 19 um long. CAPILLITIUM

Bovista type, well developed, separate units having a thick main axis (< 20 um

diam.), thick walled (walls < 2.85 um), branches originating from the main axis,

1.2-6.0 um diam. (walls < 1.0 um), with attenuate tips. EXOPERIDIUM composed

of subglobose to irregular sphaerocysts. ENDOPERIDIUM composed of hyaline,

unbranched, and aseptate hyphae. STERILE BASE composed of pigmented, thick

walled, branched, and aseptate hyphae.

MATERIAL EXAMINED: PAKISTAN: KHYBER PAKHTUN Kuwa, Nathia Gali, ca. 2500m

(8205 ft) a.s.l., solitary, on ground, under Pinus wallichiana A.B. Jacks., in Himalayan

moist temperate forest, 13 Sep. 2006, N. Yousaf NYG208 (LAH 130906).

Gasteroid fungi from Pakistan ... 263

Fia. 1: Bovistella japonica. A. Young gasterocarp. B. Mature gasterocarp. C. Pedicellate basidiospores.

D. Typical Bovista type capillitial units. E. Exoperidial elements (sphaerocysts). F. Thick, main

axis of capillitial threads. G. Aseptate endoperidial hyphae. H. Hyphae of sterile base. Scale bars:

A, B=1.5 cm; C = 2.5 um; D = 60 um; E = 3 um; F = 20 um; G, H = 10 um.

ComMENTs: Bovistella was first proposed by Morgan in 1892, and the genus

is distributed worldwide. Bovistella japonica is characterized by the presence

264 ... Yousaf, Niazi & Khalid

of short, separate capillitial units, pedicellate spores, and sterile base. The

combination of Bovista-like capillitial units and Lycoperdon-like sterile base

places the species in the intermediate genus, Bovistella, along with B. ohiensis

(Ellis & Morgan) Morgan, B. radicata (Durieu & Mont.) Pat., B. lycoperdoides

(Schwein.) Lloyd, B. paludosa (Lév.) Pat., and B. ammophila (Lév.) Lloyd.

Among these taxa, B. japonica is more closely allied with B. ohiensis and

B. paludosa, which also share the same general spore dimensions (~4—-5 tm).

Bovistella japonica can be distinguished by its more compact sterile base. This

species is a new addition to the mycoflora of Pakistan.

Lycoperdon excipuliforme (Scop.) Pers., Syn. meth. fung.: 143 (1801) Fic. 2

GASTEROCAR? elliptical to turbinate, 30-50 mm diam. x 50 mm high; spore

case broad, globose to subglobose; ostiole present, irregular. STERILE BASE

present, and forming a well-developed pseudostipe, attached to the substrata

by well developed rhizomorphs, these white, branched, thick, and heavily

encrusted with soil particles, and woody debris. PER1IprIuM layered, thickness

< 1 mm. Exoperipium dull brown, fragile, in the form of granules which later

slough off. ENDOPERIDIUM dull, golden brown, papery. GLEBA dark brown,

pulverulent.

BASIDIOSPORES globose to subglobose, dark brown, 5.3-6.7 um diam.

(4.4-5.5 um excluding ornamentation), verrucose (verrucae < 1 um),

numerous sterigmal remnants present in mounts, these up to 35.6 um long.

CAPILLITIUM Lycoperdon-type, capillitial threads 1.14-6.8 um diam., branched,

(dichotomous when occurring), thick-walled (walls < 2 um), sub undulated to

undulate, aseptate, having slit-like pores, with attenuate tips. PARACAPILLITIUM

absent. ExOPERIDIUM composed of subglobose sphaerocysts. ENDOPERIDIUM

composed of unbranched, septate hyphae.

MATERIAL EXAMINED: PAKISTAN: KHYBER PAKHTUN Kuwa, Nathia Gali, ca. 2500m

(8205 ft) a.s.L, solitary, on ground, under bamboo tree, in Himalayan moist temperate

forests, 26 Aug. 2006, N. Yousaf NYG209 (LAH 268006).

ComMENTs: Commonly called the “pestle puffball” L. excipuliforme usually

fruits in groups from summer to autumn in coniferous and deciduous

woodlands (Jordan 2006). It is characterized by large basidiomes consisting

of an upper spore case and long pseudostipe and by the presence of slit-like

pores in its capillitial threads. Lycoperdon perlatum Pers., which also develops

the same reticulate pattern over the endoperidial surface as the larger verrucae

slough off, differs in its smaller stature and the absence of a well-developed

elongated pseudostipe. Lycoperdon excipuliforme is new to the mycoflora of

Pakistan.

Gasteroid fungi from Pakistan ... 265

[Fae tas F

Fic. 2: Lycoperdon excipuliforme. A. Gasterocarps. B. Ostiole (arrow). C. Verrucose basidiospores.

D. Capillitial threads. E. Exoperidial elements (sphaerocysts). F. Endoperidial hyphae. Scale Bars:

A, B = 0.8 cm; C = 2.5 um; D, F = 10 um; E = 3 pm.

Phellorinia herculeana (Pers.) Kreisel, Ceska Mykol. 15: 196 (1961) Fic. 3

= Phellorinia inquinans Berk., London J. Bot. 2: 521 (1843)

GASTEROCARP consisting of spore case, stipe, and bulbous base, < 85 mm

tall; spore case < 50 mm diam. x 35 mm high, basal portion encrusted with soil

particles; dehiscing as apical portions wear away to expose the powdery spore

mass. STIPE woody, continuous with spore case, not uniform, having a wide,

266 ... Yousaf, Niazi & Khalid

Fic. 3: Phellorinia herculeana. A. Gasterocarps. B. Verrucose basidiospores. C. Endoperdial hyphae.

D. Exoperidial hyphae with globular elements. Scale Bars: A = 1.2 cm; B = 3 um; C, D = 10 um.

bulbous base, and tapering toward the apex, with parallel, broad ridges and

furrows running from apex to base, and permanent scales scatter intermittently;

rhizomorphs not observed. Exopertpium off-white when immature, pale

yellowish to brown with age, thick, entirely covered with hard scales of variable

sizes. ENDOPERIDIUM continuous with stipe, pale yellow. GLEBA pulverulent,

brown.

BasIpIosPporeEs globose to subglobose, apedicellate, hyaline, 4.5-8.0 um

diam., verrucose (verrucae < 1 um long). CAprLiitrum hyaline, rarely present

in mounts. EXOPERIDIUM composed of branched, hyaline hyphae, with inflated,

globose elements at their tips. ENDOPERIDIUM composed of thin, hyaline,

branching, septate hyphae.

MATERIAL EXAMINED: PAKISTAN: Punjab, Bahawalpur, Rahim Yar Khan, ca. 115 m

(380 ft) a.s.l., Cholistan Desert, solitary, on ground, 29 Sep. 2006, N. Yousaf NYG210

(LAH 290900).

ComMENTs: Phellorinia was erected by Berkeley in 1843 with P inquinans

Berk. as the type species. It is a monotypic genus that has been reported from

Gasteroid fungi from Pakistan ... 267

different regions of Pakistan by Ahmad (1952). Phellorinia herculeana is widely

distributed across arid to semi-arid regions of all continents, including countries

such as Australia (Fuhrer 2005, Williams & Woinarski 1997), Pakistan (Ahmad

1952, Ahmad et al. 1997), Spain, and Yemen (Kreisel & Fatimi 2004).

Acknowledgements

The authors wish to thank Dr. Taiga Kasuya of the Laboratory of Plant parasitic

Mycology, Graduate School of Life and Environmental Sciences, University of Tsukuba,

Japan for helping in the identification of these gasteroid fungi. We also thank Dr. Scott.

T. Bates of the Cooperative Institute for Research in Environmental Sciences, University

of Colorado at Boulder, U.S.A., and Ms. Larissa Trierveiler Pereira in the Department

of Botany, Biosciences Institute, UFRGS, Brazil for their valuable comments, and for

critically reviewing the manuscript.

Literature cited

Ahmad S. 1952. Gasteromycetes of West Pakistan. Punjab University Press. Lahore.

Ahmad §S, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Sultan Ahmad Mycological Society of

Pakistan, Lahore.

Ahmad M, Husain T, Sheikh AH, Hussain SS, Siddiqui MF. 2006. Phytosociolgy and structure of

Himalayan Forests from different climatic zones of Pakistan. Pak. J. Bot. 38(2): 361-383.

Akbar G, Khan TN, Arshad M. 1996. Cholistan desert, Pakistan. Rangelands 18(4): 124-128.

Fuhrer B. 2005. A field guide to Australian fungi. Blooming Books Pvt. Ltd. Science. 360 p.

Iqbal SH, Kasuya T, Khalid AN, Niazi AR. 2006. Lysurus pakistanicus, a new species of Phallales

from Pakistan. Mycotaxon 98: 163-168.

Jordan P. 2006. Field guide. Edible mushrooms of Britian and Europe. New Holand Publishers

(UK) Ltd. London.

Khalid AN, Iqbal SH. 1996. New Gasteromycetes from Pakistan. Sultania 1(2): 97-102.

Kreisel H, Fatimi MA. 2004. Basidiomycetes and larger Ascomycetes from Yemen. Feddes

Repertorium 115(7-8): 547-561. http://dx.doi.org/10.1002/fedr.200411053.

Long WH, Ahmad S. 1947. The genus Tylostoma in India. Farlowia 3(2): 225-267.

Moreno G, Khalid AN, Alvarado P. 2009. A new species of Phallus from Pakistan. Mycotaxon 108:

457-462. http://dx.doi.org/10.5248/108.457.

Razzaq A, Shahzad S. 2004. Pisolithus tinctorius, a new record from Pakistan. Pak. J. Bot. 36(2):

449-451.

Razzaq A, Shahzad S. 2007. Geastrum sessile and G. vulgatum, new records from Pakistan. Pak. J.

Bot. 39(6): 2193-2194.

Sultan A, Nazir K, Ayub N. 2001. Some common species of fleshy macromycetes (Discomycetes,

Gasteromycetes and Agaricales) from Margalla hills national park and adjacent areas. Pak. J. Bot.

33(special issue): 709-721.

Sultana K, Gul M, Firdous SS, Asghar R. 2007. Hymenomycetes from Multan District. Pak. J. Bot.

39(2): 651-657.

Williams JE, Woinarski J. 1997. Eucalypt ecology: individuals to ecosystems. Cambridge University

Press. United Kingdom.

Yoshimi S, Hagiwara H. 1992. Gasteroid fungi from Pakistan. 165-184, in: T Nakaike, S Malik

[eds]. Cryptogamic Flora of Pakistan, Vol. I. National Science Museum Tokyo.

MY COTAXON

http://dx.doi.org/10.5248/119.269

Volume 119, pp. 269-275 January-March 2012

Pseudocercosporella stellariicola, a new hyphomycete

occurring on Stellaria aquatica

MI-JEONG PARK, JI-HYUN PARK & HYEON-DONG SHIN"

Division of Environmental Science and Ecological Engineering,

College of Life Sciences and Biotechnology, Korea University, Seoul 136-701, Korea

* CORRESPONDENCE TO: hdshin@korea.ac.kr

ABSTRACT — A new cercosporoid fungus, Pseudocercosporella stellariicola, was collected

on leaves of Stellaria aquatica (Caryophyllaceae). This fungus is described, discussed, and

compared with other similar hyphomycetes occurring on Stellaria and related genera:

Cercosporella echinulata, Phacellium episphaerium, Pseudocercosporella cerastii, and

Ps. woronovii. The phylogenetic relationship of Ps. stellariicola and other Pseudocercosporella

species within Mycosphaerellaceae is also inferred from the ITS rDNA sequence data.

KEY worpDs — anamorphic Mycosphaerella, imperfect fungi, plant pathogen

Introduction

As a result of extensive studies conducted on the mycobiotas of cercosporoid

fungi in Korea from 1990 to 2000, Shin & Kim (2001) published a monograph

comprising 127 species in 14 genera on 151 species of host plants. Although

Stellaria aquatica, one of the most common wild plants in Korea, was frequently

found infected with Phacellium episphaerium, a previously undescribed leaf-

spotting hyphomycete has also been found on this plant since 2004. ‘This

fungus differed from other similar hyphomycetes known to occur on Stellaria

and related genera. It was well characterized by colorless conidiophores in

dense fascicles, inconspicuous, unthickened conidial scars and solitary, hyaline

conidia. These features agree with the concept of Pseudocercosporella proposed

by Braun (1990), and this hyphomycete is identified and characterized as a

novel species in that genus.

Materials & methods

For microscopy, free-hand sections of fresh materials were mounted in water.

Morphological characteristics were measured using a model BX51 microscope

(Olympus, Tokyo, Japan), and photographed using an Axio imager microscope (Carl

270 ... Park, Park & Shin

Zeiss, GOttingen, Germany). The voucher specimens were deposited in the Korea

University herbarium (KUS).

Monoconidial isolates were deposited at the Korean Agricultural Culture Collection

(KACC) ofthe National Academy of Agricultural Science in Koreaand the Centraalbureau

voor Schimmelcultures (CBS) in the Netherlands. Genomic DNA was extracted using

mycelia harvested from colonies following a previously described method (Lee & Taylor

1990). The ITS rDNA region was amplified using universal primers ITS1/ITS4 (White

et al. 1990). The PCR amplicons were purified using a MultiScreen HTS™ PCR filter

plates (Millipore, Carrigtwohill, Cork, Ireland) and sequenced using an ABI Prism™ 377

automatic DNA sequencer (Applied Biosystems, Foster City, CA, USA) with a BigDye™

cycle sequencing kit v. 3.1 (Applied Biosystems). The raw sequences were edited using

the DNASTAR computer package v. 5.05 (Lasergene, Madison, WI, USA). The ITS rDNA

sequences obtained from three isolates (KACC42363, KACC42395, and CPC11297)

have been deposited in GenBank with accession numbers EF600945, EF600956, and

GU214693, respectively. For phylogenetic analysis, available ITS rDNA sequences of

Pseudocercosporella spp. and closely related taxa belonging to Mycosphaerellaceae were

retrieved from GenBank. The sequences were aligned with ClustalX v. 2.1 (Larkin

et al. 2007). The Bayesian inference analysis was performed with MrBayes v. 3.1.2

(Huelsenbeck & Ronquist 2001). The GTR + G as nucleotide substitution model was

selected using jModeltest v. 0.1.1 (Posada 2008). One million generations with four

chains were run and a tree every 100th generation was saved. The first 1000 of the 10,000

saved trees were discarded as burn-in. The resulting 50% majority-rule consensus tree

was Visualized with TreeViewX (Page 1996). Branch support was calculated as posterior

probability (PP).

Taxonomy

Pseudocercosporella stellariicola M.J. Park, J.H. Park & H.D. Shin, sp.nov. Fic. 1

MycoBank MB 563164

Similar to Pseudocercosporella cerastii, but with wider (2-3 um) conidia and distinct leaf

lesions. Similar to P woronovii, but with shorter (40-80 um), straight to slightly curved,

1-3(-4)-euseptate conidia.

Type: On living leaves of Stellaria aquatica (L.) Scop. (Caryophyllaceae), KOREA,

NAMYANGJU, Experimental Farm of Korea University, 37°35'01"N 127°14'13"E, 3 May

2006, H.D. Shin & M.J. Park (Holotype, KUS-F21740; culture ex-type, KACC42363).

ErymMo ocy: The epithet is derived from the host plant genus.

LEAF SPOTS amphigenous, circular to subcircular, 2-7 mm diam., whitish to

pale brown. CaEsPITULI amphigenous, grayish-brown, punctiform, evenly

and more or less densely distributed over the spot. MyceLium internal;

hyphae septate, branched, hyaline, 3-5 um wide; superficial mycelium absent.

STROMATA well-developed, erumpent, subcuticular or intraepidermal,

sometimes substomatal, consisting of medium brown, pseudoparenchymatal

cells, ca. 30-100 um wide and 25-65 um high. CoNIDIOPHORES numerous

in a dense divergent fascicle, hyaline throughout, aseptate, straight, conic to

Pseudocercosporella stellariicola sp. nov. (Korea) ... 271

Fic. 1. Pseudocercosporella stellariicola. A: Symptoms on leaves of Stellaria aquatica. B: Close-up of

the leaf lesions showing many stromata on the infected tissue. C: Upper view of a stroma emerging

from a stoma of the host plant. D: Vertical view of a stroma protruding from the cuticle of the

host plant. E: Small stroma with a few conidiophores emerging through the substomatal cavity.

F: Conidia. G: Close-up of conidia. H: Six-week-old colonies formed on potato dextrose agar.

Scale bar: C-D = 50 um, E-F = 30 um, G = 20 um.

272... Park, Park & Shin

mildly geniculate-sinuous, 8-15(-20) um in length, 2-3 um in width; conidial

scars unthickened, not darkened. Conrp1a solitary, filiform, straight to slightly

curved, hyaline, 1-3(-4)-euseptate, non-constricted at the septa, irregularly

guttulate, obtuse at the apex, subtruncate to subobtuse at the base, 40-80 x

2-3 um; hilum unthickened, not darkened. CoLonizs on PDA attaining ca. 10

mm diam after 6 weeks at 25°C under 12-h fluorescent light, dark brownish

to blackish, somewhat crumpled, consisting of a slimy mass due to abundant

sporulation, producing sparse aerial mycelia. TELEOMORPH unknown.

ADDITIONAL SPECIMENS EXAMINED — KOREA, YANGPYEONG, Experimental Forest of

Korea University, 37°30'12"N 127°41'55"E, 5 May 2004, H.D. Shin, KUS-F20158; 26

May 2004, H.D. Shin, KUS-F20239 (culture: CPC11297); 20 May 2005, H.D. Shin, KUS-

F21102; 25 May 2006, H.D. Shin & M.J. Park, KUS-F21808 (culture: KACC42395);

Cheongun-myeon, 37°32'04"N 127°44'48"E, 20 May 2007, H.D. Shin & M_.J. Park, KUS-

F22591 (culture: KACC42830); NAMYANGJU, Experimental Farm of Korea University,

37°35'01"N 127°14'13"E, 17 May 2004, H.D. Shin, KUS-F20216; 19 April 2008, H.D.

Shin & M.J. Park, KUS-F23281 (culture: KACC43763); 25 April 2011, H.D. Shin & J.H.

Park, KUS-F25669; TAEAN, Nam-myeon, 36°41'11"N 126°17'05"E, 17 April 2007, H.D.

Shin & M.J. Park, KUS-F22539; HONGCHEON, Sangoan-ri, 37°39'25"N 127°49'18"E, 28

April 2007, H.D. Shin & MJ. Park, KUS-F22565 (culture: KACC42827); Seongdong-

ri, 37°44'41"N 127°52'01"E, 29 April 2007, H.D. Shin & M.J. Park, KUS-F22578;

HoENGSEONG, Eupha-ri, 37°29'47"N 127°58'52"E, 29 April 2011, H.D. Shin & J.H. Park,

KUS-F25676; Seowon-myeon, 37°31'33"N 127°52'29"E, 6 May 2011, H.D. Shin, KUS-

F25679.

Discussion

Two cercosporoid hyphomycetes are presently known to occur on Stellaria

species. Cercosporella echinulata Garb., described on Myosoton aquaticum (L.)

Moench (= Stellaria aquatica), differs from the new species by conidiophores

with an echinulate, denticulate apical part. Additionally, C. echinulata possesses

much longer (35-110 um) conidiophores and narrower (1-2 um) conidia.

Although the identity of C. echinulata is unclear (according to Braun 1995), it

is easily differentiated from Ps. stellariicola. Phacellium episphaerium (Desm.)

U. Braun is also readily differentiated from the present species by its synnematous

fascicles of conidiophores and aseptate conidia.

Two Pseudocercosporella species are known on other caryophyllaceous host

plants. Pseudocercosporella cerastii (H.C. Greene) U. Braun (Braun 1995) known

on Cerastium spp. from Central Asia and North America is distinguished

from the new species by much narrower (0.75-2 um) conidia. The leaf lesions

associated with Ps. cerastii are almost absent or diffuse, whereas those of

Ps. stellariicola are quite distinct by being circular to subcircular and delimited

by a dark border.

Pseudocercosporella woronovii (Siemaszko) U. Braun recorded on Silene

spp. and Lychnis spp. in Europe (see Braun 1995) also produces (sub-)circular

lesions and amphigenous colonies on the leaves. However, Ps. woronovii differs

Pseudocercosporella stellariicola sp. nov. (Korea) ... 273

0.99 AY259139 Pseudocercosporella capsellae

073 GU214662 Pseudocercosporella capseliae

AY259140 Pseudocercosporella capsellae

0.55] +Q303091 Pseudocercosporella capsellae

0.561 0.54 AY259131 Ramulispora sorghi

GU214658 Cercosporella virgaureae

AY 152553 Mycosphaerella latebrosa

AY 489277 Septoria leucanthemi

AY 152574 Septoria apiicola

GU214699 Septoria dysentericae

0.99} EF535695 Pseudocercosporelia sp.

EF535696 Pseudocercosporella sp.

0.79 GU214683 Pseudocercosporella sp.

0.96/"rGU214684 Pseudocercosporella sp.

GU214686 Pseudocercosporella sp.

AY489285 Septoria obesa

1.00} |“ GU214685 Pseudocercosporella sp.

GU214698 Septoria cucubali

AY840527 Cercospora beticola

GU214655 Cercospora sojina

GU214653 Cercospora apii

EF600956 Pseudocercosporelila stellariicola

EF600945 Pseudocercosporella stellariicola

1.00 FKACC42827 Pseudocercosporella stellariicola

1.00 GU214693 Pseudocercosporeila stellariicola

0.99] FKACC43763 Pseudocercosporelia steilariicola

KACC42830 Pseudocercosporella stellariicola

4.00 JF77045 Mycosphaerella cerastiicola

0.92-°-GU214692 Ramularia coleosporii

1.00] -—-GU214694 Ramularia uredinicola

EU019284 Ramularia pratensis var. pratensis

GU214689 Ramularia acroptili

AF181692 Mycosphaerella graminicola

0.84-AF173314 Mycosphaerella africana

0.99] “-AY251069 Passailora fulva

0.81] p~AY808302 Dothistroma pini

ae i GU570542 Pseudocercosporella bakeri

1.00(DQ302952 Pseudocercosporella endophytica

DQ302953 Pseudocercosporella endophytica

AJ244261 Penidiella columbiana

1.00 AF393723 Devriesia staurophora

EU707862 Teratosphaeria fibrillosa

0.93 GU570539 Microcyclospora pomicola

GU214682 Pseudocercosporelia fraxini

AF393707 Davidiella tassiana

aeaoejjaioeydsoohyy

1.00 0.53

0.70

0.77

Fic. 2. A 50% majority-rule consensus tree from Bayesian analysis of the ITS rDNA sequences of

Pseudocercosporella spp. and closely related taxa belonging to Mycosphaerellaceae. Numbers above

the branches indicate the posterior probability values. The number of nucleotide changes between

taxa is represented by branch length and the scale bar corresponds to the number of substitution per

site. The sequences of Ps. stellariicola are shown in bold and a well-supported clade accommodating

them is highlighted in gray box.

274 ... Park, Park & Shin

from Ps. stellariicola by longer (50-100 um), straight to flexuous, obscurely

1-5-septate conidia.

The complete ITS rDNA sequence data of the six Ps. stellariicola isolates

obtained were nearly identical, with only one base pair difference between

KACC42827 and the other isolates. They formed a separate clade distinct from

the other species of Mycosphaerellaceae (1.00 PP value) in the Bayesian tree

(Fic. 2). Our phylogenetic analysis indicated that Mycosphaerella cerastiicola

Crous is sister to the Ps. stellariicola clade (0.99 PP value); Ramularia spp.

are also phylogenetically close to the new species. In recent studies, based on

18S, 28S, and ITS-5.8S rDNA sequence analyses (Crous et al. 2009 & 2011),

isolate CPC11297 (representing Ps. stellariicola) was tentatively identified as

a Ramularia sp. based on its placement in the Ramularia clade. Nevertheless,

based on the morphological characteristics described in this study, the isolate

is better accommodated in Pseudocercosporella. In addition, the fact that

Ps. stellariicola is closely related with Ramularia spp. is not surprising, as

Pseudocercosporella is polyphyletic within Mycosphaerellaceae and Capnodiales,

as shown in previous phylogenetic studies (Crous et al. 2009, Frank et al. 2010)

and the present analysis. In conclusion, both morphological and molecular

data support Ps. stellariicola as a novel species.

Acknowledgements

The authors are grateful to Uwe Braun and Pedro W. Crous for critical comments on

the manuscript and for acting as pre-submission reviewers. This work was supported

by a grant from Regional Subgenebank Support Program of Rural Development

Administration, Republic of Korea.

Literature cited

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65-75.

Braun U. 1995. A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic

hyphomycetes). Vol. 1. IHW-Verlag, Miinchen.

Crous PW, Schoch CL, Hyde KD, Wood AR, Gueidan C, de Hoog GS, Groenewald JZ. 2009.

Phylogenetic lineages in the Capnodiales. Stud. Mycol. 64: 17—47.

http://dx.doi.org/10.3114/sim.2009. 64.02

Crous PW, Tanaka K, Summerell BA, Groenewald JZ. 2011. Additions to the Mycosphaerella

complex. IMA Fungus 2: 49—64. http://dx.doi.org/10.5598/imafungus.2011.02.01.08

Frank J, Crous PW, Groenewald JZ, Oertel B, Hyde KD, Phengsintham P, Schroers HJ. 2010.

Microcyclospora and Microcyclosporella: novel genera accommodating epiphytic fungi causing

sooty blotch on apple. Persoonia 24: 93-105. http://dx.doi.org/10.3767/003158510X510560

Huelsenbeck JP, Ronquist F. 2001. MRBAYES: Bayesian inference of phylogenetic trees.

Bioinformatics 17: 754—755. http://dx.doi.org/ 10.1093/bioinformatics/17.8.754

Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H, Valentin F,

Wallace IM, Wilm A, Lopez R, Thompson JD, Gibson TJ, Higgins DG. 2007. Clustal W and

Clustal X version 2.0. Bioinformatics 23: 2947-2948.

http://dx.doi.org/ 10.1093/bioinformatics/btm404

Pseudocercosporella stellariicola sp. nov. (Korea) ... 275

Lee SB, Taylor JW. 1990. Isolation of DNA from fungal mycelia and single spores. 282-287, in: MA

Innis et al. (eds): PCR protocols: a guide to methods and applications. Academic Press, San

Diego, CA, USA.

Page RDM. 1996. TREEVIEW: An application to display phylogenetic trees on personal computers.

Comput. Appl. Biosci. 12: 357-358.

Posada D. 2008. jMODELTEST: Phylogenetic Model Averaging. Mol. Biol. Evol. 25: 1253-1256.

Shin HD, Kim JD. 2001. Cercospora and Allied Genera from Korea. National Institute of Agricultural

Science and Technology, Suwon, Korea. 302 p.

White TJ, Bruns T, Lee SB, Taylor JW. 1990. Amplification and direct sequencing of fungal

ribosomal RNA genes for phylogenetics. 315-322, in: MA Innis et al. (eds): PCR protocols: a

guide to methods and applications. Academic Press, San Diego, CA, USA.

MYCOTAXON

http://dx.doi.org/10.5248/119.277

Volume 119, pp. 277-289 January-March 2012

A key to the peltigericolous fungi in Turkey

MEHMET GOKHAN HALIcI’**, MEHMET CANDAN?’ & AYSEN TURK’

" Erciyes Universitesi, Fen Fakiiltesi, Biyoloji Boliimti, 38039 Kayseri, Turkey

?Anadolu Universitesi, Fen Fakiiltesi, Biyoloji Boliimti, Eskisehir, Turkey

CORRESPONDENCE TO *: '" mghalici@erciyes.edu.tr,

*mecandan@anadolu.edu.tr, aturk@anadolu.edu.tr

ABSTRACT — Eleven taxa of peltigericolous fungi are reported from Turkey for the first

time - Arthonia fuscopurpurea, A. peltigerina, Corticifraga fuckelii, C. peltigerae, Nectriopsis

lecanodes, Norrlinia peltigericola, Polycoccum peltigerae, Scutula epiblastematica, Stigmidium

leucophlebiae, Thelocarpon epibolum var. epithallinum, and Zwackhiomyces kiszkianus. A key

is provided to the 19 specific and infraspecific peltigericolous fungal taxa reported from

Turkey. Also provided are additional references, a list of the distributions in Turkey, and a

host index.

Key worps — Ascomycota, lichenicolous fungi, Peltigera

Introduction

During our 2008-10 project, “Taxonomical research on Peltigera species

distributed in western and eastern Black Sea region of Turkey and their

lichenicolous fungi,’ we collected hundreds of Peltigera specimens and cited

the distribution of Peltigera species in about two thirds of Turkey in our final

project report (unpublished). The lichenized fungus genus Peltigera is a well-

known exceptionally rich host for the obligately lichenicolous fungi known as

“peltigericolous fungi” (Miadlikowska & Alstrup 1995, Candan et al. 2010).

More than 80 species of peltigericolous fungi have been reported worldwide

(Hawksworth 1980, Hawksworth & Miadlikowska 1997). Hawksworth (1980)

cited the fact that Peltigera is one of the richest genera to harbor lichenicolous

fungi as evidence that it is derived from a particularly ancient ascomycete

group.

Only eight peltigericolous species (Halici 2008; Candan et al. 2010) have

been included among the 166 lichenicolous fungal species reported from

Turkey (Candan & Halici 2011, Halici & Candan 2011, Kocakaya et al. 2011).

Our addition here of 11 peltigericolous taxa raises that total to 19 peltigericolous

and 177 lichenicolous fungi now known from the country. We provide a key

278 ... Halici, Candan & Tiirk

to the known peltigericolous fungi with their distribution and host species in

Turkey to encourage further study of the group in Turkey.

Material & methods

All materials (except Refractohilum peltigerae) of the 19 cited taxa were studied

personally and are deposited in ANES (Eskisehir, Turkey). Specimens were examined

with a Leica DM 1000 research microscope. Hand sections were examined in K (10%

KOH solution), I (Merck Lugol's iodine), and water. Ascospore measurements were

made in water. Under “Taxonomy’ we provide as much information as possible from the

Turkish specimens studied for taxa new to Turkey and refer to more detailed descriptions

for all taxa. A map showing the geographical provinces of Turkey (Fic. 1) and provincial

vehicle registration codes (TABLE 1) summarize the distribution of the taxa in Turkey. A

host index (TaBLE 2) is also provided.

Taxonomy

Arthonia fuscopurpurea (Tul.) R. Sant.

A detailed description is provided by Santesson (1960).

The Turkish specimen, collected on a thallus of Peltigera polydactylon,

seems weakly parasitic, causing bleaching of infected parts of the host thallus.

Ascomata arthonioid, sometimes grouped together. Hymenium hyaline, K/I+

blue, I+ red. Epihymenium greenish brown. Asci 8-spored, ascospores hyaline,

1-septate, 9-11 x 3-4 um. Distribution in Turkey: 29.

MATERIAL EXAMINED: TURKEY. GUMUsHANE, Torul, southern slopes of Zigana

Mountain, 40°38'28"N 39°23'50"E, alt. 2100 m, on Peltigera polydactylon, 04.09.2009,

M. Candan, M.G. Halici & O. Sezer (ANES 14199).

Arthonia peltigerina (Almq.) H. Olivier

A detailed description is provided by Almquist (1880).

Turkish specimens, collected on thalli of Peltigera canina and P. praetextata,

seem parasitic, bleaching the infected parts of the host thalli. Infection is

generally seen in the old parts of the host thalli. Ascomata arthonioid, < 0.5

mm diam., often grouped together in subconcentric rings, sometimes grouped

together. Hymenium hyaline, K/I+ blue, I+ red, dextrinoid. Epihymenium

brown, slightly greenish tones available, I+ reddish brown. Asci 8-spored, I+

yellowish orange. Ascospores hyaline, 1-septate, cells with oil droplets, with a

gelatinous sheath, 14-17 x 5-7 um. Distribution in Turkey: 03.

MATERIAL EXAMINED: TURKEY. AFYON, Sultandagi, Sultandag1 Mountains, 38°28'35"N

31°10'48"E, alt. 1622 m, in Quercus vulcanica forest, on Peltigera canina, 11.05.2010,

M. Candan (ANES 13824); 38°28'48"N 31°11'12”E, alt. 1692 m, in Quercus vulcanica

forest, on Peltigera praetextata on mosses, 11.05.2010, M. Candan (ANES 13836).

Capronia peltigerae (Fuckel) D.Hawksw.

Detailed descriptions of the species are provided by Hawksworth (1980) and

Zhurbenko (2004) and of the Turkish specimens by Candan & al. (2010).

Key to peltigericolous fungi in Turkey ... 279

Specimens were collected on thalli of Peltigera canina, P. leucophlebia,

P. monticola and P. neckeri. Distribution in Turkey: 03, 09, 26, 29, 35, 37, 48.

MATERIAL EXAMINED: TURKEY. Ayp1n, Nazilli, north of Kavaklidere, 37°27'39"N

28°20'39"E, alt. 800 m, in Pinus brutia forest, on Peltigera neckeri, 15.06.2009, M.

Candan & O. Sezer (ANES 14220); Esk1sEHIR, In6nii, south of Erenkéy Village, 39°44'N

30°01’E, alt. 1100 m, in Quercus sp. communities and Pinus nigra forest, on Peltigera

monticola, 02.05.2010, M. Candan (ANES 14216); GUMUSHANE, Torul, exit of Torul,

southern slopes of Zigana pass, 40°38'04"N 39°23'12"E, alt. 1019 m, in Picea and Abies

forest, on Peltigera canina, 04.09.2009, M. Candan & M.G. Halici (ANES 14217); Izmtr,

Kemalpasa, Nif Mountain, 38°24'04"N 27°23'52"E, alt. 869 m, in Pinus nigra forest,

on Peltigera neckeri, 16.06.2009, M. Candan & O. Sezer (ANES 14218); KasTAMONU,

Senpazar, southwest of Sada Village, Kiire Mountains, 41°43'45"N 33°29'04"E, alt. 830

m, in Abies nordmanniana and Pinus sylvestris mixed forest, on Peltigera leucophlebia,

30.07.2010, M. Candan & M.G. Halici (ANES 14219).

Corticifraga fuckelii (Rehm) D. Hawksw. & R. Sant.

A detailed description is provided by Hawksworth & Santesson (1990).

The Turkish specimen, collected on thalli of Peltigera sp. (with gyrophoric

acid determined by TLC), seems parasymbiotic (no visible damage to the

host thallus). Ascomata apothecia, 0.2-0.3 mm diam., not arising in circular

spots of the host thallus. Interascal filaments with a hyaline end cell, 1.5-2 um

diam. Hymenium 50-60 um tall. Asci 8-spored, I+ red. Ascospores hyaline,

1-2-septate, 15-18 x 5-7 um. Distribution in Turkey: 08.

MATERIAL EXAMINED: TURKEY. ARTVIN, Savsat, northeast of Savsat, 41°13'67"N

42°22'58"E, alt. 1360 m, in Picea orientalis forest, on Peltigera sp. (with gyrophoric acid)

on mosses, 31.08.2009, M. Candan, M.G. Halici & Okan Sezer (ANES 13825).

Corticifraga peltigerae (Fuckel) D. Hawksw. & R. Sant.

A detailed description is provided by Hawksworth & Santesson (1990).

The Turkish specimens, collected on thalli of Peltigera canina, P. monticola

and P. rufescens, seem parasitic (bleaching seen in the infected parts of the host

thalli). Ascomata apothecia, 0.2-0.3 mm diam., arising in circular spots of the

host thallus. Interascal filaments with a brown end cell (not a good taxonomical

character, in our opinion, as this is indistinct in sections from young ascomata).

Hymenium 50-60 um tall. Asci 8-spored. Ascospores hyaline, 1-3-septate,

12-20 x 5-7 um. Distribution in Turkey: 08, 42, 45, 66.

MATERIAL EXAMINED: TURKEY. ARTVIN, Savsat, northeast of Savsat, 41°13'67"N

42°22'58"E, alt. 1360 m, in Picea orientalis forest, on Peltigera rufescens, 31.08.2009,

M. Candan, M.G. Halici & O. Sezer (ANES 13830); Ardanuc, 41°13'52"N 42°05'31"E,

alt. 472 m, in Alnus-Quercus forest, on Peltigera canina, 31.08.2009; M. Candan, M.G.

Halici & O. Sezer (ANES14221); Manisa, Akhisar, Sindirgi road, east of Isaca Village,

39°09'35"N 28°01'29"E, alt. 490 m, in Pinus brutia forest, on Peltigera monticola ,

17.06.2009; M. Candan & O. Sezer (ANES 13829); Konya, Bozkir-Bel6ren, west of

Aydinkisla Village, 37°13'21"N 32°21'27E, alt. 1400 m, in Juniperus sp. communities,

on Peltigera monticola, 05.08.2009, M. Candan & M.G. Halici (ANES 13826); Yozeat,

10 km to Yozgat, 39°37'13"N 34°56'53"E, alt. 1015 m, on Peltigera rufescens, 25.08.2009.

M. Candan, M.G. Halici & O. Sezer (ANES 13828).

280 ... Halici, Candan & Tiirk

Dacampia rufescentis (Vouaux) D. Hawksw.

Detailed descriptions of the species are provided by Vouaux (1912) and Hawksworth

(1986) and of the Turkish specimen by Candan & al. (2010).

P. praetextata, and P. rufescens. Distribution in Turkey: 26, 32, 37, 66.

MATERIAL EXAMINED: TURKEY. Isparta, Egirdir, east of Mahmatlar Village, northern

slopes of Egirdir Lake, 37°57'11"N 30°56'41"E, alt. 940 m, in siliceous rocks, on Peltigera

kristinssonii, 31.07.2009, M. Candan & M.G. Halici (ANES 14213). KasTAMONU,

Seydiler, southeast of Seydiler, Kastamonu highway, Oyrak passage, 41°34'26"N

33°46'43"E, alt. 1200 m, in Salix sp., Populus sp., and open areas, on Peltigera rufescens,

30.07.2010, M. Candan & M.G. Halici (ANES 14214); Yozeat, Cayiralan, Cat forests,

39°19'30"N 35°50'17"E, alt. 1650 m, in Pinus sylvestris forest, on Peltigera ponojensis,

27.08.2009, M. Candan, M.G. Halici & O. Sezer (ANES 14215).

Nectriopsis lecanodes (Ces.) Diederich & Schroers

Detailed descriptions are provided by Booth (1959) and Sérusiaux et al. (1999).

Specimens were collected on thalli of Peltigera kristinssonii, PR. ponojensis,

Turkish specimens, collected on thalli of Peltigera leucophlebia, seem

parasitic (extensive damage seen in the infected parts of the host thalli).

Ascomata perithecioid, 150-300 um diam., pinkish to orange, often covered

with a whitish tomentum, arising on a small stalk. Hymenium orange. Asci

8-spored. Ascospores oblique, uniseriately arranged in the asci, hyaline,

1-septate, one or two oil droplets present in each cell, ellipsoid, round in the

apices, more or less verruculose, 10-12 x 4-5 um. Distribution in Turkey:

of.

MATERIAL EXAMINED: TURKEY. ERzuRuM, Olur, southeast of Yukarikaracasu Village,

40°49'26"N 42°17'11"E, alt. 1800 m, in Pinus sylvestris forest, on Peltigera leucophlebia

on mosses, 30.08.2009, M. Candan, M.G. Halici & O. Sezer (ANES 14203); KASTAMONU,

Azdavay, southwest of Sada Village, Kiire Mountains, 41°43'45"N 33°29'04"E, alt. 830 m,

in Abies nordmanniana and Pinus sylvestris forest, on Peltigera leucophlebia on mosses,

30.07.2010, M. Candan & M.G. Halici (ANES 13837).

Neolamya peltigerae (Mont.) Theiss. & Syd.

Detailed descriptions of the species are provided by Keissler (1930) and Ertz et al.

(2004) and of the Turkish specimen by Candan & al. (2010).

Turkey: 08, 55.

MATERIAL EXAMINED: TURKEY. SAMSuN, Ayvacik, northwest of Hasan Ugurlu Dam

Lake, 40°56'33"N 36°38'21"E, alt. 80 m, in siliceous rocks, on Peltigera didactyla,

07.09.2009, M. Candan, M.G. Halici & O. Sezer (ANES 14202).

Norrlinia peltigericola (Nyl.) Theiss. & Syd.

Detailed descriptions are provided by Hawksworth (1980) and Martinez & Hafellner

(1998).

Specimens were collected on thalli of Peltigera didactyla. Distribution in

The Turkish specimen, collected on a Peltigera praetextata thallus, seems

parasymbiotic (no visible damage to the host lichen). Ascomata perithecioid,

Key to peltigericolous fungi in Turkey... 281

black, aggregated in small groups, immersed only at the base, wall blackish in

the outer part, brown in the inner part. Interascal filaments not present, oil

droplets present. Asci 2-spored. Ascospores hyaline, rarely very pale brownish,

muriform with 12 transsepta and 4 longisepta, 42-52 x 14-28 um. Distribution

in Turkey: 09.

MATERIAL EXAMINED: TURKEY. AypINn, Bozdagan, Madran Mountain, south of

Altintas Village, 37°35'40"N 28°21'00"E, alt. 800 m, in Pinus brutia forest, on Peltigera

praetextata, 15.06.2009, M. Candan & O. Sezer (ANES 13833).

Phoma peltigerae (P. Karst.) D. Hawksw.

Detailed descriptions of the species are provided by Keissler (1930) and Hawksworth

(1981) and of the Turkish specimens by Candan & al. (2010).

Specimens were collected on thalli of Peltigera canina, P. elisabethae, and

P. rufescens. Distribution in Turkey: 11, 16, 43.

Polycoccum crassum Vézda

A detailed description of the species is provided by Hawksworth & Diederich (1988)

and of the Turkish specimens by Halici et al. (2007) and Candan & al. (2010).

Specimens were collected on thalli of Peltigera canina, P. elisabethae,

P. monticola, and P. neckeri. Distribution in Turkey: 08, 29, 35, 44, 48, 66.

MATERIAL EXAMINED: TURKEY. ARTVIN, Savsat, northeast of Savsat, 41°13'67"N

42°22'58"E, alt. 1360 m, in Picea orientalis forest, on Peltigera elisabethae, 31.08.2009,

M. Candan, M.G. Halici & O. Sezer (ANES 14206); GUMUSHANE, Torul, exit of Torul,

southern slopes of Zigana pass, 40°38'04"N 39°23'12"E, alt. 1019 m, in Picea and Abies

forest, on Peltigera neckeri, 04.09.2009, M. Candan, M.G. Halici & O. Sezer (ANES

14207); Izmir, Kemalpasa, Nif Mountain, 38°24'50"N 27°23'40"E, alt. 710 m, in Pinus

brutia forest, on Peltigera neckeri, 16.06.2009, M. Candan & O. Sezer (ANES 14208);

YOZGAT, Cayiralan, Cat forest enterance, Sivas province border, 39°19'30"N 35°50'17"E,

alt. 1650 m, in Pinus sylvestris forest, on Peltigera monticola, 27.08.2009, M. Candan, M.

G. Halici & O. Sezer (ANES 14209).

Polycoccum peltigerae (Fuckel) Vézda

A detailed description is provided by Hawksworth & Diederich (1988).

Turkish specimens, collected on thalli of Peltigera leucophlebia and P.

rufescens, seem slightly parasitic (swellings seen on the host lichens; heavy

infections may kill parts of the host thallus). Ascomata perithecioid, ~ 200

um diam., entirely immersed in the host lichens, becoming erumpent only in

the ostiolar region. Asci 8-spored. Ascospores brown, 1-septate, ellipsoid to

fusiform, cells generally attenuated at the ends, 14-18 x 5-6 um. Distribution

in Turkey: 08, 37.

MATERIAL EXAMINED: TURKEY. ARTVIN, Savsat, northeast of Savsat, 41°13'67"N

42°22'58"E, alt. 1360 m, in Picea orientalis forest, on Peltigera rufescens, 31.08.2009, M.

Candan, M. G. Halici & O. Sezer (ANES 14204); KASTAMONU, near Azdavay, southwest

of Sada Village, Kiire Mountains, alt. 830 m, in Abies nordmanniana and Pinus sylvestris

forest, on Peltigera leucophlebia , 30.07.2010. M. Candan & M.G. Halici (ANES 14205).

282 ... Halici, Candan & Tiirk

Pronectria robergei (Mont. & Desm.) Lowen

A detailed description of the species is provided by Keissler (1930) and of the

Turkish specimen by Candan & al. (2010).

The specimen was collected on thallus of Peltigera rufescens. Distribution in

Turkey: 08.

Refractohilum peltigerae (Keissl.) D. Hawksw.

Detailed descriptions of the species are provided by Hawksworth (1977, 1979) and of the

Turkish specimen by John & Breuss (2004) and Halic1 (2008).

The specimen was collected on thallus of Peltigera polydactylon. Distribution

in Turkey: 61.

Scutula epiblastematica (Wallr.) Rehm

Detailed descriptions are provided by Triebel et al. (1997) and Clauzade et al.

(1989).

The Turkish specimen, collected on thallus of Peltigera praetextata, seems

parasitic (host algal photobiont depleted in the infected parts of the thallus).

Ascomata apothecioid, often aggregated, cream coloured when young,

becoming greenish black when mature. Inner part of exciple purplish brown

Hymenium hyaline, I+ red, hypothecium pale brownish. Asci Scutula-type

(with a fuzzy K/I+ blue tube-like structure in the tholus). Ascospores hyaline,

1-septate, with oil droplets, 8-11 x 3-4 um. Distribution in Turkey: 67.

MATERIAL EXAMINED: TURKEY. ZonGuLDAK, Devrek, north of I[sabeyli Village,

Babadag Mountain, 41°13'23"N 31°52'57"E, alt. 600 m, in Carpinus sp. and Fagus

orientalis forest, on Peltigera praetextata on mosses, 29.07.2010, M. Candan & M.G.

Halici (ANES 14198).

Stigmidium leucophlebiae Cl. Roux & Triebel

A detailed description is provided by Roux & Triebel (1994).

Turkish specimens, collected on thalli of Peltigera leucophlebia, seem parasitic

(infected host thallus parts brownish or blackish). Ascomata perithecioid,

50-60 um diam., semi-immersed. Vegetative hyphae sometimes visible on the

host thalli. Asci 8-spored. Ascospores hyaline, 1-septate, cells not equal in size,

with a gelatinous sheath, 9-14 x 4-5 um. Distribution in Turkey: 37, 78.

MATERIAL EXAMINED: TURKEY. KaraBtx, Yenice, Seker canyon, 41°11'26"N

32°21'35"E, alt. 230 m, in mixed Pinus nigra and broad leaved tree forest, on Peltigera

leucophlebia, 01.06.2009, A. Tiirk & M. Candan (ANES 13834); KasTAMONU, Senpazar,

southwest of Sada Village, Kiire Mountains, 41°43'45"N 33°29'04"E, alt. 830 m, in Abies

nordmanniana and Pinus sylvestris mixed forest, on Peltigera leucophlebia, 30.07.2010,

M. Candan & M. G. Halici (ANES 13835).

Stigmidium peltideae (Vain.) R.Sant.

A detailed description of the species is provided by Roux & Triebel (1994) and of the

Turkish specimens by Candan & al. (2010).

Key to peltigericolous fungi in Turkey ... 283

Specimens were collected on thalli of Peltigera collina, P. neckeri, and

P. praetextata. Distribution in Turkey: 03, 09, 26, 29, 48, 66.

MATERIAL EXAMINED: TURKEY. Aypin, Nazilli, Semailli Village road, 38°01'12"N

28°18'24"E, alt. 850 m, in Quercus forest and siliceous rocks, on Peltigera neckeri,

15.06.2009, M. Candan & O. Sezer (ANES 14210); Eskisenrr, Tiirkmen Dagi, Golciik

road, 39°29'N 30°21'E, alt. 1320 m, in siliceous rocks, on Peltigera collina, 24.03.2009,

M. Candan (ANES 14211); GUMUsHANE, Torul, exit of Torul, southern slopes of Zigana

pass, 40°38'04"N 39°23'12"E, alt. 1019 m, in Picea and Abies forest, on Peltigera neckeri,

04.09.2009, M. Candan, M.G. Halici & O. Sezer (ANES 14212); MuGxa, Yatagan,

Yatagan old road, Gokbel pass, 37°25'34"N 28°08'52"E, alt. 323 m, on Peltigera neckeri,

15.06.2009, M. Candan & O. Sezer (ANES 14222); Yozeat, Akdagmadeni, east of

Akdagmadeni, 39°40'00"N 35°56'07’E, alt. 1603 m, in Pinus sylvestris forest, on Peltigera

neckeri, 26.08.2009, M. Candan, M.G. Halici & O. Sezer (ANES 14223).

Thelocarpon epibolum var. epithallinum (Leight. ex Nyl.) G. Salisb.

Detailed descriptions are provided by Salisbury (1966) and Ahti (1973).

The Turkish specimen, collected on a Peltigera leucophlebia thallus,

seems parasymbiotic (no visible damage to the host lichen seen). Ascomata

perithecioid, superficial, bright yellow, citrine green when fresh, pruinose,

150-250 um diam. Interascal filaments present, hyaline, oil droplets present.

Asci more than 50-spored. Ascospores hyaline, simple, 9-11 x 3-4 um.

Distribution in Turkey: 78.

MATERIAL EXAMINED: TURKEY. KaraBuxK, Seker, Yenice, Seker canyon, 41°11'26"N

32°21'35"E, alt. 230 m, in mixed Pinus nigra and broad leaved tree forest, on Peltigera

leucophlebia, 01.06.2009, A. Tiirk & M. Candan (ANES 13831).

Zwackhiomyces kiszkianus D. Hawksw. & Miadl.

A detailed description is provided by Hawksworth & Miadlikowska (1997).

The Turkish specimen, collected on a Peltigera elisabethae thallus, seems

parasymbiotic (no visible damage to the host lichen seen). Ascomata perithecioid,

scattered, immersed, exposed only at the ostiole. Asci (6—)8-spored. Ascospores

hyaline, ellipsoid to somewhat soleiform, 1-septate, constricted at the septum,

25-27 x 11-12 um. Distribution in Turkey: 08.

MATERIAL EXAMINED: TURKEY. ARTVIN, Savsat, northeast of Savsat, 41°13'67"N

42°22'58"E, alt. 1360 m, in Picea orientalis forest, on Peltigera elisabethae, 31.08.2009, M.

Candan, M.G. Halici & O. Sezer (ANES 14201).

Key to the peltigericolous fungi species in Turkey

TA) Spates produce dats Cie ta. sca tr seteg deh seteg teh fied bh Med hh Mie hh feed dee beets 2

Le “BSpOres NOb Produced 18icayot a cose a cesste a cea uiehee vaseiie creeds saaieen uMietd 9 Bleed 18

Dl Te FASCOMMAta PCLIENCCLOIG 9-2) aja Bea wind eect Pa wlad 2 Rated Ola Lda, 8 Ilan conte dled 3

2 | siscomata arthonioidiorapothecioid 22 7 1o.00.t 189th ne nt ttl Cath 14

32): PAscospores brown when atime, .. a4. ee. ed oateie hh ated see een ae aaah 4

3. Ascomata hyaline or rarely very pale brown when mature ................... 7

284 ... Halici, Candan & Tiirk

A(3). Ascespores ] septate 2o:6 2.86 92-45-00 hens hGe shes Mane ache 8 5

4. Ascospores 3-septate or muriform with trans- and longisepta ................ 6

5(3). Perithecia 300-500 um diam., in general on the lower surface of the host lichen.

Asci 4-spored. Ascospores 24-30 x 8-10 um.............. Polycoccum crassum

5. Perithecia ~ 200 um diam., entirely immersed in the host lichen, becoming

erumpent only in the ostiolar region. Asci 8-spored. Ascospores 14-18 x 5-

Sg LEE Avi bt be Bache Bede Sit LOL bene tN Nae De Polycoccum peltigerae

6(4). Perithecia covered with stiff, smooth-walled, and dark brown hairs

Bp Pegmah » hast sitlaae sient eats ea ge ce ths Sl Se oe Capronia peltigerae

6: » “Pétithectaswithout Watrscy nF.) Fo eee eh ee Dacampia rufescentis

WS). Ascrcontainingm ote: thats SpOrese tent. wats Wade as Meo std ee vice Wad 6 ahd Ri nal ie acl Dy 8

FS EASEH UP LORSRGOTER toons cast ish donaensuten dae tatew tise isn a uteb lates nal ates nabtates wae cee aay ous nak Oy 9

8(5). Perithecia immersed to slightly erumpent, black. Asci 16-spored

Steerer cere Pear A: Sarit Pe gat 2 etre ret s faerie: tones eres Neolamya peltigerae

8. Perithecia superficial, bright yellow, citrine green when fresh, pruinose.

Asci more thatt 50+spored «Ne. laieg./ ens Thelocarpon epibolum var. epithallinum

OU) IP ASCE BSP OEM sacs ah dod s want x eGo oh lope FRG F Map ¥ NG ON b aaa Lagan nears 4 10

Om “ASCE D-spotedinn ©. cares 1. wares 0. aes i. cae 20 Rar, 0, oe oh Norrlinia peltigericola

LOCO Peritheciati piksjorange- oF reddish Conesicnes: aya. Ras wlth asta Lonoke F 11

LO.» “Peritheeiatblaches, «44th, +2 dike +2 dined dth tera di asee dt see-g ih pete’ Insnew A Rede pets 12

11(10). Perithecia pinkish to orange, often covered with a whitish tomentum,

atising-on-a sinvall. shally, 8 p59 gents Page nse Sngale st Peet ht aps use Nectriopsis lecanodes

11. Perithecia orange-red, becoming yellowish when old; immersed to

SOMAP IM BICLS EM, OF ose nie Rage con Pegl e ge eBay MPa hes Pronectria robergei

L2G). MiniterascalilamenfsabSent 3, . tote oo fads tole sk ole sbatele Wolaslold wali 13

12. einterascal-filamients presents 6.21. semctlose eer testes Zwackhiomyces kiszkianus

13(12). Perithecia 50-60 um diam. Ascospores 9-14 x 4-5 um

Eee ic CSatrin SenehAe Een A> Maire 4h SPs Wik Ste be Stigmidium leucophlebiae

13. Perithecia 35-40 um diam. Ascospores 11-12 x 2.5-3 um

Es Ene ALPES LIER AD BEN A Oe SENS lh Then OER ie Ue A hie Stigmidium peltideae

FAD), Ascomataapothetioid -. yo. oyu eeioe y waieye # ea ee F Rapley # Map lege BF Ma le hg ee Fale Be 15

14 Asconiat a arthomioid.s necs! aces! wenn (eka ee ae 10m Coe, Re Leos oe Ose re Eee 17

WGA ay AS CIMDE- Scr isla sy Per Re ks Re whe a oll Bl a Red eels al showed Redectid Piste 15

Se SASCESCUTMA CTY R ta ee. ae a cetiem a wets nusas toh peonabit sea bis Scutula epiblastematica

16(15). Ascomata not arising in circular spots of the host thallus, not bleaching

the thallus. Interascal filaments with a hyaline end cell, 1.5-2 um diam

bn geht + ES ites gg atiaeee-g Lith 2-2 Saber g dinbee Fd tnaer 4 Azer g S'Réet Corticifraga fuckelii

16. Ascomata arising in circular bleached spots of the host thallus.

Interascal filaments with a brown end cell ............. Corticifraga peltigerae

Key to peltigericolous fungi in Turkey ... 285

17(14). Ascomata often grouped together in subconcentric rings.

AScOspores.14=17. X5=F (Id my daca tam, ties sim wes tery atvateryaatlons tac Arthonia peltigerina

17. Ascomata sometimes grouped together. Ascospores 9-11 x 3-4 um

FO Se Mee hie ly eis, 9 Plate, 9 atid ae Plbeche St Arthonia fuscopurpurea

18(1).Conidia produced within pycnidia (coelomycetes). .......... Phoma peltigerae

18. Conidia not produced within pycnidia (hyphomycetes)

PE EE Le Oe, Se Bee Be Ae Bee Refractohilum peltigerae

Acknowledgements

Kerry Knudsen (UCR) and Wolfgang von Brackel (Germany) are thanked for

reviewing this paper. Shaun Pennycook is also thanked for his very useful comments

on formatting the manuscript. This study was supported by TUBITAK (108T556 coded

project).

Literature cited

Ahti T. 1973. Notes on the lichens of Newfoundland. 2. Thelocarpon epibolum Nyl. Ann. Bot. Fenn.

10: 66-67.

Almquist S. 1880. Monographia Arthoniarum Scandinaviae. Kongl Svenska Vetensk. Akad. Hand.

17: 1=69:

Booth C. 1959. Studies of Pyrenomycetes IV. Nectria (part 1.). Mycol. Pap. 73: 1-115.

Candan M, Halici MG. 2011. New Cercidospora records for Turkey. Turkish Journal of Botany 35:

625-629.

Candan M, Halici MG, Ozdemir Tiirk A. 2010. New records of peltigericolous fungi from Turkey.

Mycotaxon 111: 149-153. http://dx.doi.org/10.5248/111.149

Clauzade G, Diederich P, Roux C. 1989. Nelikenigintaj fungoj likenlogaj. Bull. Soc. Linn. Provence,

numéro spécial 1: 1-142.

Ertz D. 2004. Neolamya. pp. 677-678, in: TH Nash III et al. (eds). Lichen flora of the Greater

Sonoran Desert Region.Vol. 2. Tempe: Arizona State University.

Halici MG. 2008. A key to the lichenicolous Ascomycota (including mitosporic fungi) of Turkey.

Mycotaxon 104: 253-286.

Halici MG, Candan M. 2011. Arthonia anatolica sp. nov. (Arthoniaceae) on Aspicilia contorta subsp.

hoffmanniana, a new lichenicolous species from Turkey. Mycotaxon 116: 335-339.

Halici MG, Ozdemir Tiirk A, Candan M. 2007. New records of pyrenocarpous lichenicolous fungi

from Turkey. Mycotaxon 99: 201-206.

Hawksworth DL. 1977. Three new genera of lichenicolous fungi. Bot. J. Linn. Soc. 75: 195-209.

http://dx.doi.org/10.1111/j.1095-8339.1977.tb01484.x

Hawksworth DL. 1979. The lichenicolous hyphomycetes. Bull. Brit. Mus. (Nat. Hist.), Bot. 6:

183-300.

Hawksworth DL. 1980. Notes on some fungi occurring on Peltigera, with a key to accepted species.

Trans. Br. Mycol. Soc. 74: 363-386. http://dx.doi.org/10.1016/S0007-1536(80)80167-7

Hawksworth DL. 1981. The lichenicolous coelomycetes. Bull. Brit. Mus. (Nat. Hist.), Bot. 9: 1-98.

Hawksworth DL. 1986. Notes on British lichenicolous fungi: V. Notes Roy. Bot. Gard. Edinburgh

43: 497-519.

286 ... Halici, Candan & Tiirk

Hawksworth DL, Diederich P. 1988. A synopsis of the genus Polycoccum (Dothideales), with a key

to accepted species. Trans. Brit. Mycol. Soc. 90: 293-312.

http://dx.doi.org/10.1016/S0007-1536(88)80101-3

Hawksworth DL, Miadlikowska J. 1997. New species of lichenicolous fungi occurring on Peltigera

in Ecuador and Europe. Mycol. Res. 101: 1127-1134.

http://dx.doi.org/10.1017/S0953756297003778

Hawksworth DL, Santesson R. 1990. A revision of the lichenicolous fungi previously referred to

Phragmonaevia. Biblioth. Lichenol. 38: 121-143.

John V, Breuss O. 2004. Flechten der éstlichen Schwarzmeer-Region in der Tiirkei (BLAM-

Exkursion 1997). Herzogia 17: 137-156.

Keissler K. 1930. Die Flechtenparasiten. Rabenhorst’s Kryptogamen-Flora von Deutschland,

Oesterreich und der Schweiz 8: i- xi, 1-712.

Kocakaya M, Halict MG, Aksoy A. 2011. Zwackhiomyces turcicus sp. nov. (Ascomycota,

Xanthopyreniaceae) from Turkey. Mycotaxon 116: 329-333. http://dx.doi.org/10.5248/116.329

Martinez I, Hafellner J. 1998. Lichens and lichenicolous fungi on Peltigerales in the Iberian

Peninsula and the Canary Islands. Mycotaxon 69: 271-310.

Miadlikowska, J, Alstrup V. 1995. Some peltigericolous fungi and lichens mainly from Poland.

Graphis Scripta 7: 7-10.

Roux C, Triebel D. 1994. Révision des espéces de Stigmidium et de Sphaerellothecium (champignons

lichénicoles on lichénisés, ascomycetes) corresponsant 4 Pharcidia epicymatia sensu Keissler ou

a Stigmidium schaereri auct. Bull. Soc. Linn. Provence 45: 451-542.

Salisbury G. 1966. A monograph of the lichen genus Thelocarpon. Lichenologist 3: 175-196.

http://dx.doi.org/10.1017/S0024282966000197

Santesson R. 1960. Lichenicolous fungi from northern Spain. Svensk bot. Tidskr. 54: 499-522.

Sérusiaux E, Diederich P, Brand AM, van den Boom P. 1999. New or interesting lichens and

lichenicolous fungi from Belgium and Luxembourg. VIII. Lejeunia 162: 1-95.

Triebel D, Wedin M, Rambold G. 1997. The genus Scutula (lichenicolous ascomycetes, Lecanorales):

species on the Peltigera canina and P. horizontalis groups. Symb. Bot. Upsal. 32(1): 323-337.

Vouaux A. 1912. Synopsis des Champignons parasites de Lichens. Bull. Trimestriel Soc. Mycol.

France 29: 33-128.

Zhurbenko MP. 2004. Lichenicolous and some interesting lichenized fungi from the Northern

Ural, Komi Republic of Russia. Herzogia 17: 77-86.

Key to peltigericolous fungi in Turkey ... 287

TABLE 1. Provincial vehicle registration codes.

CODE

PROVINCE

Adana

Adiyaman

Afyon

Agri

Amasya

Ankara

Antalya

Artvin

Aydin

Balikesir

Bilecik

Bingél

Bitlis

Bolu

Burdur

Bursa

Canakkale

Cankin

Corum

Denizli

Diyarbakir

Edirne

Elazig

Erzincan

Erzurum

Eskisehir

Gaziantep

Giresun

Giimiishane

Hakkari

Hatay

Isparta

Icel

Istanbul

Izmir

Kars

Kastamonu

Kayseri

Karklareli

Kirsehir

Kocaeli

CODE

PROVINCE

Konya

Kiitahya

Malatya

Manisa

Kahramanmaras

Mardin

Mugla

Mus

Nevsehir

Nigde

Ordu

Rize

Sakarya

Samsun

Siirt

Sinop

Sivas

Tekirdag

Tokat

Trabzon

Tunceli

Sanhurfa

Usak

Van

Yozgat

Zonguldak

Aksaray

Bayburt

Karaman

Karikkale

Batman

Surnak

Bartin

Ardahan

Igdir

Yalova

Karabiik

Kilis

Osmaniye

Diizce

288 ... Halici, Candan & Tiirk

TABLE 2. Host index.

HOST LICHEN LICHENICOLOUS FUNGUS

Peltigera canina Arthonia peltigerina

Capronia peltigerae

Corticifraga peltigerae

Phoma peltigerae

Polycoccum crassum

Peltigera collina Stigmidium peltideae

Peltigera didactyla Neolamya peltigerae

Peltigera elisabethae Phoma peltigerae

Polycoccum crassum

Zwackhiomyces kiszkianus

Peltigera kristinssonii Dacampia rufescentis

Peltigera leucophlebia Capronia peltigerae

Nectriopsis lecanodes

Polycoccum peltigerae

Stigmidium leucophlebiae

Thelocarpon epibolum var. epithallinum

Peltigera monticola Capronia peltigerae

Corticifraga peltigerae

Polycoccum crassum

Peltigera neckeri Capronia peltigerae

Polycoccum crassum

Stigmidium peltideae

Peltigera polydactylon Arthonia fuscopurpurea

Refractohilum peltigerae

Peltigera ponojensis Dacampia rufescentis

Peltigera praetextata Arthonia peltigerina

Dacampia rufescentis

Norrlinia peltigericola

Scutula epiblastematica

Stigmidium peltideae

Peltigera rufescens Corticifraga peltigerae

Dacampia rufescentis

Phoma peltigerae

Polycoccum peltigerae

Pronectria robergei

Peltigera sp. (with gyrophoric acid) Corticifraga fuckelii

MYCOTAXON

http://dx.doi.org/10.5248/119.291

Volume 119, pp. 291-299 January-March 2012

Taxonomy of the Gymnopus inusitatus group

and the new G. inusitatus var. cystidiatus from Hungary

VLADIMIR ANTONIN’, PETER FINY? & MICHAL TOMSOVSKY?

' Moravian Museum, Dept. of Botany, Zelny trh 6, CZ - 659 37 Brno, Czech Republic

*Zsombolyai u. 56, H - 8000 Székesfehérvar, Hungary

* Faculty of Forestry and Wood Technology, Mendel University in Brno,

Zemédeélskd 3, CZ - 613 00 Brno, Czech Republic

CORRESPONDENCE TO *: vantonin@mzm.cz

AsBstTRAcT — The authors describe a new bisporic basidiomycete, Gymnopus inusitatus

var. cystidiatus, collected in Hungary. It differs morphologically from var. inusitatus by the

presence of well-developed cheilocystidia and ecologically by growing in the sandy grassland

under Salix. Its taxonomic position is confirmed by DNA studies.

Key worps — Agaricomycetes, Agaricales, Omphalotaceae, phylogeny, ecology

Introduction

During his mycological field research in Hungary, the second author found

a fungus macroscopically similar to Mycetinis scorodonius (Fr.) A.W. Wilson

& Desjardin, but lacking a garlic-like smell. Moreover, microscopic studies

showed that it belongs to the genus Gymnopus (Pers.) Roussel, species group

of G. inusitatus/bisporus/catalonicus of sect. Levipedes (Fr.) Halling, subsect.

Alkalivirentes Antonin & Noordel. Species of sect. Levipedes are characterized

by a pileipellis composed of inflated, often lobed to coralloid, elements

(dryophila-type) and the taxa of subsect. Alkalivirentes by wall encrustations

that turn green in alkali (Antonin & Noordeloos 2010), a division supported by

molecular studies. DNA sequences of nuclear ITS and partial LSU ribosomal

RNA genes from holotypes of the above-mentioned taxa were identical.

Based on this result and morphological similarities, we propose synonymy of

G. inusitatus, G. bisporus, and G. catalonicus. We propose a new variety for the

Hungarian collection, which differs microscopically and ecologically from the

typical G. inusitatus.

292 ... Antonin, Finy & TomSovsky

Materials & methods

Macroscopic description of collected specimens is based on a photo and dry

specimens. Colour abbreviations follow Kornerup & Wanscher (1983); herbarium

abbreviations follow Holmgren & Holmgren (1998). Nomenclatural authorities are cited

according to the International Plant Names Index Authors website (http://www.ipni.org/

ipni/authorsearchpage.do). Microscopic features are described from dried material

mounted in H,O, c. 5% KOH, Melzer’s reagent, and Congo red using an Olympus BX-

50 light microscope with a magnification of 1000x. Measurements are designated for

basidiospores by E (quotient of length and width in any one spore) and Q (mean of

E-values) and for lamellae by L (number of entire lamellae) and | (number of series of

lamellulae between each pair of entire lamellae).

The nuclear ribosomal RNA ITS and LSU regions from the specimen were sequenced

as well as those from the types the type specimens of Gymnopus bisporus (BCN-SCM B-

4065), G. catalonicus (BCN-SCM B-4057), and G. inusitatus (BCN-SCM B-4058). Both

DNA extraction and PCR amplification were conducted twice, with new PCR reagents

used each time.

DNA was isolated from dried herbarium specimens using PowerSoil™ DNA Isolation

Kit (Mo-Bio). PCR amplifications were performed in the Mastercycler® ep thermocycler

(Eppendorf) according to TomSovsky & Jankovsky (2008). Macrogen Inc. (Seoul, Korea)

sequenced the amplified fragments.

Newly obtained sequences were edited manually using BioEdit version 7.0.9. (T. Hall,

Ibis Biosciences). Selected sequences from Gymnopus sect. Levipedes species. published

by Mata et al. (2006) were retrieved from GenBank and added to the dataset (Fic.

1). Gymnopus brassicolens and G. impudicus were selected as outgroup taxa. The ITS

alignment with gaps consisted of 790 characters including 630 constant and 160 variable

sites.

Bayesian analysis used MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003) and best-

fit model (GTR+G) and parameters given by MrModeltest (Nylander 2004). Markov

chains initiated from a random tree were run for 2,000.000 generations with samples

taken every 100th generation. The generation number excluded a burn-in of 200 000

generations; Tracer v1.4.1 (http://tree.bio.ed.ac.uk/software/tracer/) determined the

statistics.

The additional PHYML analysis for estimating maximum likelihood phylogenies was

run at the server Phylogeny.fr (Dereeper et al. 2008) in “A la Carte” mode. Alignments

were treated with Gblock to eliminate poorly aligned positions and ambiguous regions;

GTR substitution model was selected for the ITS dataset. Bootstrap branch support

values (BP) were estimated in PHYML under the maximum likelihood criterion using

default 100 replicates. There were 659 (89% of the original 790 positions) positions

analysed in PHYML after Gblock treatment.

The LSU sequences were not analysed due to the limited number of similar sequences

available in the Genbank.

Results

Both ITS and LSU sequence analyses showed 100% similarity among the

Hungarian collection and the holotypes of Collybia inusitata, C. bispora, and

Gymnopus inusitatus var. cystidiatus (Hungary) ... 293

bisporus JN247551

95/1 | catalonicus JN247552

inusitatus JN247553

inusitatus var. cystidiatus JN247550

88/0.72

hybridus DQ449980

37/0.68 aff. nivalis DQ449982

98/0.89 earleae DQ480094

19/0.34 earleae DQ449994

alkalivirens DQ480112

100/1 alkalivirens DQ450000

100/1 spongiosus DQ449993

spongiosus AF505784

ah 76/1/— erythropus DQ449996

erythropus DQ449998

impudicus DQ480109

brassicolens DQ449990

0.02

Fic. 1. Bayesian phylogram obtained from the ITS rDNA sequences of Gymnopus inusitatus and

other taxa of Gymnopus sect. Levipedes. Gymnopus brassicolens and G. impudicus were selected

as outgroup. Bootstrap support values of Maximum likelihood analysis (Phyml) and Bayesian

posterior probability values are given at branches. The bar indicates number of substitutions per

nucleotide position.

C. catalonica (Fic. 1), indicating that they are conspecific. Both phylogenetic

analyses produced almost identical tree topologies. The Bayesian tree had a

likelihood value -2192.343. Empirical base frequencies were pi(A) = 0.247,

pi(C) = 0.167, pi(G) = 0.210, and pi(T) = 0.377. Model information: parameter

a = 0.235; substitution rates A-C = 0.062, A-G = 0,495, A-T = 0.070, C-G = 0.012,

C-T = 0.286, G-T = 0.073; tree-length = 0.488.

294 ... Antonin, Finy & TomSovsky

Fic. 2. Gymnopus inusitatus var. cystidiatus (holotype). Scale bar = 10 mm. Photo P. Finy.

Taxonomy

Microscopical examination of the types and Hungarian collection noted

above confirmed the synonymy indicated by the sequence analyses. We propose

the following taxonomy for this species group of bisporic taxa:

Gymnopus inusitatus (Vila & Llimona) Vila & Llimona, Rev. Catal. Micol. 28: 180.

2006.

= Collybia inusitata Vila & Llimona, Rev. Catal. Micol. 24: 84. 2002.

= Collybia bispora J. Carbo & Pérez-De-Greg., Rev. Catal. Micol. 24: 278. 2002.

= Gymnopus bisporus (J. Carb6é & Pérez-De-Greg.) Vila &

Llimona, Rev. Catal. Micol. 28: 180. 2006.

= Collybia catalonica Vila & Llimona, Rev. Catal. Micol. 24: 80. 2002.

= Gymnopus catalonicus (Vila & Llimona) Vila & Llimona,

Rev. Catal. Micol. 28: 180. 2006.

Gymnopus inusitatus var. cystidiatus Antonin, Finy & Toméovsky, var. nov.

MycoBank MB 563276 FIGS 2-3

A varietate typica cheilocystidiis presentibus et oecologiae (sub Salice) differt.

Type: Hungary, Székesfehervar-Sdst6, 14.1X.2010, leg. P. Finy (Holotype, BRNM

737257; GenBank JN247550, JN247554).

Erymo ocy: having well-developed cheilocystidia.

BASIDIOMES similar to Mycetinis scorodonius. PILEUS up to 10 mm broad,

convex to applanate, with plane to depressed centre, smooth, slightly crenulate

Gymnopus inusitatus var. cystidiatus (Hungary) ... 295

Fic. 3. Gymnopus inusitatus var. cystidiatus (holotype).

a. cheilocystidia, b. basidiospores, c. basidia, d. pileipellis. Scale bar = 20 um.

296 ... Antonin, Finy & TomSovsky

at margin especially when old, brown to reddish brown (7-8E7) at centre, paler

towards margin. LAMELLAE rather distant, L = 14-17, | = 2-3, emarginate

and attached with tooth, + ventricose, whitish, with concolorous edge. STIPE

up to 35 x 0.75 mm (dry specimens), cylindrical, slightly broadened at apex,

cylindrical to slightly clavate at base, smooth, + concolorous with pileus at

base with paler upper part; basal mycelium brown to reddish brown. CONTEXT

without any distinct smell; taste not tested.

BASIDIOSPORES (10-)10.5-13 x 4.5-6.0 um, average = 11.3 x 5.3 um,

E = 1.9-2.7, Q = 2.15 (40 spores, 2 basidiomes, 1 collection), ellipsoid-

fusoid, pip-shaped, thin-walled. Basip1aA 26-33 x 7.5-8.5 um, 2-spored,

clavate. BASIDIOLES up to 43 x 4.0-9.0 um, cylindrical, clavate, subfusoid.

CHEILOCYSTIDIA 18-34(-40) x 7.0-12(-20) um, clavate, subutriform, often

capitate, sometimes irregular, smooth or with projection(s), thin-walled.

TRAMA HYPHAE + cylindrical, smooth or minutely incrusted, up to 12 um

wide. PILEIPELLIS a cutis composed of cylindrical to slightly inflated elements,

sometimes irregular or with projections, + thin-walled, coarsely incrusted

(zebroid), 3.0-10 um wide hyphae with greenish brown incrustation in KOH;

terminal cells cylindrical, clavate, fusoid, only minutely incrusted or smooth,

often irregular or with projections; some elements transient to poorly developed

dryophila-structure. STIPITIPELLIS a cutis of parallel, cylindrical, slightly

thick-walled, incrusted, up to 5.0 um wide hyphae. CauLocysTip1A absent,

scattered appressed to (sub)erect, cylindrical, rare terminal cells with scattered

projections present. CLAMP CONNECTIONS present.

CHEMICAL REACTIONS — No part of basidiocarps amyloid or dextrinoid.

ECOLOGY & DISTRIBUTION— Between Salix and grass remnants in open

sand calcareous dune near S. cinerea and S. purpurea. So far known only from

the type locality.

ADDITIONAL SPECIMENS EXAMINED — Gymnopus bisporus: Spain, Catalonia, Girona,

La Fonollera, Torroella de Mongri, 1 Oct. 2000, J. Carbo (holotype of Collybia bispora,

BCN-SCM B-4065, GenBank: JN247551, JN247555).

Gymnopus catalonicus: Spain, Catalonia, Barcelona, Can Ferrer, 23 Oct. 2000, J.

Vila & X. Llimona (holotype of Collybia catalonica, BCN-SCM B-4057), GenBank:

JN247552, JN247556.

Gymnopus inusitatus: Spain, Catalonia, Roses, Coll de la Perafita, 6 Oct. 1998, J. Vila

& X. Llimona (holotype of Collybia inusitata, BCN-SCM B-4058), GenBank: JN247553,

JN247557.

Discussion

Gymnopus inusitatus var. cystidiatus is characterized macroscopically (Fic.

2) by having basidiomes similar to Mycetinis scorodonius, and microscopically

(Fic. 3) by rather large basidiospores, long, 2-spored basidia, well-developed

clavate, subutriform, often capitate cheilocystidia, a pileipellis composed of

Gymnopus inusitatus var. cystidiatus (Hungary) ... 297

Fic. 4. Gymnopus inusitatus var. inusitatus. Marginal cells of lamellae (holotypes).

a. Collybia inusitata, b. C. bispora and c. C. catalonica). Scale bar = 20 um.

298 ... Antonin, Finy & TomSovsky

hyphae with greenish brown incrustation in KOH forming a cutis or only a

poorly developed dryophila-structure, and cylindrical, clavate, fusoid, often

irregular terminal cells, an absence of caulocystidia, and ecologically by growing

in sandy grasslands under Salix.

In this group of bisporic taxa, Antonin & Noordeloos (2010) distinguished

two species — Gymnopus bisporus (syn. G. catalonicus) with a brown to reddish

brown (marginally paler) pileus and basidiospores measuring 9.0-11(-12)

x 4.5-5.5(-6.0) um and G. inusitatus with a reddish brown to violaceous

brown pileus (cream to ochraceous-brown when drying), and slightly

narrower basidiospores measuring 10-14 x 4.0-5.0 um. Both species lack true

cheilocystidia (Fic. 4). The basidiospore size variation in this group of taxa is

distinctly wider than admitted by the current anatomic-morphological species

concepts. The same situation was noted for European species in the Psilocybe

cyanescens group (Borovicka 2008, Borovicka et al. 2010).

The most important diagnostic characters of G. inusitatus var. cystidiatus

are the presence of well-developed cheilocystidia and a different. All three taxa

synonymized under of G. inusitatus var. inusitatus were collected in xerothermic

acidophilous stands with various Cistus species (Pérez-De-Gregorio & Carbo

2002, Vila & Llimona 2002). Gymnopous inusitatus vat. cystidiatus was collected

in calcareous sandy grasslands under Salix. Its type locality lies in the Great

Hungarian Plain in the northwestern part of the Pannonic sand steppes where

the open xeric grassland is dominated by the Pannonian endemic Festuca

vaginata on dune tops and communities of Salix rosmarinifolia and other Salix

species in the moist interdune depressions. The Hungarian collection represents

the first record of G. inusitatus in central Europe.

Species from the G. dryophilus group differ from G. inusitatus by a well-

developed dryophila-structure in the pileipellis, tetrasporic basidia, and smaller

basidiospores. Gymnopus erythropus (Pers.) Antonin, Halling & Noordel. has

red-brown coloured hairs on the stipe base like G. inusitatus but differs in

basidiome colour, smaller basidiospores ((5.0-)5.5-8.0(-9.0) x 3.5-4.5(-5.0)

um), tetrasporic basidia, and differently shaped cheilocystidia (Antonin &

Noordeloos 2010).

The macroscopically similar Gymnopus earlei Murrill from North

America has a larger (< 35 mm broad) pileus, buff or pale orangish yellow

(later orangish buff) lamellae, a more robust stipe (10-46(-90) x 2-5 mm),

smaller basidiospores (5.6-7 x 2.8-3.5 um), mostly tetrasporic basidia, and

inconspicuous cheilocystidia (Halling 1983, Vilgalys & Miller 1983). Gymnopus

bicolor A.W. Wilson et al. has similar cheilocystidia and a poorly developed

dryophila-structure, but it has crowded lamellae, a larger (22-28 mm broad)

pileus, a more robust stipe (30-42 x 2-3 mm), tetrasporic basidia, and smaller

basidiospores (5.2-8 x 2.4-3.6 um; Wilson et al. 2004).

Gymnopus inusitatus var. cystidiatus (Hungary) ... 299

Acknowledgments

The authors are very grateful to the curator of the BCN herbarium for a loan of the

type specimens, to Zdenék Pouzar (Prague, Czech Republic) for correcting the Latin

diagnosis, and to Alexander Urban (University of Vienna, Vienna, Austria) and Alfredo

Vizzini (Universita degli Studi di Torino, Torino, Italy) for their critical review of the

manuscript. Studies of the first author were financially supported by the Ministry of

Culture of the Czech Republic (MK 00009486201).

Literature cited

Antonin V, Noordeloos ME. 2010. A monograph of marasmioid and collybioid fungi in Europe.

IHW- Verlag, Eching, 480 p.

Borovicka J. 2008. The wood-rotting bluing Psilocybe species in Central Europe - an identification

key. Czech Mycol. 60: 173-192.

Borovicka J, Noordeloos ME, Gryndler M, Obornik M. 2010. Molecular phylogeny of Psilocybe

cyanescens complex in Europe, with reference to the position of the secotioid Weraroa novae-

zelandiae. Mycol. Progress 10: 149-155. http://dx.doi.org/10.1007/s11557-010-0684-3

Dereeper A, Guignon V, Blanc G, Audic S, Buffet S, Chevenet F, Dufayard JF, Guindon S, Lefort

V, Lescot M, Claverie JM, Gascuel O. 2008. Phylogeny.fr: robust phylogenetic analysis for the

non-specialist. Nucleic Acids Res. 36(suppl 2): W465-W469.

Halling RE. 1983. The genus Collybia (Agaricales) in the northeastern United Sates and adjacent

Canada. Mycol. Mem. 8: 1-147.

Holmgren PK, Holmgren NH. 1998 (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/.

Kornerup A, Wanscher JH. 1983. Methuen handbook of colour. 3" edn. Methuen Co., London.

Mata JL, Hughes KW, Petersen RH. 2006. An investigation of /omphalotaceae (Fungi: Euagarics)

with emphasis of the genus Gymnopus. Sydowia 58: 191-289.

Nylander JAA. 2004. MrModeltest v2. Evolutionary Biology Centre,Uppsala.

Pérez-De-Gregorio MA, Carbé J. 2002. Una nueva especies de Collybia (Fr.: Fr.) Staude, encontrada

en Catalufia. Rev. Catal. Micol. 24: 277-282.

Ronquist F, Huelsenbeck JP. 2003. Mr. Bayes 3: Bayesian phylogenetic inference under mixed

models. Bioinformatics 19:1572-1574.

TomSovsky M, Jankovsky L. 2008. Validation and typification of Laetiporus montanus. Mycotaxon

106: 289-295.

Vila J, Llimona X. 2002. Noves dades sobre el component ftingic de les comunitats de Cistus de

Catalunya. Rev. Catal. Micol. 24: 75-121.

Vilgalys R, Miller OK Jr. 1983. Biological species in the Collybia dryophila group in North America.

Mycologia 75(4): 707-722.

Wilson AW, Desjardin DE, Horak E. 2004. Agaricales of Indonesia. 5. The genus Gymnopus from

Java and Bali. Sydowia 56(1): 137-210.

MY COTAXON

http://dx.doi.org/10.5248/119.301

Volume 119, pp. 301-306 January-March 2012

New records of Pezizales from Pakistan

T. ASHRAE*! & A.N. KHALID?

Department of Botany, University of the Punjab, Quaid-e-Azam Campus, Lahore, 54590, Pakistan

CORRESPONDENCE TO *: *'tayibamaan01@gmail.com e& *drankhalid@gmail.com

ABSTRACT — Geopora cooperi f. cooperi and Peziza succosa have been described and

illustrated morphologically and by amplifying ITS1-5.8S-ITS2 region of rDNA. Both taxa are

new records from Pakistan.

KEY WORDS — ascocarp, ascospores, operculate ascus

Introduction

Pezizales are characterized by unitunicate asci that generally open by

rupturing to form a terminal or eccentric lid or operculum (Hansen & Pfister

2006). Of the 1638 pezizalean species known, Geopora Harkn. and Peziza Dill.

ex Fr. are represented by 13 and 104 species respectively (Kirk et al. 2008).

From Pakistan three Geopora species — G. arenicola (Lév.) Kers, G. arenosa

(Fuckel) S. Ahmad, G. foliacea (Schaeff.) S. Ahmad) — and eight Peziza species

— P. badiofusca (Boud.) Dennis, P cerea Bull., PR gerardii Cooke, P. micropus

Pers., P. pakistanica (S. Ahmad) S. Ahmad, P. repanda Pers., P. vesiculosa Bull.,

P. violacea Pers.) — have been reported (Ahmad 1978; Ahmad et al. 1997).

This paper describes two taxa of Pezizales newly recorded from Pakistan. The

specimens were first identified morphologically, after which the ITS1+5.8S+

ITS2 rDNA regions were amplified for comparison with GenBank sequences

of related species. Eighty-seven taxa of Pezizales were previously reported from

Pakistan; the addition of our recently collected Geopora cooperi f. cooperi and

Peziza succosa) raises that total to 89.

Materials & methods

Collected ascocarps were photographed and necessary ecological data were recorded

in field. The specimens were dried with the help of fan heater and preserved with

tentative field No., for further studies. Apothecia were rehydrated and free hand sections

were made in the laboratory. Microscopic (morpho-anatomical) characters were noted

at 10x and 40x and drawings were made with the help of camera lucida.

302 ... Ashraf & Khalid

DNA was extracted from dried ascomata, using Extract N. Amp.™ Plant kit (SIGMA).

nrDNA of the ITS1-5.8S-ITS2 was amplified using fungal specific and universal primers

pairs (ITSIF/ITS4 and ITS1/ITS4, respectively) under the following parameters:

denaturation at 94°C for 4 min, then with 35 cycles of 45 s at 94°C, 45 s at 54°C and 1

min 30 s at 72°C, and a final amplification at 72°C for 2 min. Bidirectional sequencing

of the purified PCR products was done by Macrogen Inc. (South Korea). ITS sequences

were compared using BLAST in GenBank to confirm our identifications.

The specimens have been deposited in the Pezizales collection of LAH Herbarium,

Department of Botany, University of the Punjab, Lahore, Pakistan. Sequences have been

accessioned in GenBank.

Taxonomy

Geopora cooperi Harkn., Bull. Calif. Acad. Sci. 1: 168 (1885), f. cooperi PLATE 1

Ascocarp:5-5.5cm broader, globose to oblong, irregularly convoluted, outer

surface dark brown and covered with light brown hairs, inner surface pinkish

white to white, highly convoluted, convolutions close but separate. HYMENIUM:

200-250 um thick. Asci: Long cylindrical, operculate, centric operculum,

unitunicate, 8-spored, uniseriate, non-amyloid, tapering at base, (213-)

221-260 x (16-)19-21 um. Ascospores: Broadly oval-elliptical, hyaline,

smooth, thin walled, with a large central guttule, cytoplasm dense around

the guttule, (19-)20-23(-25) x 13-14 um. Paraphyses: Long, slender, septate,

swollen at the tips, nearly of same level as ascus, apical cell 14.5 um broad.

ExcIpuLuM: Ectal excipulum: textura globulosa, 120 um, 3-4 cells thick, cells

oblong to circular in section, smooth, walls thick and dark brown, 24-40 x

18-27 um, cells on the outside darker than the inner cells. Outer cells of the

excipulum forming abundant, hyaline to light brown hairs, that are flexuous,

branched, 11-13 um wide, thick-walled, (walls up to 3 um). Medullary

excipulum of textura intricata, thin walled hyphae, hyaline to light brown.

MATERIAL EXAMINED: PAKISTAN: GILGIT-BALTISTAN, Fairy Meadows, 3036 m

(9960ft) a.s.l., under Pinus wallichiana A.B. Jacks., solitary, on ground, 24 July 2010,

A.N. Khalid TA-50. (LAH 240710; GenBank JN558642).

COMMENTS: Geopora cooperi differs from other epigeous Geopora species by

producing truffle-like ascocarps with a small basal opening and a heavily folded

layer (Tamm et al. 2010). It differs from most other truffle-like fungi because

of the forcible discharge of ascospores: truffle spores are usually not forcibly

expelled from the ascus. Mycorrhizal interaction of Geopora cooperi with Pinus

ponderosa P. Lawson & C. Lawson and the orchid Epipactis atrorubens (Hoftm.)

Besser has been established (Sheffereson et al. 2008). Geopora cooperi f. cooperi

differs from G. cooperi f. gilkeyae Burds. in the narrower oval to elliptical

ascospores (Burdsall 1968). Our DNA sequence analyses matched the 625

nucleotide ITS region (ITS1-5.8S-ITS2) from the Pakistani collection with that

Pezizales new to Pakistan ... 303

PLATE 1. Geopora cooper f. cooperi: A-B. Different views of apothecia (arrow indicates outer hairy

layer of fruiting body). C. Section of hymenium showing asci and paraphyses. D. Ascospores.

E. Ectal excipulum. Scale bars: A = 1.5 cm; B = 1 cm; C = 40 um; D = 13.5 um; E= 24 um

304 ... Ashraf & Khalid

oo.

\ x

\, \

\ H

\ \

. \

H n.

ym)

i t

it \

ea)

i i |

\ 5 /

1 ij

i ‘i

PLATE 2. Peziza succosa: A. Apothecia. B. Section of apothecium showing hymenium, subhymenium

and excipulum. C. Asus and paraphysis. D. Ascospores. Scale bars: A = 1.1 cm; B = 190 um; C = 49.1

um; D = 9.5 um.

Pezizales new to Pakistan ... 305

from Geopora cooperi f. cooperi (FR694202.1) at 98% similarity and 90% Query

coverage, which we consider sufficient to refer our collection to this taxon.

Peziza succosa Berk., Ann. Mag. Nat. Hist. 6: 358 (1841) PLATE 2

APOTHECIA: Usually gregarious, cupulate, deeply concave, medium-

sized, disc concave at first, hemispherical and later slightly irregular,

2.5-3cm in diam., sessile to subsessile, margins entire turning inwards,

purplish brown hymenium with slight yellowish tinge, outer surface slightly

lighter than hymenium, fresh apothecia exudes yellowish juice when damaged,

Subhymenium plectenchymatous. HyMENIUM: up to 304-315 um. AscrI:

Unitunicate, operculate, 8-spored, amyloid at tip, cylindrical slightly narrow

near base with flexuous stalk, 260-300 x 17-19um. AscosporEs: Uniseriate,

elliptical, ornamented with warts, biguttulate,19-20 x 10-11(-12) um.

PARAPHYSES: Slender, straight, septate, 3.5-4 um in diam. ExcrpuLum: Ectal

excipulum 95 um thick, textura globulosa to angularis, cells hyaline to slightly

brown, smaller thin-walled cells also present; medullary excipulum 285 um

thick epidermoid to polygonal cell, hyaline, thin-walled, 14-25 um in diam.

MATERIAL EXAMINED: PAKISTAN: KHYBER PAKHTUNKHWA, Himalayan Moist

Temperate Forests, Khanspur, 2575 m (8205 ft) a.s.l, gregarious, on ground, mossy

substrate, 21 Aug 2010, T. Ashraf TA-134 (LAH 210810; GenBank JN588568).

ComMENTs: Peziza succosa has strongly amyloid asci, with the bluing reaction

strongest at the apex, although no pronounced bluing ring is present. With other

ectomycorrhizal fungi, P. succosa has been reported in deciduous forests and is

widespread in Europe, North America (Hansen et al. 2001, Michell 2006), and

Asia (DH Pfister, pers. comm.). ITS sequence analyses of our collection showed

it to be 99% identical with P succosa (DQ200840.1) from Fredericksburg Have,

Zealand, Denmark.

Acknowledgements

The authors would like to thank Prof. Dr. D. H. Pfister (Harvard University Herbaria,

Cambridge USA) and Dr. A. R. Niazi (Department of Botany, University of the Punjab,

Lahore) for their helpful comments and critical review of this manuscript.

Literature cited

Ahmad §S, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Sultan Ahmad Mycological Society of

Pakistan, Lahore. 248 p.

Ahmad S. 1978. Ascomycetes of Pakistan. Biological Society of Pakistan, Monograph 7: [i]-vii,

1-236.

Burdsall HH. 1968. A revision of the genus Hydnocystis (Tuberales) and of the hypogeous species of

Geopora (Pezizales). Mycologia 60(3): 496-525. http://dx.doi.org/10.2307/3757418

Hansen K, Pfister DH. 2006. Systematics of the Pezizomycetes—the operculate discomycetes.

Mycologia 98(6): 1029-1040. http://dx.doi.org/10.3852/mycologia.98.6.1029

306 ... Ashraf & Khalid

Hansen K, Lzessge T, Pfister DH. 2001. Phylogenetics of the Pezizaceae, with an emphasis on Peziza.

Mycologia 93(5): 958-990. http://dx.doi.org/10.2307/3761760

Kirk PM., Cannon PF, David JC, Stalpers JA. 2008. Ainsworth and Bisby’s dictionary of the fungi,

10" ed. CABI: Wallingford, Oxon. 771 p.

Michell K. 2006. Field guide to mushrooms and other fungi of Britain and Europe. New Holland

Publishers. 192 p.

Shefferson RP, Kull T, Tali K. 2008. Mycorrhizal interaction of orchid colonizing Estonian mine

tailings hills. American Journal Botany 95: 156-164. http://dx.doi.org/10.3732/ajb.95.2.156

Tamm H, Poldmaa K, Kullman B. 2010. Phylogenetic relationships in genus Geopora

(Pyronemataceae, Pezizales). Mycological Progress 9:509-522.

http://dx.doi.org/10.1007/s11557-010-0659-4

MY COTAXON

http://dx.doi.org/10.5248/119.307

Volume 119, pp. 307-313 January-March 2012

A new Hysterostomella species from the Cerrado

in Brasilia National Park

C.A. INAcIOo*, R.C. PEREIRA-CARVALHO, E.S.C. SouZA,

H.B. SALES & J.C. DIANESE.

Departamento de Fitopatologia, Universidade de Brasilia, Distrito Federal, 70.910-900, Brazil

* CORRESPONDENCE TO: inacio@unb.br

ABSTRACT — A new Hysterostomella species found on leaves of Connarus suberosus is

illustrated and described as Hysterostomella connari.

Key worps — Ascomycota, biodiversity, Parmulariaceae, taxonomy, tropical fungi

Introduction

Parmulariaceae, including 34 genera of foliicolous biotrophic fungi, occurs

primarily in the Neotropics and Paleotropics (Inacio & Cannon 2008, Kirk et

al. 2008). The type genus of the family, Parmularia Lév. and its type species

P. styracis Lév., was described on Styrax ferrugineus (Léveillé1846), a plant

species endemic to the Brazilian Cerrado. For decades this family was set aside

with occasional contributions (Miller & von Arx 1973, von Arx & Miller 1975,

Barr 1987, Sivanesan 1970, Sivanesan & Hsieh 1989, Sivanesan et al. 1998).

Recently the type species of all accepted genera were re-examined, consolidating

our understanding of the family on the basis of morphological features (Inacio

2003, Inacio & Cannon 2003, 2008). Now a new species belonging to the genus

Hysterostomella, previously observed by Sales (2003) and Silva et al. (2006)

from the Brazilian Cerrado, is formally described and illustrated.

The revision by Inacio (2003) of Hysterostomella Speg., type species

H. guaranitica Speg., included a re-description of several species. The genus is

characterized by subcuticular internal stromata when host cells are incorporated

by the infective mycelium. Sometimes these structures are subepidermal,

connected to superficial stromatic ascomata by peg-like or tubular hyphal

filaments. The discoid multilocular stromatic ascomata contain spherical

or elongate locules dispersed in various orientations (Inacio 2003, Inacio &

Cannon 2008). Hysterostomella is close to Cycloschizon Henn., which differs by

308 ... Inacio & al.

forming internal stromata that become erumpent via a column with a wart-like

expanded top containing locules distributed in complete or incomplete rings.

Dothidasteroma Hohn., also close to Hysterostomella, differs mainly by forming

flat internal stromata, never subepidermal, that connect to the superficial

ascomata by either bundles of dark colored hyphae, or single hyphae.

Materials & methods

Leaf samples of Connarus suberosus containing dark brown to black crustose fungal

structures were collected from different areas of Cerrado in Parque National, Brasilia.

These were pressed and dried for 3-4 days at + 50°C, and deposited in the Mycological

Collection of the Herbarium UB. The leaves were observed under a stereomicroscope

to describe the lesions and location and characteristics of the fungal colonies. For the

morphological studies, each specimen was mounted on slides using lacto-glycerol/

cotton blue or glycerol KOH/basic phloxine, to observe the internal structures. Pieces

of the dried material containing stromata were re-hydrated for 4-12 h using an aqueous

solution containing 10% ethanol and 0.1% Tween-20. Then 7-15 um thick sections

were produced using a freezing microtome (MICROM GMBH, HM 500 OM, Micron

Laborgerate, Walldorf, Germany). A compound light microscope (Zeiss Ultraphot

I, Carl Zeiss, Oberkochen, Germany) was used for the photographic documentation

and to obtain morphometrical data. Pieces of leaves with one or more lesions showing

representative samples of fruiting bodies were examined with SEM after treatment in a

0.1 M sodium cacodylate buffer (pH 7.4) containing 2% glutaraldehyde for at least 24 h.

The samples were dehydrated in an increasing acetone concentration series (15%, 30%,

50%, 75%, 100%; 15 min per concentration). Leaf pieces were then dried at the critical

point before being covered by a thin layer of gold in a sputter coater for 2 min. Finally,

the samples were observed in a scanning electron microscope (Jeol, model JSM 840-A

Taxonomy

Hysterostomella connari Inacio, Pereira-Carvalho, E.S.C. Souza, H.B. Sales &

Dianese sp. nov FIGs 1-21

MycoBank MB 561556

Species haec ab Hysterostomella tetracerae differt conidiis non fasciatis et grandioribus.

Type: BRAZIL. Distrito Federal, Brasilia, Parque Nacional, near Parada do Cristal, on

leaves of Connarus suberosus Planch. (Connaraceae), 18 Jan. 2010, leg. Carlos Antonio

Inacio (holotype, UB Mycol. Coll. 21492).

EryMo_oey: referring to the host genus.

Lesions 3-12 mm diam, black, circular encircled by a reddish linear band,

adaxially more evident. Colonies sparse, containing black circular or irregular

superficial stromata that are discoid, mostly epiphyllous, crustose, sheltering

conidiomata, spermogonia and/or ascomata.

INTERNAL PSEUDOSTROMATA as mycelium with colourless to reddish-brown

angular cells invading and incorporating one or more host cell layers, to 30 um

Hysterostomella connari sp. nov. (Brazil) ... 309

Fics 1-4. Hysterostomella connari on leaves of Connarus suberosus. 1. Adaxial colony shown as

a group of stromatic ascomata circular to irregular in shape. 2. Crustose stromatic ascomata (H)

together with a colony of an Asterina species (A). 3. A stromatic ascoma bordered by a reddish

band (bar). 4. An older opened ascoma (left) and another still closed (right) seen in SEM. Scale

bars: 1 = 10 mm; 2 = 2 mm; 3,4 =0.5 mm.

deep. Superficial stromata 0.1-3.5 mm diam., 50-113 x 150-680 um in vertical

section, subepidermal, erumpent, brown to dark brown, shiny, with laciniate

edges; upper surface composed of textura prismatico-radiata, the hyphae with

brown prismatic cells that proliferate dichotomously towards the margins to

form a circular or irregular 1-2 cell-layered multilocular shield that houses the

sexual and asexual fruiting bodies; rather prominent, opening by irregular slits;

upper wall 10-30 um thick, brown to dark brown, with surface consisting of

5-7 um diam brown cells; lower wall 30 um thick, brown to dark brown, covering

the host epidermal cells, dense, bearing a basal cushion of mingled colourless,

light brown to brown hyphae, bearing the asci or lining the internal surface

of the conidiomata and spermogonia. Mycelium internal. Hyphae colourless

to pale brown, 1-2 um diam, septate, branched, penetrating and occupying

the epidermal and first layers of mesophyll, sometimes with coiled intracellular

structures; to 100 um deep.

ConripiIoMaTA included in the superficial stromata, gregarious, intermixed

with the ascomata and spermogonia, globose to subglobose, opening by

diametral or irregular fissures, uni- or multilocular, 40-160 um high in vertical

310 ... Inacio & al.

Fics 5-8. Hysterostomella connari on leaves of Connarus suberosus. 5. Superficial textura of a

stromatic ascoma, with arrows showing the dichotomous proliferation of component hyphae.

6. Surface of an ascoma seen under SEM. 7. Vertical section through an ascoma showing parallel

asci and branched sterile filaments, and transcuticular hyphae connecting the flat portion of the

pseudostroma with the superficial ascoma. 8. Detail of the interconnection between the components

of the internal pseudostroma with the superficial ascoma. Scale bars: 5, 6, 8 = 10 um; 7 = 50 um.

section, with a brown to dark brown upper wall; UPPER WALL dense, 10-25

(-30) um thick, composed of textura prismatico-angularis; CELLS 3-7 um diam.;

CONIDIOMATAL LOCULES 25-150 x 120-610 um, single or sometimes multiple

separated by a stromatic wall, with conidiogenous cells and sterile filaments

attached to the inner face of the upper locular wall; CONIDIOGENOUS CELLS

8-20(-25) x 3-7 um, ampulliform, cylindrical to lageniform, colourless to

pale brown, smooth, holoblastic; conip1A 9-16 x 6-10 um, initially colourless,

becoming pale brown to brown, guttulate, obovoid to widely clavate, aseptate,

smooth, with a truncate base. SPERMOGONIA structurally and dimensionally

similar to the conidiomata; sPERMOGENOUS CELLS 2-7 x 2-3 um, phialidic,

hyaline, cylindrical; sPERMATIA 3-5 x 1-2 um, aseptate, hyaline, bacilliform.

ASCOMATA 50-95 x 155-580 um, intra-stromatic, occurring separately

from, or mixed with conidiomata and/or spermogonia occupying the same

stromatic crust, containing parallel asci and sterile filaments on a basal cushion

of colourless to light brown hyphae covered by a thin gelatinous layer; YOUNG

ASCI variable in shape, cylindrical, clavate to broadly clavate, with a short

Hysterostomella connari sp. nov. (Brazil) ... 311

: fe .

Fics 9-11. Hysterostomella connari on leaves of Connarus suberosus. 9. Immature asci. 10. A mature

bitunicate ascus with dark brown ascospores. 11. Ascospores. Scale bars: 9-11 = 10 um.

pedicel, thick-walled particularly towards the apex where, before spores are

visible, a sub-apical chamber is present; FULL-SIZED ASCI showing 60-70 x

18-30 um, 6- to 8-spored with ascospores arranged in one to three rows,

broadly-clavate, thick-walled particularly in the upper part, collapsing after

spore release showing a large apical crack in a typically fissitunicate fashion;

ASCOSPORES 17-21 x 7-11 um, becoming light to dark brown, 1-septate,

guttulate, covered by a mucilaginous sheath, ellipsoidal, round at both ends,

narrower at the septum, tapering towards the lower tip; INTERASCAL FILAMENTS

septate, smooth, colourless to light brown, thin-walled, 2-4 um diam, filiform,

round or slightly attenuate at the tip, branched above the middle portion,

immersed in a gelatinous matrix.

ADDITIONAL SPECIMENS EXAMINED: BRAZIL. DISTRITO FEDERAL, BRASILIA, Parque

Nacional: 2 km North from Parada do Cristal—on leaves of Connarus suberosus, 24 Jan.

2007, leg. Rafael Guimaraes, collection number 6 (UB Mycol. Coll. 20.420); Parada do

Cristal ‘s pathway—on leaves of Connarus suberosus, 06 Feb. 2006, Siomara Vasconcelos

da Silva (UB Mycol. Coll. 20.115); Unknown sites within Parque Nacional—08 Jan.

1996, Zuleide Martins Chaves 302 (UB Myc. Coll. 10.656) & 321 (UB Myc. Coll. 10.675);

15 Jan.1996, Zuleide Martins Chaves 350 (UB Myc. Coll. 10.738); 24 Jan. 1996, Mariza

Sanchez 1476 (UB Myc. Coll. 10.911); 20 Jan. 2010, Gustavo Puga Lemes (UB Myc. Coll.

21529) and Carlos Antonio Inacio (UB Mycol. Coll. 21516)

Discussion

Inacio (2003) and Inacio & Cannon (2008) have partially revised

Hysterostomella. All species are assumed to show clear host specificity, and no

312 ... Inacio & al.

Fics 12-17. Hysterostomella connari on leaves of Connarus suberosus. 12. An ascoma (left) beside

a spermogonium (right). 13. A conidioma with a typically inverted hymenium. 14. Holoblastically

formation of a conidium (arrow). 15. A group of immature hyaline conidia. 16. Mature dark brown

aseptate conidium (arrow). 17. Spermatia. Scale bars: 12, 13 = 50 um; 14, 15, 17 = 10 um; 16 = 5

um.

Hysterostomella species has previously been recorded associated with members

of the Connaraceae. Among the species with re-examined type materials, only

H. tetracerae (Hohnel 1909) and H. gymnosporiae (Hansford 1947) have a

conidial anamorph similar to that found in H. connari. However both differ

because their conidia have a clear hyaline median band. In the remaining species

there are no records of anamorphs. Spermogonia as shown in H. connari were

previously reported only in H. tetracerae, which, however, differs additionally

by its larger ascomata (88-188 um diam.), darker and easily discernible lower

ascomatal wall, and narrower asci. Hysterostomella gymnosporiae forms

more prominent ascomata (170-250 um high) and more restricted internal

pseudostromata than H. connari.

Hysterostomella connari sp. nov. (Brazil) ... 313

Acknowledgments

The authors wish to thank Ms. Mariza Sanchez (Herbarium UB, Mycological

Collection), Siomara Vasconcellos Silva and Gustavo Puga Lemes for technical assistance,

CNPq (Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico), FAPDF,

FINATEC (Universidade de Brasilia) and Paul FE Cannon (CABI Bioscience, UK) and

Richard Hanlin (University of Georgia, U.S.A.) for peer reviewing this manuscript. The

first author thanks the Universidade de Brasilia (UnB), particularly to the Department

of Plant Pathology for supporting this study.

Literature cited

Barr ME. 1987. Prodromus to the class Loculoascomycetes. Printed by Margaret E. Barr Bigelow.

USA. 168 p.

Hansford CG. 1947. New tropical fungi. Proceedings of the Linnean Society of London 159:

21-42.

Hoéhnel FXR von. 1909. Fragmente zur Mykologie (VIII. Mitteilung, Nr. 407 bis 467).

Sitzungsberichten der Kaiserliche Akademie der Wissenschaften in Wien Mathematische-

naturwissenschaftliche Klasse, Abt.1, 118: 1461-1552.

Inacio CA. 2003. A monograph of the Parmulariaceae. PhD thesis, University of London, London,

UK.

Inacio CA, Cannon PF. 2003. Viegasella and Mintera, two new genera of Parmulariaceae

(Ascomycota) with notes on the species referred to Schneepia. Mycological Research 107(1):

82-92. http://dx.doi.org/10.1017/S0953756202007013

Inacio CA, Cannon PF. 2008. The genera of Parmulariaceae. CBS, Utrecht, The Netherlands.

192 p.

Kirk PM, Cannon PF, Minter DW, Stalpers JA (Eds). 2008. Dictionary of the fungi, 10th edition.

CABI Publishing, UK.

Léveillé JH. 1846. Description des champignons d‘herbier du Muséum de Paris. Annales des

Sciences Naturelles, Botanique, Série 3, 5: 249-304.

Miller E, von Arx JA. 1973. Pyrenomycetes: Meliolales, Coronophorales, Sphaeriales. 135-219, in:

CG Ainsworth et al. (eds). The Fungi, 1V A. Academic Press, New York, London.

Sales HB. 2003. Alguns fungos associados a plantas do cerrado. Brasilia: Universidade de Brasilia.

78 p. [Master dissertation].

Silva, SV, Inacio CA, Sanchez M. 2006. Study of Parmulariaceae from cerrado vegetation of Parque

Nacional-Brasilia, DE In: XXXIX Congresso Brasileiro de Fitopatologia, 2006, Salvador, BA.

Fitopatologia Brasileira 31: $345-S345.

Sivanesan A. 1970. Parmulariopsella burseracearum gen. et sp. nov. and Microcyclus placodisci sp.

nov. Transactions of the British Mycological Society 55(3): 509-514.

Sivanesan A, Hsieh WH. 1989. Kentingia and Setocampanula, two new ascomycete genera.

Mycological Research 93(1): 83-90. http://dx.doi.org/10.1016/S0953-7562(89)80141-8|

Sivanesan A, Hsieh WH, Chen CV. 1998. A new monotypic genus of a parmulariaceous

dictyosporous ascomycetes from Taiwan. Botanical Journal of the Linnean Society 126:

323-326. http://dx.doi.org/10.1111/j.1095-8339.1998.tb01384.x

von Arx JA, Miller E. 1975. A re-evaluation of the bitunicate ascomycetes with keys to families and

genera. Studies in Mycology 9: 1-159.

MY COTAXON

http://dx.doi.org/10.5248/119.315

Volume 119, pp. 315-328 January-March 2012

Penicillium mallochii and P. guanacastense,

two new species isolated from Costa Rican caterpillars

KAROL G. RIVERA’, JOEL Diaz’, FELIPE CHAVARRIA-DIAz?’,

MarIA GARCIA’, MIRJAM URB?*, R. GREG THORN3,

GERRY LOUIS-SEIZE', DANIEL H. JANZEN® & KEITH A. SEIFERT"

‘Biodiversity (Mycology), Eastern Cereal and Oilseed Research Centre,

Ottawa, Ontario K1A 0C6, Canada

?Area de Conservacén Guanacaste, Apartado Postal 169-5000, Liberia, Guanacaste, Costa Rica

*Department of Biology, University of Western Ontario, London, Ontario, N6A 5B7, Canada

‘Present address: Department of Microbiology and Immunology, McGill University,

Montreal, Quebec H3A 2B4, Canada

° Department of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA

* CORRESPONDENCE TO: keith. seifert@agr.gc.ca

ABSTRACT — Twenty-five strains of monoverticillate Penicillium species were isolated from

dissected guts and fecal pellets of leaf-eating caterpillars reared in the Area de Conservacién

Guanacaste, Costa Rica, or from washed leaves of their food plants. Phylogenetic analyses

of B-tubulin, nuclear ribosomal internal transcribed spacer (ITS), cytochrome c oxidase

subunit 1, translation elongation factor 1-a and calmodulin gene sequences revealed two

phylogenetically distinct, undescribed species closely related to P sclerotiorum. Penicillium

mallochii was isolated from Rothschildia lebeau and Citheronia lobesis (Saturniidae) and

their food plant Spondias mombin (Anacardiaceae) and P. guanacastense from Eutelia sp.

(Noctuidae). Both fungi produce greenish conidial masses and orange pigments in agar culture,

have smooth-walled, monoverticillate conidiophores with moderately vesiculate apices, and

globose to subglobose conidia. The species morphologically resemble P. sclerotiorum but

differ subtly in vesicle width and conidial shape.

Key worps — Penicillium subg. Aspergilloides, sect. Sclerotiora, DNA barcoding, All Taxa

Biological Inventory (ATBI)

Introduction

In 2001, a microbial observatory was established in Sector Santa Rosa

of the UNESCO World Heritage Site, Area de Conservacién Guanacaste,

northwestern Costa Rica, to culture microbiota (including fungi) associated

with caterpillars reared on the tree Spondias mombin as a model for elucidating

ecological roles of insects and associated microbes in a tropical forest ecosystem.

316 ... Rivera & al.

The fungi isolated from dissected caterpillar guts and fecal pellets were the

subject of an MSc thesis (Urb 2006) and included many Penicillium strains that

required detailed taxonomic study to identify. Several strains were identified

as the common and widespread biverticillate Penicillium citrinum Thom, but

a significant number had unbranched, monoverticillate conidiophores and

colonies reminiscent of P. sclerotiorum J.-H. Beyma (Pitt 1979).

As part of the Canadian Barcode of Life Network, we explored DNA

barcoding of Penicillium species using cytochrome oxidase 1 (cox1; Seifert

et al. 2007) and the proposed fungal DNA barcode, the internal transcribed

spacer (ITS) of the nuclear ribosomal DNA. In this paper, two new species,

P. mallochii and P. guanacastense are formally described from the Area de

Conservacion Guanacaste, as part of a broader revision of the P. sclerotiorum

complex (Rivera & Seifert 2011). The fungi were characterized using cultural

and microscopic characters, DNA barcode data, and additional DNA sequence

data from B-tubulin (BENA), translation elongation factor 1-a (TEF1-a), and

calmodulin (cmp) genes.

Materials & methods

Isolates

Caterpillars of Rothschildia lebeau and Citheronia lobesis (Saturniidae) feeding on

Spondias mombin (Anacardiaceae), Cochlospermum vitifolium (Bixaceae), or Exostema

mexicanum (Rubiaceae) were collected by Joel Diaz, Felipe Chavarria-Diaz, and

Maria Garcia in the tropical dry forest of Sector Santa Rosa of Area de Conservacién

Guanacaste, northwestern Costa Rica. Caterpillars were surface sterilized with dH,O and

ethanol and guts were removed aseptically and, when possible, partitioned into foregut

plus midgut and hindgut sections. Gut contents or feces (obtained from caterpillars kept

in sterile Petri dishes) were suspended in sterile H,O and serially diluted in sterile dH_,O,

then plated on soil extract-rose bengal agar (Bills & Foster 2004). Fungal colonies were

transferred to Malt Extract Agar (MEA, Samson et al. 2004) until pure cultures were

obtained. Cultures are preserved in the Canadian Collection of Fungal Cultures (CCFC,

Table 1) and ex-types are also deposited in the CBS-KNAW Fungal Biodiversity Centre

culture collection (CBS), Utrecht, the Netherlands. Cultures lacking DAOM numbers

are preserved in the personal collection of K. Seifert (KAS) or R.G. Thorn (RGTHC).

Details for cultures of other species are given by Rivera & Seifert (2011).

BLAST searches of ITS and BENA sequences suggested that the new species were

related to Penicillium sclerotiorum. When possible, we obtained five strains per species of

this clade (Peterson 2000) for comparison, including P. sclerotiorum, P. adametzii K.M.

Zalessky, P. bilaiae Chalab., P herquei Bainier & Sartory and P. hirayamae Udagawa.

Strains were obtained from CBS and the USDA-ARS, National Center for Agricultural

Utilization Research culture collection (NRRL).

Morphology

For standardized descriptions, spore suspensions in 0.1% semi-solid agar were

inoculated at three points on Czapek Yeast Agar (CYA, Pitt 1979) with added micro-

Penicillium mallochii & P. guanacastense spp. nov. (Costa Rica) ... 317

nutrients (Samson et al. 2004), and MEA in 9 cm polystyrene Petri dishes, and incubated

at 25 °C for 7 din the dark (Samson & Pitt 1985). To determine whether sclerotia could

be induced, some strains were incubated for up to 6 months in incident light at room

temperature on Oatmeal Agar (OA, Samson et al. 2004). Colony colors were determined

using Kornerup & Wanscher (1978); alphanumeric color codes refer to this work.

Colony photographs were taken with a Nikon CoolPix E 5000 camera with incandescent

lighting and a copy stand.

Microscopic examinations employed an Olympus BX 50 light microscope on material

removed from MEA colonies and mounted in 85% lactic acid. Microphotographs were

taken with an Evolution MP Camera and microscopic characters were measured from

digital images using Image-Pro Plus v. 6 (Media Cybernetics, MD, U.S.A.). For each

strain, ten phialides, stipes, metulae and vesicles were measured manually, and 25

automated measurements were made for conidia.

DNA isolation, PCR amplification, DNA sequencing and phylogenetic analyses

Detailed protocols are provided in the revision of the P sclerotiorum complex by

Rivera & Seifert (2011), along with details of PCR and sequencing of some strains, and

only a summary of relevant details is provided here. DNA was extracted from agar

cultures using UltraClean™ Microbial DNA Isolation Kits (MO BIO Laboratories, Inc.,

Montreal, Canada) following the manufacturer's protocol. Genes were amplified by PCR

and sequenced with the following primers: BENA Bt2a and Bt2b (Glass & Donaldson

1995); ITS ITS1 and ITS4 (White et al. 1990); cox1 PF and AR (Seifert et al. 2007); TEF1-a

EF Ic and EF6 (Peterson et al. 2004); cmp CMD5 and CMD6 (Hong et al. 2006).

PCR reactions were performed in 10 ul reaction mixtures with 1 ul genomic DNA,

1X PCR Buffer, 0.1 mm of dNTPs, 0.08 uM of each primer, and 0.5X Taq polymerase, in

a TGradient (Biometra, Montreal, Canada) or a Genius (Techne, Duxford, Cambridge)

thermocycler. Sequencing reactions were performed directly from PCR products in both

directions, with templates precipitated by ethanol/EDTA/sodium acetate precipitation.

Sequences were determined on an ABI 3130XL Genetic Analyzer (Applied Biosystems,

Foster City, California, USA).

Consensus sequences were assembled using Sequencher v. 4.8 (Genes Codes

Corporation, MI, USA). Alignments were constructed using the online version of

MAFFT v. 6 (Katoh et al. 2009), and adjusted to optimize homology using BioEdit

7.0.9 (Hall 1999). Maximum parsimony (MP) analyses were performed using heuristic

searches in PAUP v. 4 (Swofford 2002) with the tree bisection-reconnection (TBR)

branchswapping algorithm. Maximum likelihood (ML) analyses were performed

using Phylogenies for Maximum Likelihood (PhyML) v. 2.4.4 (Guindon et al. 2010).

Tree searches for each alignment were run under the nucleotide substitution model

obtained from ModelTest 3.7, using the Akaike Information Criteria (AIC) (Posada

and Crandall, 1998), and starting from a tree obtained for branch swapping using the

modified neighbor joining algorithm BIONJ (Gascuel 1997) as implemented by PhyML.

Bayesian inference (BI) analyses were performed using MrBayes 3.1.2 (Huelsenbeck and

Ronquist 2001), under the same models selected for ML. All Bayesian analyses were

performed using random starting trees, and for each gene, four chains were run for one

million generations, sampling every 100 generations, generating 10,001 trees, with the

first 2,500 discarded as ‘burnin’ for each chain.

318 ... Rivera & al.

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320 ... Rivera & al.

All sequences for the two new species are deposited in GenBank with the accession

numbers JN625955-JN626181 (TABLE 1). Details of strains of other species, and the

GenBank accession numbers for their sequences, are provided by Rivera & Seifert

(2011).

Results

Representative ML trees for each of the five genes (ITS, coxl, BENA, CMD

and TEF1-a) are presented in Fic. 1. The MP and BI trees are not shown but

differences in topology are discussed, and support values for each analysis are

mapped onto the ML trees and included in TABLE 2. A combined gene tree is

included in the revision of the entire P. sclerotiorum complex (Rivera & Seifert

2011).

TABLE 2. Statistical support for the monophyletic clades representing Penicillium

mallochii and P. guanacastense in five gene data alignments. *

P. mallochii , P. guanacastense

ITS MP bootstrap (%) : 99 100

ML probability 0.998 1.00

BI posterior probability 1.00 1.00

COX1 MP bootstrap (%) 85 100

| ML probability 0.986 0.996

BI posterior probability 0.53 1.00

TEF1- MP bootstrap (%) 91 100

ML probability 0.975 1.00

BI posterior probability 0.87 1.00

BENA MP bootstrap (%) 100 100

ML probability 0.998 1.00

BI posterior probability 1.00 1.00

CMD MP bootstrap (%) 100 100

ML probability 1.00 1.00

BI posterior probability 1.00 1.00

*analyzed by Maximum Parsimony (MP), Maximum Likelihood (ML) and Bayesian Inference.

Both P mallochii and P. guanacastense received strong support as distinct

monophyletic clades in all analyses of all five genes (TABLE 2). The MP BENA

trees revealed two groups of strains within P. mallochii 86% and 70% bootstrap

support, which are absent in the other gene trees. These groups had 0.866 and

0.814 ML probabilities, and BI posterior probabilities of 0.74 and 0.66. Long

branch lengths subtending these two clades on the MP tree appear to be an

artifact, sometimes experienced with maximum parsimony analyses.

Penicillium mallochii & P. guanacastense spp. nov. (Costa Rica) ... 321

P. levitum

Tp hirayamae

P. adametzii

P. bilaiae

P levitum

Tp herquei

P. levitum P. levitum

Fic 1. Maximum likelihood (ML) gene trees for ITS, cox1, TEF1-a, BENA and cmp data sets.

The negative-log likelihood (-InL) of the ML trees are -1497.83287, -1610.84725, -3299.83157,

-2712.50962, -3454.53854. Branches with ML-bootstrap support >95%, MP-bootstrap support

>95%, and Bayesian Inference partition probabilities >0.99 are thickened.

322 ... Rivera & al.

Taxonomy

Penicillium mallochii K.G. Rivera, Urb & Seifert, sp. nov. FIGs 2, 4-8

MycoBaAnk MB 563043

Growth on CYA 29-39 mm, on MEA 24-35 mm diam. after 7 days. Conidiophores

monoverticillate, stipes 50-380 x 2-4 um, terminal vesicle 4-6 um wide, rarely with a

single metula 15-45 x 1.5-2 um. Phialides 7-10(16) x 2-3(4) um. Conidia (sub)globose,

2:9 SRD 25 wn:

Type: Costa Rica, Santa Rosa, Area de Conservacién Guanacaste, isolated from leaf of

Spondias mombin, 2003, leg. Joel Diaz, Felipe Chavarria-Diaz, and Maria Garcia no. 03-

RGTHC-903, isol. M. Urb (Holotype, DAOM 239917, dried culture on Blakeslee’s MEA;

ex-type culture, CCFC 239917).

Erymotoey: ‘mallochii’ named for Prof. David Malloch, retired mycologist from the

University of Toronto, now a visiting fellow at the Natural History Museum in St. John,

New Brunswick, an expert on ascomycetes, and a mentor to KGR, RGT and KAS.

Colonies on CYA after 7 days at 25°C: typically (24-)29-39 mm diam.; dense

and velutinous, sometimes with aerial mycelium in the center; moderately

sulcate, with 4-7 grooves, often wrinkled in the center; approximately 1 mm

deep; conidia Turquoise Grey to Greenish Grey (24-25D2-3), concentric

rings of different shades of these colors present; clear yellow exudate droplets

produced moderately in some strains, absent in others; vivid orange soluble

pigment present in some strains, absent in others; 2-3 mm of white mycelia at

the margin, margin entire; reverse deep Yellow (4A8), Reddish Orange (4A6-7)

or Golden Yellow to Orange (5-6A-B6-8) in the center, Light to Pastel Yellow

(3-4A4-5) at the margins.

Colonies on MEA after 7 days at 25°C: (19-)24-35 mm diam; planar,

velutinous, with no aerial mycelium; conidia Greyish Green to Dull Green

(25-30B-E3-7), sometimes forming crusts that dislodge when the colony is

disturbed, no exudates or soluble pigments present; reverse Greyish Green

(30C5), Greyish Yellow (1-2B-C3-8), Yellow (2-3A6-7), Deep Yellow (4A8)

or Orange (6A-B6-8), near the margins paler shades of these colors; margin

entire.

Conidiophores monoverticillate, borne from agar surface, stipes septate,

53-380 x 2-4 um, smooth to finely roughened; mostly unbranched, but in

some strains ~10% of the conidiophores have single branches 15-45 x 1.5-2

um; vesicles 4-6 um diam (mean for different strains = 4.5-6.0 + 0.71 um, n =

25). Phialides 7-10(-16) x 2-3(- 4) um, ampulliform with a collula. Conidia

globose to subglobose, finely roughened, 2.5-3.5 x 2-2.5 um (range of means

for different strains = 2.6-3.4 + 0.01 x 2.3-2.4 + 0.01, n = 25), mean L/W ratio

AWA oa §,

DistriBuTION: Area de Conservacién Guanacaste, Sector Santa Rosa, Guanacaste

Province, northwestern Costa Rica.

Habitat: Rothschildia lebeau and Citheronia lobesis (Saturniidae) caterpillar guts and

faeces, and leaves of Spondias mombin (Anacardiaceae).

ADDITIONAL MATERIAL EXAMINED: See TABLE 1.

Penicillium mallochii & P. guanacastense spp. nov. (Costa Rica) ... 323

—

2 DAOM239917 DAOM 229924 KAS 2611 KAS 3195

= — —

DAOM239912 DAOM239913 DAOM239912 DAOM239913

Fics 2-3. Seven day old cultures of Penicillium mallochii (2) and P. guanacastense (3) on CYA and

MEA, showing variation in colony characters.

324 ... Rivera & al.

Fics 4-8. Penicillium mallochii, photomicrographs from the holotype. 4-5 and 7-8. Monoverticillate

conidiophores. 6. Conidia. Scale bar = 10 um.

Penicillium guanacastense K.G. Rivera, Urb & Seifert, sp. nov. FIGS 2, 9-12

MycoBank MB 563044

Growth on CYA 25-33 mm, on MEA 29-33 mm diam. after 7 days. Conidiophore

monoverticillate or rarely with one metula, stipes 85-100 x 2.5-3 um, terminal vesicle

4-5.5 um wide. Phialides 8.5-12 x 2-3.5 um. Conidia globose, 2.5-3.0 um diam.

Penicillium mallochii & P. guanacastense spp. nov. (Costa Rica) ... 325

Type: Costa Rica, Santa Rosa, Area de Conservacién Guanacaste, isolated from Eutelia

sp. reared on leaf of Spondias mombin, 2002, leg. Joel Diaz, Felipe Chavarria-Diaz, and

Maria Garcia no. 02-RGTHC-462, isol. M. Urb (Holotype, DAOM 239912, dried culture

on Blakeslee’s MEA; ex-type culture, CCFC 239912).

ETYMOLOGY: ‘guanacastense’ named for the Area de Conservacién Guanacaste, where

the fungus was discovered.

Colonies on CYA after 7 days at 25°C: 25-33 diam, dense, velutinous, sulcate,

with 5-8 grooves, sometimes with sectors with poor sporulation; conidia

Greenish Grey (25-30B-E2), with yellow concentric ring near the margins

and a bluish grey concentric ring towards the center, clear exudate droplets

produced moderately and yellow soluble pigments produced by one strain; 2-3

Fics 9-12. Penicillium guanacastense, photomicrographs from the holotype. 9. Conidia. 10-12.

Monoverticillate conidiophores. Scale bar = 10 um.

326 ... Rivera & al.

mm of white mycelium at the margin, margin entire; reverse Orange Yellow to

Orange (4-6B7-8).

Colonies on MEA after 7 days at 25°C: 29-33 mm diam., planar, velutinous,

no aerial mycelia observed; conidia Greenish Grey (25-30B-E2), sometimes

forming crusts that dislodge when the colony is disturbed, with no exudates or

soluble pigments present; reverse Yellowish White (2-4B2), Orange (6B7-8) or

Light Brown (6-7D4-8), with paler shades of these colors near the margin.

Conidiophores monoverticillate on MEA, borne directly from agar surface;

stipes finely roughed, septate 85-100 x 2.5-3 um; unbranched or sometimes

up to ~10% have single branches 15-21 x 2.5-3 um; vesicles 4—5.5 um (mean

4.5-4.65 + 0.5 um). Phialides 8.5-12 x 2-3.5 um, ampulliform with

distinguishable collula. Conidia globose, finely roughened, 2.5-3 um diam

(range of means for different strains 2.7-2.8 + 0.01 x 2.5-2.6 + 0.01 um, n=25),

mean L/W ratio 1.1:1.

DIstRIBUTION: Area de Conservacién Guanacaste, Sector Santa Rosa, Guanacaste

Province, northwestern Costa Rica.

Habitat: Guts of Eutelia caterpillar (Noctuidae) feeding on foliage of Spondias mombin

(Anacardiaceae).

ADDITIONAL MATERIAL EXAMINED: See TABLE 1.

Discussion

Penicillium mallochii and P. guanacastense are phylogenetically related and

morphologically very similar to P sclerotiorum. All three species have vesiculate,

monoverticillate conidiophores and vivid orange to red colony reverses

on CYA. Morphological characters place both species in Penicillium subg.

Aspergilloides ser. Glabra (Pitt 1979), and the recently erected, phylogenetically

and morphologically defined sect. Sclerotiora (Houbraken & Samson 2011). All

species in this series have long, monoverticillate conidiophores with a swollen

or vesiculate apex. The conidiophores of P. sclerotiorum sensu stricto are

conspicuously vesiculate (mean 3.6 + 0.4 um), while those of P mallochii and

P. guanacastense are only moderately vesiculate, but with the vesicles of

P. mallochii (3.1 + 0.2 um) being slightly more swollen than those of

P. guanacastense (3.0 + 0.3 um). Furthermore, P. sclerotiorum has ellipsoidal

conidia, while P mallochii and P. guanacastense have globose to subglobose

conidia. Some strains of P sclerotiorum produce pale colored sclerotia (Pitt

1979), which were not observed in any of our P. mallochii and P. guanacastense

strains.

There are ecological differences among these three species. Penicillium

mallochii strains were isolated from saturniid caterpillars and leaves of their food

plant Spondias mombin, whereas P. guanacastense was isolated from noctuid

caterpillars, also feeding on Spondias mombin. In contrast, P. sclerotiorum

strains have mostly been isolated from the soil and the air.

Penicillium mallochii & P. guanacastense spp. nov. (Costa Rica) ... 327

Based on our five-gene analysis of multiple strains of each species, P. mallochii,

P. guanacastense, and P. sclerotiorum are phylogenetically distinct species. Each

species was resolved as a strongly supported monophyletic group with each

gene, with each kind of analysis, except for the ITS MP and BI analyses, which

did not resolve P. mallochii from the branch that grouped P. guanacastense and

P. sclerotiorum.

In the study of microfungi isolated from guts and frass of two species of

caterpillars feeding on the tree Spondias mombin, Penicillium was the fourth

most commonly isolated of the 48 identified genera, comprising 88 of 835

isolates (Urb 2006). Twenty-four strains of P. citrinum were isolated, and

this was the most frequent species isolated from guts of Citheronia lobesis

(with three $-tubulin variants, represented by GenBank accession numbers

JN637993-JN637995). A similar number of strains of P. mallochii were isolated

from Rothschildia lebeau (Urb 2006). Whether these species are phylloplane

inhabitants or endophytes of S. mombin leaves is unknown, and the significance

of their frequent association with these caterpillars remains a subject for future

research.

Acknowledgments

We are grateful for the critical reviews of the manuscript by R.A. Samson and S.W.

Peterson, to Rafik Assabgui for processing sequences, and the curators of the CBS, DAOM,

and NRRL culture collections for providing strains for this study. Walter Gams kindly

corrected our Latin diagnoses for an early draft. The fungal collection and isolations

were funded by a subcontract to RGT under the U.S. National Science Foundation grant

NSF DEB-0084224 to Anna-Louise Reysenbach (Portland State University) and NSF

DEB 0515699 to Daniel Janzen. The taxonomic studies were funded by Genome Canada

through the Ontario Genomics Institute, NSERC, and other sponsors listed at www.

BOLNET.ca.

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Gascuel O. 1997. BIONJ: An improved version of the NJ algorithm based on a simple model of

sequence data. Molecular Biology and Evolution 14: 685-695.

Glass NL, Donaldson GC. 1995. Development of primer sets designed for use with the PCR

to amplify conserved genes from filamentous ascomycetes. Applied and Environmental

Microbiology 61: 1323-1330.

Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. 2010. New algorithms and

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Hall TA. 1999. Bioedit: a user-friendly biological sequence alignment editor and analysis program

for Windows 95/98/NTT. Nucleic Acids Symposium Series. 41: 95-98.

328 ... Rivera & al.

Hong SB, Cho HS, Shin HD, Frisvad JC, Samson RA. 2006. Novel Neosartorya species isolated from

soil in Korea. International Journal of Systematic and Evolutionary Microbiology 56: 477-486.

http://dx.doi.org/10.1099/ijs.0.63980-0

Houbraken J, Samson RA. 2011. Phylogeny of Penicillium and the segregation of Trichocomaceae

into three families. Studies in Mycology 70: 1-51. http://dx.doi.org/10.3114/sim.2011.70.01

Huelsenbeck JP, Ronquist F. 2001. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics

17: 754-755. http://dx.doi.org/10.1093/bioinformatics/17.8.754

Katoh K, Asimenos G, Toh H. 2009. Multiple alignment of DNA sequences with MAFFT. Methods

in Molecular Biology 537: 39-64. http://dx.doi.org/10.1007/978-1-59745-251-9_3

Kornerup A, Wanscher JH. 1966. Methuen handbook of color, 3"' ed. Denmark, Sankt Jorgen

Tryk.

Peterson SW. 2000. Phylogenetic analysis of Penicillium species based on ITS and LSU-rDNA

nucleotide sequences. 163-178, in: Samson RA, Pitt JI. (eds). Integration of modern taxonomic

methods for Penicillium and Aspergillus classification. Amsterdam, Harwood Academic

Publishers.

Peterson SW, Bayer EM, Wicklow DT. 2004. Penicillium thiersii, Penicillium angulare and Penicillium

decaturense, new species isolated from wood-decay fungi in North America and multilocus

DNA sequence analysis. Mycologia 96: 1280-1293. http://dx.doi.org/10.2307/3762145

Pitt JI. 1979. The genus Penicillium and its teleomorphic states Eupenicillium and Talaromyces.

London, Academic Press.

Posada D, Crandall KA. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics

14: 817-818. http://dx.doi.org/10.1093/bioinformatics/14.9.817

Rivera KG, Seifert KA. 2011. A taxonomic and phylogenetic revision of the Penicillium sclerotiorum

complex. Studies in Mycology 70: 139-158. http://dx.doi.org/10.3114/sim.2011.70.03

Samson RA, Pitt JI. (eds). 1985. Advances in Penicillium and Aspergillus systematics. New York,

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Samson RA, Hoekstra ES, Frisvad JC. 2004. Introduction to food borne fungi, 7” ed. Centraalbureau

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Seifert KA, Samson RA, Waard JR de, Houbraken J, Levesque CA, Moncalvo JM, Louis-Seize

G, Hebert PDN. 2007. Prospects for fungus identification using CO1 DNA barcodes, with

Penicillium as a test case. Proceedings of the National Academy of Sciences of the United States

of America 104: 3901-3906. http://dx.doi.org/10.1073/pnas.0611691104

Swofford DL. 2002. PAUP”. Phylogenetic Analysis Using Parsimony (*and Other Methods), version

4, Sunderland, Massachusetts, Sinauer Associates.

Urb, M. 2006. Fungi associated with Costa Rican caterpillars. MSc thesis, University of Western

Ontario.

White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal

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methods and applications. San Diego, Academic Press.

MY COTAXON

http://dx.doi.org/10.5248/119.329

Volume 119, pp. 329-335 January-March 2012

Two new species of Terriera from Yunnan Province, China

JUAN-FANG SONG, LING LIU, YUAN-YUAN LI & CHENG-LIN Hou’

College of Life Science, Capital Normal University,

Xisanhuanbeilu 105, Haidian, Beijing 100048, China

* CORRESPONDENCE TO: houchenglincn@yahoo.com

ABSTRACT —Two new species of Terriera are described and illustrated from Yunnan Province,

China. Terriera rotundata on leaves of Quercus sp. is recognized by its more or less circular

ascomata in vertical section. Terriera petrakii on Smilax bracteata is similar to Lophodermium

camelliicola but differs from the latter by the pale conidiomata and ellipsoidal conidia and the

well-developed textura prismatica between the covering and the basal stroma.

Key worps —Ascomycota, Rhytismatales, Fagaceae, Smilacaceae

Introduction

The genus Terriera B. Erikss. was first segregated from Lophodermium

Chevall. based mainly on paraphyses swollen at the tip to form an

epithecium and the lack of lip cells (Eriksson 1970). Johnston (1988, under

L. multimatricum; 1989, under L. minus) considered ascomatal development

and structure as important features for Terriera. Molecular analyses support

Terriera as monophyletic (Ortiz-Garcia et al. 2003, Lantz et al. 2011). Many

Lophodermium species with ascomatal structures typical of Terriera have been

transferred to Terriera (Johnston 2001, Ortiz-Garcia et al. 2003). Of the 18

species recognized worldwide, only two species have been reported for China

(Frohlich & Hyde 2000, Yang et al. 2011). We describe here two new species of

Terriera from Yunnan Province, China.

Materials & methods

Hand sections of different thickness of ascomata were prepared using a razor

blade and mounted in water, Melzer’s reagent, 5% KOH, or 0.1% (w/v) cotton blue in

lactic acid. For the observation of ascomatal outlines in vertical section, sections were

pretreated in water before being mounted in lactic acid or cotton blue. Gelatinous

sheaths surrounding ascospores and paraphyses were observed in water or cotton blue.

Ascospore contents were drawn based on observations in water mounts. Measurements

330 ... Song & al.

were taken of 20 ascospores and asci for each specimen from tissues mounted in 5%

KOH or Melzer’s reagent.

Taxonomy

Terriera rotundata C.L. Hou, sp. nov. Fics. 1-5

MycoBANnk 561752

T. rotundata differs from the known species in Terriera by its more or less globular

ascomata in vertical section.

Type: China. Yunnan, Chuxiong, Zixishan, alt. ca. 2400 m, on fallen leaves of Quercus

sp. (Fagaceae). 2 August 2001, C.-L. Hou et al. 172 (Holotype BJTC 201102).

EryMo.oey: rotundata refers to the almost rounded shape of the ascoma as seen in

vertical section.

Ascomarta developing on fallen leaves, epiphyllous, occasionally hypophyllous,

not associated with bleached, pale brown areas. Ascomata (450—)600-900

x 330-450 um, elliptical but often strongly curved at maturity, occasionally

triangular, young ascomata more or less elliptical in outline, black, shiny, with

a blurred margin, both ends rounded and often with pale spots, the central part

of the ascomata strongly raising above the surface of the substrate, opening

by a longitudinal split or occasionally by teeth. Immature, unopened ascomata

macroscopically visible as two black areas separated from each other by a

broad, pale longitudinal zone. In median vertical section, ascomata 140-180

um deep in the host tissue, more or less circular in section. COVERING STROMA

consistently 28-35 um thick, composed of an outer layer of host cuticle,

epidermal and hypodermal cells filled with dark brown, thick-walled fungal

cells 2-3 um diam., and an inner layer formed by 3-4 rows of hyaline, thin-

walled, angular cells. Horizontal portion of the covering stroma 20-30 um

long, 10-25 um thick, composed of periphysoids arising from the inner part

of the covering stroma, their tips forming a hard black crust. A triangular

space in section between the covering stroma and the basal stroma is filled

with vertically arranged rows of prismatic, hyaline to slightly brown, thin-

walled cells 6-9 x 3-5 um. BASAL STROMA medium to well-developed, + flat,

composed of brown to dark brown textura globulosa-angularis, 15-20 um thick.

EXCIPULUM and Lip CELLS absent. SUBHYMENIUM 6-10 um thick, composed

of small cells 2-3 um diam. PARAPHySES 120-140 x 1 um, filiform, septate

or aseptate, unbranched, slightly swollen at the apex, embedded in gelatinous

sheaths. Asci ripening sequentially, 90-120 x 4—5.5 um, narrow-cylindrical,

thin-walled, J”, without circumapical thickening, discharging spores through a

small apical hole, 8-spored. Ascospores fasciculate, 70—90(-95) x 0.8-1 um,

filiform, hyaline, aseptate, with numerous guttules, gelatinous sheaths not seen.

Liberated ascospore often strongly bent.

Terriera petrakii & T. rotundata spp. nov. (China) ... 331

Figs 1-5. Terriera rotundata on leaves of Quercus sp. 1. A leaf bearing ascomata. 2. Ascomata

observed under the dissecting microscope (top right corner, two immature ascomata of Coccomyces

sp. 3. Ascoma in vertical section. 4. Structure of an ascoma in vertical section. 5. Paraphyses,

an ascus after the liberation of the ascospores, a mature ascus with ascospores, and discharged

ascospores.

332 ... Song & al.

ZONE LINES only present between colonies of this and taxonomically distinct

species.

CONIDIOMATA not observed.

ComMENTs — Terriera rotundata is distinctive within the genus because of its

more or less globular ascomata in vertical section. The developmental pattern in

T. rotundata is similar to that of T. minor (Tehon) P.R. Johnst. in vertical section

with the textura prismatica between covering and basal stroma composed of

vertically oriented cells (Johnston 1988, 1989, 2001). However, the shelf-

like ridge along ascomatal opening typical of Terriera is poorly developed in

T. rotundata. In addition, liberated ascospores of T’ rotundata are often strongly

bent and lack gelatinous sheaths.

Terriera petrakii C.L. Hou, sp. nov. FIGS. 7-13

MycoBANnk 561753

T. petrakii is similar to T. minor but differs by conidiomata and conidia.

Type: China, Yunnan, Chuxiong, Zixishan, alt. ca. 2400 m, on fallen leaves of Smilax

bracteata Presl (Smilacaceae), 2 August 2001, C.-L. Hou et al. 170 (Holotype, BJTC

201103).

Erymo.oey: In honour of Franz Petrak, a well-known mycologist who studied a similar

but undescribed species, “Lophodermium smilacinum” Petr.

AscoMaTA developing on leaves, epiphyllous, occasionally hypophyllous,

in slightly bleached pale brown areas, associated with thin, black zone lines.

Ascomata (600-)800-1300 x 380-450 um, elongate-elliptical, often strongly

curved or triangular, often coalesced, black in the central part, margin blurred,

shiny, both ends often with pale spots, the central part of the ascomata strongly

raising above the surface of the substrate, forming an elongate, flat zone at the

opening, lips absent, opening by a longitudinal split. Immature, unopened

ascomata macroscopically appearing as two black areas separated by a broad,

pale longitudinal zone, young ascomata more or less elliptical in outline. In

median vertical section, ascomata subepidermal, 120-150 um deep, + trapezoid

in section. COVERING STROMA 23-30 um thick over its entire extent, consisting

of an outer layer of host epidermal cells filled with dark brown, angular fungal

cells, a layer of dark brown textura angularis of different thickness and an

inner layer of hyaline, angular cells, horizontal portion of the covering stroma

80-100 um long and 15-25 um thick, composed of black textura angularis with

cells 4-6 um diam. The triangular zone in section between the covering stroma

and the basal stroma composed of thin-walled, cylindrical cells arranged in

vertical rows. Basal stroma, medium to well-developed, + flat, composed of

brown to dark brown textura globulosa-angularis, 10-15 um thick. ExcIpULUM

present, formed by paraphysis-like cells 1-2 um diam., thickened near the

Terriera petrakii & T. rotundata spp. nov. (China) ... 333

SIS RES

OP REOS TOES

8 x

XKBOUOSO

50um

wings

mae

Fics 1-7. Terriera petrakii on leaves of Smilax bracteata. 6. A leaf bearing ascomata, conidiomata

and zone lines. 7. Ascomata observed under the dissecting microscope (right, an ascoma of

Coccomyces occultus). 8. A conidioma observed under the dissecting microscope 9. Ascoma in

vertical section. 10. Structure of an ascoma in vertical section. 11. Structure of a conidioma in

vertical section. 12. Conidiogenous cells and conidia. 13. Paraphyses, a young ascus, two mature

asci with ascospores, and discharged ascospores.

334 ... Song & al.

opening, embedded in a gelatinous matrix. SUBHYMENIUM consisting of textura

angularis with cells 2-4 um diam., 6-10 um thick. PARAPHysES 100-125 x 1

um, filiform, septate, branched, or unbranched, not swollen at the apex. Ascr

ripening sequentially, 85-110 x 4—5 um, cylindrical, thin-walled, J—, without

circumapical thickening, discharging spores through a small apical hole,

8-spored. Ascosporss fasciculate, (60-)70-85 x 0.8 um, filiform, hyaline,

aseptate, with a thin gelatinous sheath.

CONIDIOMATA in pale brown areas near the ascomata, round, pale, with

conspicuous black perimeter line, 140-300 um diam., opening by ostioles. In

vertical section, conidiomata intraepidermal, 40-55 um deep. UPPER LAYER

composed of host cuticle, remains of host epidermal cells, and hyaline textura

angularis with cells 3-5 um diam. BASAL LAYER 50-58 um thick, composed of

light brown textura angularis. CONIDIOGENOUS CELLS ovoid-cylindrical, 9-13

x 1.5-2 um tapering towards the apex, hyaline. Conrp1a broadly ellipsoidal,

hyaline, 1.5-2 x 1 um.

ADDITIONAL SPECIMEN EXAMINED —UNITED STATES, Hawaii: Olinda Pipe Line, on

fallen twigs of Smilax sp., coll. C.L. Shear & N.E. Stevens, det. F Petrak, 29 December

1928 (W 618).

CoMMENTS — The ascomatal structure of Terriera petrakii is very similar to that

of T. minor, which differs in shorter, uncurved ascomata, broader 0-1-septate

ascospores, a less well developed excipulum, and an unknown anamorph.

The conidioma shape and structure and ascospore shape in T. petrakii are

very similar to those of Lophodermium camelliicola Minter (which should

be transferred to Terriera). That species differs in producing black instead of

pale conidiomata, cylindrical to ellipsoidal conidia, a less developed textura

prismatica between the covering and basal stroma, and its habit on Camellia

sinensis (L.) Kuntze. The herbarium specimen W 618 (labelled by Petrak as

“Lophodermium smilacinum Petr. n. sp. but never described), with a structure

typical of Terriera, is somewhat similar to T: petrakii but occurs on twigs, has

narrower and more confluent ascomata, and lacks an anamorph.

Ascomata of Terriera petrakii often occur together with Hypoderma

smilacicola C.L. Hou & M. Piepenbr. and Coccomyces occultus Y.L. Lin & Z.Z.

Li (Hou & Piepenbring 2006, Lin et al. 1999).

Acknowledgments

We are grateful to Dr. PF. Cannon and Dr. R Kirschner for serving as pre-submission

reviewers and to the curators and staff of the herbarium W for specimen loans. The

study was supported by the Federal Ministry of Education and Research of Germany,

the National Natural Science Foundation of China (31170019, 30770006), and PHR

(KZ201110028036).

Terriera petrakii & T. rotundata spp. nov. (China) ... 335

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19: 1-71.

Frohlich J, Hyde KD. 2000. Palm microfungi. Fungal Diversity Press. Hong Kong. 364 p.

Hou CL, Piepenbring M. 2006. Five new species of Hypoderma with a key to Hypoderma species for

China. Nova Hedwigia 82: 91-104. http://dx.doi.org/10.1127/0029-5035/2006/0082-0091

Johnston PR. 1988. An undescribed pattern of ascocarp development in some non-coniferous

Lophodermium species. Mycotaxon 31: 383-394.

Johnston PR. 1989. Lophodermium (Rhytismataceae) on Clusia. Sydowia 41: 170-179.

Johnston PR. 2001. Monograph of the monocotyledon-inhabiting species of Lophodermium.

Mycol. Pap. 176: 1-239.

Lantz H, Johnston PR, Park D, Minter DW. 2011. Molecular phylogeny reveals a core clade of

Rhytismatales. Mycologia 103: 57-74. http://dx.doi.org/10.3852/10-060

Lin YR, Li ZZ, Chen Y, Li Z, Wu WJ. 1999. A new species of Rhytismatales, Coccomyces occultus sp.

nov. J. Anhui Agricult. Univer. 26: 37-39.

Ortiz-Garcia $, Gernandt DS, Stone JK, Johnston PR, Chapela IH, Salas-Lizana R, Alvarez-Buylla

ER 2003. Phylogenetics of Lophodermium from pines. Mycologia 95:846-859.

http://dx.doi.org/10.2307/3762013.

Yang ZZ, Lin YR, Hou CL. 2011. A new species of Terriera (Rhytismatales, Ascomycota) from

China. Mycotaxon 117: 367-371. http://dx.doi.org/10.5248/117.367

MY COTAXON

http://dx.doi.org/10.5248/119.337

Volume 119, pp. 337-343 January-March 2012

Gloeocystidiellum kenyense in Azores and Madeira

M. TERESA TELLERIA’, MARGARITA DUENAS’, IRENEIA MELO?,

ESPERANZA BELTRAN-TEJERA?, J. LAURA RODRIGUEZ-ARMAS',

ISABEL SALCEDO‘ & MARIA P. MARTIN?

‘Real Jardin Botdnico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain

2Jardim Botdnico (MNHN), Universidade de Lisboa/CBA-FCUL,

Rua da Escola Politécnica 58, 1250-102 Lisboa, Portugal

°Dpto. de Biologia Vegetal (Botanica), Universidad de La Laguna, 38071 La Laguna, Tenerife, Spain

‘Dpto. de Biologia Vegetal y Ecologia (Botanica), Universidad del Pais Vasco,

Aptdo. 644, 48080 Bilbao, Spain

* CORRESPONDENCE TO: felleria@rjb.csic.es

ABSTRACT — The aim of this study was to clarify the identity of several collections of

Gloeocystidiellum from Azores Archipelago and Madeira Island that were initially related to

the Gloeocystidiellum porosum-clavuligerum group. Three new ITS nrDNA sequences were

aligned and analysed phylogenetically with 18 homologous sequences from GenBank. The

two main lineages, with very high bootstrap value, that were detected correspond to two well-

separated species, G. kenyense and G. clavuligerum. The sequences obtained from Azores and

Madeira specimens clustered in the G. kenyense clade. Gloeocystidiellum kenyense is described

and illustrated from the specimens collected in Macaronesian region, and its affinities with

G. clavuligerum are discussed.

Key worps — Basidiomycota, Gloeocystidiellaceae, corticioid fungi, oceanic islands,

taxonomy

Introduction

In the framework of our studies about corticioid fungi from oceanic islands

(Duenias et al. 2008a, b, Melo et al. 2008, Telleria et al. 2008, 2009a, b, c), we

found several collections of a Gloeocystidiellum species. These collections from

Azores Archipelago and Madeira Island are related, according to Larsson &

Hallenberg (2001), to the Gloeocystidiellum porosum-clavuligerum group.

Donk (1931) described Gloeocystidiellum (Russulales, Gloeocystidiellaceae),

selecting G. porosum (Berk. & M.A. Curtis) Donk as type, to accommodate the

corticioid fungi with monomitic hyphal system, gloeocystidia, and smooth or

ornamented amyloid spores. Larsson & Larsson (2003) accept Gloeocystidiellum

338 ... Telleria & al.

in a strict sense to include a few species with resupinate basidiomes with

smooth hymenophores, monomitic hyphal systems, hyphae with clamps, SA+

or SA- gloeocystidia, and ellipsoid, ornamented and amyloid spores.

Several morphological and molecular studies as well as cultural studies

and intercompatibility tests have analyzed species delimitation in the

Gloeocystidiellum porosum-clavuligerum group (Nakasone 1982, Hallenberg

1984, 1988, Boidin et al. 1997, Larsson & Hallenberg 2001, Larsson & Larsson

2002, 2003). ITS2 and LSU nrDNA phylogenetic analyses were congruent with

intercompatibility tests and gave higher resolution in differentiating this species

complex than could be found in morphologically based analyses. Larsson &

Larsson (2002) and Larsson & Hallenberg (2001) found that G. porosum and

G. clavuligerum (Hohn. & Litsch.) Nakasone are phylogenetically widely separated

and that G. clavuligerum should be divided into two taxa, G. clavuligerum and

G. kenyense.

Our goal was to clarify the identity of Gloeocystidiellum specimens from

Azores and Madeira mentioned above. ITS nrDNA sequences from three

collections were compared with homologous sequences from GenBank

published in Larsson & Hallenberg (2001) and Larsson & Larsson (2003). As

mentioned in Seifert (2009), the ITS region has became the default marker for

species identification in fungi. Morphological characters were compared with

the taxa described in Larsson & Larsson (2002).

Materials & methods

Morphological studies

Colors of basidioma are according to ISCC_NBS Centroid Color Charts. Dried

specimens were used for light microscope study. Measurements and drawing were

made from microscopic sections mounted in 3% KOH solution or Melzer’s reagent and

examined at up to 1250x with an Olympus BX51 microscope provided with a drawing

tube. Lengths and widths of 30 spores, 10 basidia, and 10 cystidia were measured from

each sample. Voucher specimens have been deposited in TFC, MA, LISU, O.

DNA extractions and sequencing

Genomic DNA was isolated from herbarium specimens using an EZNA Fungal

DNA MiniPrep kit (Omega Biotek, Doraville, Georgia, USA) or a DNeasy Plant Mini

Kit (Qiagen, Hilden, Germany) following instructions of the manufacturers. Lysis

buffer incubation was done overnight at 55°C. The primer pair ITSIF and ITS4 was

used to obtain amplifications of both ITS regions, including the 5.88 of the ribosomal

RNA gene cluster and small flanking parts of the SSU and LSU genes (White et al.

1990, Gardes & Bruns 1993). Amplifications were done using illustra“ PuReTaq™

Ready-To-Go™ PCR beads (GE Healthcare, Buckinghamshire, UK) as described in

Winka et al. (1998) following thermal cycling conditions in Martin & Winka (2000).

Negative controls lacking fungal DNA were run for each experiment to check for

contamination of reagents. Results of amplifications were assayed from 5 ul aliquots

Gloeocystidiellum kenyense ... 339

by gel electrophoresis of 2% Pronadisa D-1 Agarose (Lab. Conda, Spain). When the

primer pair ITS1F/ITS4 failed, the ITS regions were amplified in two parts as mentioned

in Telleria et al. (2010). Prior to sequencing the amplification products were cleaned

using a QIAquick gel PCR purification kit (QIAGEN, Valencia, California, USA). Both

strands were sequenced separately using primers mentioned above at Secugen S.L.

(Madrid, Spain). When products were faintly visualized on agarose gels (less that 20 ng/

ul), cloning was conducted with pGEM?’-T Easy Vector System II cloning kit (Promega,

Madison, Wisconsin, USA). From each cloning reaction, three clones were selected for

sequencing, using vector specific primers T7 and SP6 at Macrogen (Korea). Sequences

were edited and assembled using Sequencher™ version 4.2 (Gene Codes Corporations,

Ann Arbor, Michigan, USA). BLAasTN searches with MEGABLAST option were used

to compare the sequences obtained against the sequences in the National Center of

Biotechnology Information (NCBI) nucleotide database. The consensus sequences have

been lodged in the EMBL-EBI database.

Sequences were manually aligned using Se-Al v2.0al1 Carbon (Rambaut 2002) for

multiple sequences and compared with homologous sequences of Gloeocystidiellum spp.

retrieved from the EMBL Nucleotide Sequence Databases. The alignment was analyzed

using the program PAUP* Version 4.0b10 for Macintosh (Swofford 2003). The maximum

parsimony analysis (MP) was inferred using the heuristic search option in PAUP. Gaps

were treated as missing data. Branch lengths equal to zero were collapsed to polytomies.

Nonparametric bootstrap (BP) support (Felsenstein 1985) for each clade was tested

based on 10,000 replicates, using the fast-step option.

Results & discussion

From direct PCR with pair primer ITS1F/ITS4, only collections TFC 15278

(GenBank FR878082), TFC 15309 (GenBank FR878083) and MA-Fungi 80408

(GenBank FR878084) were amplified and sequenced directly.

The new sequences included the last 5 bp of 18S nrDNA, the complete

ITS1—5.8S—ITS2 sequence, and the first 46 or 50 bp of the 28S nrDNA.

The BLAST search of the three new direct sequences showed an e-value

of 0.0 and a maximum similarity of 99.0% with G. clavuligerum sequences

(AF310080, AF310077, AF310079, AF310084, AF310082; AF310083) and

G. purpureum Sheng H. Wu (AF441338). Those GenBank sequences lacked the

ITS1 region and the first part of 5.88 nrDNA, thus restricting the BLAsT search

to the last 369 bp at the 3’ end.

The other G. clavuligerum sequences from Larsson & Hallenberg (2001) and

Larsson & Larsson (2003) as well as the two outgroup G. porosum sequences

(AF310101 and AF31009) were included in full length in the alignment,

which contained 21 sequences and 388 positions, of which 293 were constant,

13 parsimony-uninformative, and 82 parsimony-informative. The 100 most

parsimonious trees were obtained using the heuristic search option (tree length

= 114; CI = 0.9211; HI = 0.0789, RI = 0.9726). The bootstrap 50% majority-rule

consensus tree is shown in Fic. 1.

340 ... Telleria & al.

FR878082, TFC 15278, Azores, Portugal

FR878083, TFC 15309, Azores, Portugal

FR878084, MA-Fungi 80408, Madeira, Portugal

AF310080, FCUG 1039, NH8150, Romania

AF310079, FCUG 2238, NH11471, Turkey

100 AF441338, Wu9310-45, Taiwan

AF310082, FCUG 2567, NH12810, Canada

AF310077, FCUG 677, NH7060, Canada

AF310084, JS16976, Norway

G. kenyense

AF310083, FCUG 2731, NH13159, Russia

AF310078, FCUG 676, NH7058, Canada

AF310081, FCUG 2248, NH1687, France

67 AF310085

AF310087

AF310088

100 100 AF310086 ,

714 G. clavuligerum

AY048875

AY048877

AY048878

AF310104

arstoos7 | G- porosum

FIGURE. 1. Strict consensus tree of 100 ITS trees obtained from heuristic search in the parsimony

analyses for 21 Gloeocystidiellum collections. Bootstrap support values (10,000) above 50% are

indicated above branches. The three new sequences generated in this study are labeled with the

accession number in bold.

Using G. porosum as outgroup, two main clades were detected, both with

very high bootstrap values (100%). According to Larsson & Hallenberg

(2001) and Larsson & Larsson (2002, 2003), the clades correspond with two

well-distinguished species, G. kenyense and G. clavuligerum. As shown in

Fic. 1, the three Azores and Madeira sequences cluster in the G. kenyense clade.

Comparison of our collections with the G. kenyense holotype (L. Ryvarden

8996, herb. O) and protologue description (Hjortstam 1987) revealed no

morphological differences.

Gloeocystidiellum kenyense Hjortstam, Mycotaxon 28: 29. 1987 FIG. 2

The Azores and Madeira specimens have resupinate basidiomata that are

closely adnate, ceraceous, thick and hard with a smooth cream to ochraceous

(89. p. Y-70.LOY) hymenophore that is cracked when dried and an

undifferentiated margin. The hyphal system is monomitic with densely

interwoven, richly branched, thin-walled hyphae and clamped at all septa.

Gloeocystidia are variable, originate from all levels, 35-85 x 7-16 um, often

with moniliform apical appendices, SA-. Basidia are cylindrical, (13-)21-27 x

4-5 um, with four sterigmata. Basidiospores are subglobose to ellipsoid, 3.5-5

x 3-3.5 um [L = 4.1, W = 3, L/W = 1.36], amyloid, ornamented.

MATERIAL EXAMINED: PORTUGAL. Azores: Pico IsLANb, S. Roque, Mistério

da Prainha, 38°29'42"N 28°16'01"W, 380 m a.s.l., on Pittosporum undulatum Vent.

Gloeocystidiellum kenyense ... 341

Darah A Ob Or

Leave GN eS) uy

ees

‘\ (

iif

100 ym| | !

1 i

ED =

Ser See ee et

FIGURE. 2. Gloeocystidiellum kenyense (LISU 210747). a: section through basidiome; b: basal

gloeocystidia; c: hyphae; d: hymenial layer with basidia and gloeocystidia; e, f: basidiospores.

(Pittosporaceae), 25 Febr. 2005, E. Beltran-Tejera & J.L. Rodriguez-Armas (TFC 15278,

GenBank FR878082); S. Roque, Meia Encosta de Santa Luzia, Travessa da Cima,

38°31'13"N 28°22'14"W, 483 m a.s.l, on Pittosporum undulatum, 25 Febr. 2005, E.

Beltran-Tejera & J.L. Rodriguez-Armas (TFC 15309, GenBank FR878083). MADEIRA:

Porto Moniz, Seixal, Chao da Ribeira, 32°47'56"N 17°06'52"W, 485 m a.s.l., on Ocotea

342 ... Telleria & al.

foetens (Aiton) Benth. & Hook.f. (Lauraceae), 27 Jun. 2000, M.T. Telleria, 14140Tell. (MA-

Fungi 80408, GenBank FR878084); 428 m a.s.l., on unidentified wood, 01 May 1998, I.

Melo, 7572IM (LISU 210747). KENYA. EASTERN PRov.: Aberdare Mounts. Kimakia

Forest sta., 0°45'S 36°50'W), alt. c. 2200-2400 m a.s.l., 16-18 Jan. 1973, L. Ryvarden

8996 (holotype, O). TANZANIA. ArusHaA PRov.: Arusha Nat. Park, Mountain Meru

east slope, 1800-2300 m a.s.1., 08 Febr. 1973, L. Ryvarden 10027 (O). REUNION. Cilaos,

28 March 1990, G. Gilles, Boidin 14289 (O).

According to Larsson & Larsson (2002), G. kenyense and G. clavuligerum differ

in their spore sizes, as well as in gloeocystidial contents, which are SA- and

SA+ respectively.

Gloeocystidiellum kenyense is widely distributed and known from Europe,

North America, Africa, and Taiwan on decaying angiosperm wood (Larsson

& Larsson 2002). In the Azores Archipelago it was found on Pittosporum

undulatum, an introduced evergreen tree native to SE Australia that today

dominates most of the low altitude forests of the Azores archipelago. On

Madeira Island the species was found on Ocotea foetens, an indigenous species

characteristic of the laurisilva, the Macaronesian subtropical mountain moist

forest known from Madeira and Canary Islands.

Acknowledgments

Thanks to Dr Ellen Larsson for sending us a copy of her Ph-Dissertation and

information related to G. kenyense and to the curator of Mycological Herbarium of

University of Oslo (O) for his help. We are also grateful to Nils Hallenberg and Karl-

Henrik Larsson for comments and corrections to improve the manuscript. This study

was supported by the project CGL2009-07231.

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MY COTAXON

http://dx.doi.org/10.5248/119.345

Volume 119, pp. 345-350 January-March 2012

A new species of Skeletocutis (Polyporaceae) on bamboo

in tropical China

Li-WEI ZHOU & WEN-MIN QIN

State Key Laboratory of Forest and Soil Ecology, Institute of Applied Ecology,

Chinese Academy of Sciences, Shenyang 110164, P. R. China

* CORRESPONDENCE TO: liwei_zhou1982@163.com

AxBsTRACT — Skeletocutis bambusicola sp. nov. is described and illustrated from the

tropical region of Yunnan, southern China. The species is characterized by the annual,

resupinate cream-colored basidiocarps, small pores, a dimitic hyphal system, small ellipsoid

basidiospores, and habit on dead bamboo. The combination of characters distinguishes

S. bambusicola from other known species of Skeletocutis.

Key worps — Polyporales, polypore, taxonomy

Introduction

Encrustation of hyphae at the dissepiment edges of polypores is a special and

steady morphological character in taxonomy. Species with encrusted hyphae

have been referred to three genera: Tyromyces P. Karst., Piloporia Niemela,

and Skeletocutis Kotl. & Pouzar (Niemela 1998). Niemela (1998) differentiated

Skeletocutis from the other two genera by its strong tendency to form effused

basidiocarps with a homogeneous to duplex context and hyaline hyphae.

Niemela (1998) reviewed the collections of resupinate Skeletocutis species

from East Finland and North Sweden, while Dai (1998) provided a key to all

accepted Skeletocutis species from China. More recently, Nufiez & Ryvarden

(2001) delimit Skeletocutis based on its dimitic system and encrustation of the

hyphae at dissepiment edges.

Phylogenetically, Skeletocutis is closely related to Tyromyces (Kim et al. 2001)

and clusters within the Polyporales (Miettinen & Larsson 2011).

Two species, Skeletocutis lenis (P. Karst.) Niemela and S. vulgaris (Fr.)

Niemela & Y.C. Dai, were transferred to a new genus, Sidera Miettinen & K.H.

Larss. (Hymenochaetales) based on ITS (internal transcribed spacer) and LSU

(large subunit) nuclear ribosomal DNA (nrDNA) sequence analyses (Miettinen

& Larsson 2011). Although representing different orders, Skeletocutis and

346 ... Zhou & Qin

Sidera share many morphological characters, including whitish resupinate

basidiocarps (in many species) with small pores, allantoid basidiospores,

and narrow skeletal hyphae (Miettinen & Larsson 2011). The main difference

between the two genera is that Skeletocutis usually has hyphae encrusted by fine

crystals on the tube mouths, whereas in Sidera the dissepiment edge hyphae are

smooth or covered with a few faceted crystal clusters (Niemela 1998, Miettinen

& Larsson 2011). Therefore species with faceted, rather than fine, crystals at the

dissepiment edges should be excluded from Skeletocutis. Moreover, molecular

data do not support the concept of Skeletocutis as restricted to dimitic hyphal

structure but one that includes species with a monomitic hyphal system

(Miettinen & Larsson 2011).

Extensive studies on Chinese wood-decaying fungi have recorded 704

polypores and 505 corticioid or hydnoid fungi for China (Dai 2011, 2012a),

among which many new taxa have been described from tropical China (Cui

& Dai 2008, Cui et al. 2009, 2010, Yuan & Dai 2009, Dai & Cui 2011, Dai & Li

2010, Dai 2012b, Dai et al. 2009, 2010, 2011, Zhou & Jia 2010). Nonetheless,

the diversity and richness of tropical wood-decaying fungi are still not well

known. During a study on polypores from tropical Yunnan, an unknown

species with finely encrusted hyphae growing on dead bamboo was identified.

It is described and illustrated here as Skeletocutis bambusicola following the

generic concepts of Niemela (1998) and Nufiez & Ryvarden (2001) according

to its latest delimitation (Miettinen & Larsson 2011). This represents the 22"

Skeletocutis species recorded in China (Dai 2012a).

Materials & methods

The studied specimens were deposited at the herbarium of the Institute of Applied

Ecology, Chinese Academy of Sciences (IFP, Shenyang, China). The microscopic

procedure follows Dai (2010). Sections were studied at magnifications up to x1000

using a Nikon Eclipse 80i microscope with phase contrast illumination. The spore size

variation excludes 5% of measurements (given in parentheses) from each end of the

range. Abbreviations used are: IKI = Melzer’s reagent, IKI- = negative in Melzer’s, KOH

= 5% potassium hydroxide, CB = Cotton Blue, CB- = acyanophilous, L = mean spore

length (average ofall spores), W = mean spore width (average of all spores), Q = variation

in L/W ratios among all specimens studied, and n = number of spores measured from

given number of specimens. Drawings were made with the aid of a drawing tube. Special

color terms follow Petersen (1996).

Taxonomy

Skeletocutis bambusicola L.W. Zhou & W.M. Qin, sp. nov. FIG. 1

MycoBank MB 563066

Differs from Skeletocutis alutacea, S. borealis, S. krawtzewii, and S. percandida by its

habit on dead bamboo.

Skeletocutis bambusicola sp. nov. (China) ... 347

20)

’ \

Wer

Y oS

ray \

\\)

Mes } hands

BEAN \W/

FiGurE 1. Skeletocutis bambusicola (holotype).

a: Spores. b: Basidia and basidioles. c: Cystidioles. d: Hyphae from subiculum.

e: Hyphae from trama. f: Hyphae at dissepiment edges.

348 ... Zhou & Qin

Type: China, Yunnan Province, Xishuangbanna, Menglun County, on dead bamboo,

12.1X.2007, Yuan 3493 (Holotype IFP 015814).

ErymMo.ocy: bambusicola (Lat.): refers to growth on bamboo.

FRurTBopy Basidiocarps annual, resupinate, soft when fresh, becoming corky

upon drying, < 13 cm long, 3.5 cm wide, 1 mm thick at center. Pore surface

cream when fresh, buff when dry, more or less glancing; pores round to angular,

8-11 per mm; dissepiments thin, entire to slightly lacerate. Subiculum cream

and soft corky when dry, < 0.2 mm thick; tubes buff-yellow, paler than pore

surface, < 0.8 mm long.

HYPHAL STRUCTURE Hyphal system dimitic; generative hyphae mostly

bearing clamp connections, some at dissepiment edges with simple septa; all

hyphae IKI-, CB-, unchanged in KOH.

SUBICULUM Hyphal structure interwoven. Generative hyphae infrequent,

hyaline, thin-walled, encrusted by fine crystals, occasionally branched, winding,

1-4 um diam; skeletal hyphae dominant, hyaline, thick-walled to distinct

thick-walled with a narrow lumen, unbranched, strongly winding, occasionally

bearing clamp connections, 1.5-3 um diam.

TuBEs Hyphal structure interwoven. Generative hyphae infrequent, hyaline,

thin-walled, encrusted by fine crystals, occasionally branched, 1.5-3 um in diam;

skeletal hyphae dominant, hyaline, slightly thick-walled, occasionally branched,

strongly winding, occasionally bearing clamp connections, 2-3 um diam.

Dissepiment edge with both generative and skeletal hyphae; generative hyphae

mostly covered by fine, sharp-pointed encrustations. Hyphal pegs occasionally

present. Cystidia absent, fusoid cystidioles present in the hymenium, hyaline,

thin-walled, 10-17 x 3-5 um. Basidia clavate, with four sterigmata and a basal

with clamp connections, 12-20 x 4-5 um; basidioles in shape similar to basidia,

but slightly smaller.

Basipiospores Ellipsoid, hyaline, thin-walled, smooth, IKI-, CB-, (2.6-)

2.7-3.1(-3.2) x 1.5-1.9(-2.0) um, L = 2.92 um, W = 1.74 um, Q = 1.66-1.70

(n = 60/2).

TYPE OF ROT White rot.

ADDITIONAL SPECIMEN EXAMINED: CHINA, YUNNAN PROVINCE, Xishuangbanna,

Menglun County, on dead bamboo, 12.1X.2007, Yuan 3500 (IFP 015815).

REMARKS — Skeletocutis bambusicola, collected in tropical China, produces

annual resupinate cream basidiocarps, finely encrusted hyphae not restricted

to the dissepiment edges, and small pores and basidiospores. This combination

of characters is typical for Skeletocutis.

The new species belongs to the thick-spored Skeletocutis group, which

includes five other species — S. alutacea (J. Lowe) Jean Keller, S. borealis

Niemela, S. krawtzewii (Pilat) Kotl. & Pouzar, S. percandida (Malencon &

Bertault) Jean Keller, and S. perennis Ryvarden (Niemela 1998). Skeletocutis

Skeletocutis bambusicola sp. nov. (China) ... 349

TABLE 1. Comparison of six thick-spored Skeletocutis species based on Niemela (1998)

and specimens examined in the present study.

Gua oene BASIDIOCARP PorESs/MM DISSEPIMENT BASIDIOSPORES

EDGE [Lx W]

S. alutacea pena pie 9-10 Dimitic O eee

se aie ge tse ie ered ay Mamie o 2 re i

Sbowalis Peennish (636-768) Monomite 40 "5

Skcrawigewis aL ee ‘ ee ak 2 é 20 <=

S. percandida ie cal i 5-6 Dimitic 5 : ae

Speermis Fern 5.6 imi ea

borealis and S. percandida are distinguished from the new species by their

cylindrical basidiospores (Niemela 1998). The other three similar species

also have ellipsoid basidiospores, S. alutacea is distinguished by its longer

basidiospores (L = 2.9-3.8 um, Niemela 1998) and rhizomorphic basidiocarps

while S. krawtzewii and S. perennis differ in larger basidiospores (3.4-4.1 x

1.7-2 in S. krawtzewii; 2.9-3.6 x 1.8-2.1 in S. perennis) and larger pores (5-6

per mm; Niemela 1998). In addition, S. krawtzewii has monomitic hyphae

at dissepiment edge and thin-walled skeletal hyphae (Niemela 1998) and

S. perennis has perennial basidiocarps with subparallel hyphae in the subiculum

(Niemela 1998). The main morphological differences among the six species are

compared in TABLE 1.

Niemela (1998) proposed that, “Ecological characteristics are important

when identifying the species of Skeletocutis.” Skeletocutis bambusicola was

found on dead bamboo, which might be the most important evidence that

separates it from other species of Skeletocutis.

Acknowledgements

We thank Dr. Hai-Jiao Li (BJFC, China) for preparing the microscopic structure

and express our gratitude to Drs. Bao-Kai Cui (BJFC, China) and Deepika Kumari

(Directorate of Mushroom Research, India) who reviewed the manuscript. The research

was financed by the National Natural Science Foundation of China (Project No.

30910103907).

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MY COTAXON

http://dx.doi.org/10.5248/119.351

Volume 119, pp. 351-359 January-March 2012

Paxillus albidulus, P. ammoniavirescens, and P. validus revisited

ELSE C. VELLINGA’ , ERIN P. BLANCHARD’,

STEPHEN KELLY’? & MARCO CONTU?

"Department of Plant and Microbial Biology, University of California,

111 Koshland Hall #3102, Berkeley CA 94720-3102, U.S.A.

71 Church Hill, Bedmond, Abbotts Langley, Hertfordshire. WD5 ORW, U.K.

°Via Marmilla, 12, I-07026 Olbia (OT), Italy

*CORRESPONDENCE TO: ecvellinga@comcast.net

ABSTRACT—Comparison of nrITS sequences of the type specimens of Paxillus ammonia-

virescens and P. validus show them to be identical. The name Paxillus ammoniavirescens was

published in ‘Spring 1999’ [northern hemisphere] and that of Paxillus validus in August 1999,

making P ammoniavirescens the correct name for this species. A variant without pigments

fits into P. involutus, suggesting that P albidulus might be an albino variant of any species

in the P involutus complex. Paxillus ammoniavirescens is widespread in Europe and has

been introduced in the southern hemisphere; P obscurosporus is known from China and

Europe; and the two taxa in the P involutus clade are reported from Europe and from North

America.

Keyworps—Paxillaceae, phylogenetic species, pigment-free variants

Introduction

The genus Paxillus Fr. is widely distributed in the northern hemisphere,

forms ectomycorrhiza with a range of host trees, is extensively used in

ectomycorrhizal research (e.g. Ek 1997, Leake et al. 2001, Prendergast- Miller

et al. 2011, Wilkinson et al. 2010), and is a nightmare for species recognition.

There are two main species clusters — P. rubicundulus P.D. Orton and allies,

associating with Alnus species, and the P. involutus complex, associating with a

range of host trees (Pinaceae, Fagaceae, Salicaceae, Corylaceae, Tiliaceae, etc.).

Hedh et al. (2008), who recently addressed species concepts of the European

taxa in the P involutus complex, recognized four species based on five gene

genealogies and nrITS sequence analyses. Three of the four phylogenetic

species correspond with the intercompatibility groups identified by Fries

(1985). Hedh et al. (2008) recognized Paxillus involutus (Batsch) Fr., P. validus,

P. obscurosporus C. Hahn, and a fourth species, sister to P involutus, which they

refrained from naming.

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Three Paxillus species revisited ... 353

Two other species have been described from Europe, P. albidulus Sutara

(Sutara 1992), a predominantly white species, and P ammoniavirescens (Dessi

& Contu 1999), characterized by a strong green reaction of the pileus surface

with ammonia. As these species were not included in the study by Hedh et

al. (2008), their identity and relationship to the phylogenetic species remained

unresolved.

Here, we investigate the identity of P ammoniavirescens and the phylogenetic

placement of taxa without pigments using nrITS sequence data and phylogenetic

methods.

Material & methods

MaTERIAL—Type material of P. ammoniavirescens, additional collections of P. ammonia-

virescens from the type locality, various Paxillus collections from the UK and France,

and Paxillus collections growing with introduced Betula in western North America were

used for analyses (TABLE I).

DNA EXTRACTION TO SEQUENCING—DNA was extracted from dried material using a

Qiagen DNeasy® Blood and Tissue kit (Qiagen, Valencia, CA, USA). PCR was done

under standard conditions using the following primers: ITS-1F and ITS-4 for nrITS1,

5.88 and nrITS2 (Gardes & Bruns 1993). PCR reactions were performed with an MJ

PTC-100™ thermocycler (Applied Biosystems, Foster City, CA, USA) or an Eppendorf

Mastercycler® gradient (Eppendorf, North America, Inc.,Westbury, NY, USA). PCR

products were cleaned using 0.5 ul of ExoSAP IT (USB Corp, Cleveland, OH, USA)

per reaction and cycled at 37°C for 45 min, followed by 80°C for 15 min. Sequencing

was performed using Big Dye chemistry, with the same primers as for PCR, and an ABI

PRISM 3100 Genetic Analyzer (both from Applied Biosystems, Foster City, CA, USA).

Sequences were edited and contigs assembled using Sequencher 4.2.2 (Gene Codes

Corporation, Ann Arbor, MI, USA). All newly produced sequences were deposited in

GenBank (see TABLE I).

PHYLOGENETIC METHODS—nrlITS sequences of the European species in the P. involutus

species complex available in GenBank and the UNITE database, from vouchered

specimens and from environmental samples, were downloaded and added to the

database. The sequences were aligned with the program MAFFT version 6 (Katoh et

al. 2002; Katoh & Toh 2008), using default settings. The database was analyzed by a

maximum likelihood (ML) method using RAXML version 7.2.3 (Stamatakis et al. 2008).

Paxillus rubicundulus was chosen as outgroup.

Results

The four phylogenetic European species recognized by Hedh et al. (2008)

are also recovered in our nrITS sequence analyses (FIG. 1), with high bootstrap

support for P. validus/P. ammoniavirescens (95%), P. involutus I (100%), and

P. involutus II (100%). The type collection of P ammoniavirescens and the type

collection of P. validus have identical ITS sequences. Collections from England

354 ... Vellinga & al.

and France with sequences newly generated for this study also belong to

P. ammoniavirescens/P. validus, plus one collection from New Zealand.

Sister to the P. ammoniavirescens/P. validus clade is P obscurosporus, with

collections from all over Europe, and one from China. The other two taxa

represent P. involutus clade I, with specimens in Europe and North America

(Wisconsin, Tennessee, British Columbia, Alaska), and P. involutus clade II,

with representatives in Europe and in western North America where they grow

with introduced Betula. Paxillus involutus I includes a specimen from England

that lacks pigments.

Discussion

Hedh et al. (2008) used five genes, AcTA, GPIA, HYDA, RABA, and ss-TUBA,

for phylogenetic species recognition in the P. involutus group, but concluded

that nrITS on its own discriminated the four phylogenetic species recovered

from the multi-gene analyses. Hence, for this study, nrITS was used to assign

specimens to species. However, nrITS sequences alone are not sufficient to

differentiate the North American species P. vernalis Watling from P. validus

(Hedh et al. 2008), nor western North American collections that are very close

to P. obscurosporus and grow with native and introduced tree species in native

and anthropogenic habitats (data not shown).

The nrITS sequences of the type collections of P ammoniavirescens

(Genbank JN661718) and P. validus (Genbank EU084666) are identical. The

two original descriptions emphasize different characters, e.g. Hahn & Agerer

(1999) described the relatively big crystals on the rhizomorphs of P. validus,

whereas Dessi & Contu (1999) did not comment on this character. The reactions

of the various parts of the basidiocarps with a set of chemicals is given in both

articles, with slight differences. Dessi & Contu (1999) emphasized the green

reaction of the pileipellis in ammonia, whereas Hahn & Agerer described this

reaction as ‘schwarz tiberhaucht;’ this difference in reaction might be caused

by differences in age of the basidiocarp or by different concentrations of

the reagent, but neither set of authors provided that information. For KOH

different concentrations were used: 15% by Hahn & Agerer (1999), 5% by Dessi

& Contu (1999). In both descriptions the pileus is introduced as ribbed, the two

species are similar in general stature, and the spore sizes and spore print colour

of the two are comparable. Thus we conclude here, based on the molecular

evidence, that the two taxa are conspecific. The next question is which name

was validly published first. Both names were published in 1999. Paxillus validus

Fic. 1. nrITS phylogeny of European Paxillus species inferred by maximum likelihood (ML) analysis.

Numbers at internodes refer to confidence estimates based on 100 rapid ML bootstraps. Type

collections are in bold; * indicates newly created sequences and “white’ indicates the variant

without pigment in the pileus. Paxillus rubicundulus is used as outgroup.

Three Paxillus species revisited ... 355

JN673718

FR852277

JN661725 *

100 f JN673719

GQ389624

EU078742

JN673720

AY585922

JN661728 *

Nee oot involutus II

EU346878

AY585921

JN661726 *

FR750011

UDBO001205

Nee fet fSes

aEAG On involutus |

AF167696

HM015471

96 HM015494

JN673721

FJ816734

EU078738

EU488709

HQ604826

FJ816736

FJ188353

HQ999961

AJ438984

EU488707

HQ021863

HM1646457

HM164646

EU078735

AF167698

DQ068961

FJ158057

100 HM146842

AY 230243

EU819416

AF167695

DQ367909

JN661723 *

UDB001204

JN673722

FN565299

AY585920

AF 167697

EU078739

100 AY585918

JN661722 *

DQ233903

DQ233899

JN673723

DQ179126

EU486436

FR877522

JN661721 *

HM164647

‘$U799609

GU799607 obscurosporus

AF 167691

EU078714

AY585912

JN172978

GU799610

UDB001480

GU799608

EU84665 obscurosporus Type

GU799612

DQ647827

V3 AF167693

FJ820574

AF 167692

EU078715

AF 167694

GU799614

GU799613

GQ389623

JN661714 *

WNO6T716>

1716* bia

EU346879 ammonlavirescens

AM084700

JN661711 *

AY585917

GQ154499

9 5 EU084666 validus Type

JN661717 *

JN661713 *

JN661719 *

JN661718 * ammoniavirescens Type

AY585915

JN673717 NZ

JN661715 *

0.06 euosera

0.06 AF 167688

EU084664 f

100 FM993258 rubicundulus

AF 167687

FM993114

356 ... Vellinga & al.

Fic. 2. Paxillus involutus s. str. White variant (coll. & photo S. Kelly).

was proposed in Nova Hedwigia 69(1-2), published in August 1999 (according

to the subsequent journal issue). Paxillus ammoniavirescens was published in

Micologia e Vegetazione Mediterranea 13(2) with the nominal date “1998,” but

actually printed and distributed to libraries in Spring [northern hemisphere]

1999. Micologia e Vegetazione Mediterranea 13(2) arrived at L on May 20 1999,

whereas Nova Hedwigia 69(1-2) arrived on 28 October 1999. This indicates

that P ammoniavirescens has priority as the correct name for this taxon.

Paxillus albidulus was described as ‘very similar’ to P involutus but

distinguished from that species by the white pileus, stipe, flesh, and basal

mycelium. Pigments were present in the hymenium, and the spore print was

‘ochreous to pale brown’ (Sutara 1992). At the time P. albidulus was described,

P. obscurosporus and PB ammoniavirescens had not yet been recognized as

separate species, but the basidiocarp stature and light spore print colour exclude

the possibility that P. albidulus is a close relative of either of these two species.

Hedh et al. (2008) were not able to get sequence data from P. albidulus, so its

identity will remain unresolved. However, the whitish collection (FIG. 2) from

the U.K., also lacking pigments in pileus and stipe, fits well into P involutus I

(Fic. 1). Paxillus albidulus was reported from a predominantly coniferous forest,

and as this also fits well with P. involutus I, we tentatively place P albidulus in

Three Paxillus species revisited ... 357

Fic. 3. Paxillus ammoniavirescens (coll. & photo L. Perrone)

Collection: Italy, Sardinia, prov. Olbia-Tempio P., Olbia, Park “Fausto Noce’,

11 Jan. 2008, det. L. Perrone & M. Contu (herb. Perrone).

synonymy with P involutus. Recently another pale Paxillus variant was studied

that also fits into P involutus s. str. (Gelardi et al. 2011); that variant, described

as P. involutus f. eburneus Gelardi et al., has a yellow colouration lacking in the

variant from England that we studied.

The most common name used for Paxillus sequences deposited in GenBank

is P. involutus; however, in many cases this is not the correct name. The option for

third-party annotations is unfortunately not available, and this case emphasizes

the necessity for it (Bidartondo et al. 2008).

Most taxa are widely distributed throughout the northern hemisphere,

but unfortunately there are very few data from Asia. Paxillus involutus | is

recorded from many European countries and North America (from Alaska to

Tennessee in the southeast). Paxillus involutus II is widespread in Europe (and

possibly also in boreal North America) but is better known from introduced

Betula planted in the urban areas of the North American western states

(Washington to California); its occurrence in the Great Smoky Mountains NP

(Tennessee) is also confirmed (Fic. 1). Paxillus obscurosporus is widespread

in Europe and known from China (GenBank DQ647827, as ‘P. vernalis’).

Lastly, P ammoniavirescens is known from Europe and introduced into New

Zealand. [Southern hemisphere Paxillus species have been shown to belong

358 ... Vellinga & al.

to a separate lineage and family for which Bresinsky et al. (1999) erected

the genus Austropaxillus Bresinsky & Jarosch]. Vizzini (2008) identified a

Moroccan collection growing under Eucalyptus as P ammoniavirescens. After

publication of Hahn & Agerer (1999) many more distribution and habitat data

have become available for P ammoniavirescens and P. obscurosporus; Hahn &

Agerer’s speculations that these are introduced species in Europe no longer

seem to be justified, as both species have been recovered from native habitats

within Europe and records from outside Europe are scarce. Hedh et al. (2008,

2009), who recorded all four species from the same county in Sweden, have

shown that strains representing all four phylogenetic species were able to form

ectomycorrhizas with Betula and with Picea seedlings, but not all tested strains

succeeded in forming mycorrhizas and the success rate among strains varied

greatly as well. How the strains behave under natural conditions is, of course,

a different question.

Taxonomy

Paxillus ammoniavirescens Contu & Dessi, in Dessi & Contu, Micol. Veg. Medit.

13(2): 123. March 1999 [“1998”]. Fig. 3

HOLOTYPE: IB, 21 Nov. 1997, P. Dessi; GenBank nrITS JN661718.

= Paxillus validus C. Hahn, in Hahn & Agerer, Nova Hedwigia 69(1-2): 261. August 1999.

HotortyPe: M, 8 Oct. 1997, C. Hahn, CH 243-97; GenBank nrITS EU84666.

Acknowledgements

The loan of the type collection of P ammoniavirescens from IB is greatly appreciated.

Alan Hills was instrumental in the understanding of P ammoniavirescens and we

warmly thank him for sending many specimens from different habitats. We also thank

Luigi Perrone for permission to use his photo of P. ammoniavirescens. Michael Kuo and

Darvin DeShazer contributed North American specimens, and Manfred Binder and

Karen Hughes allowed us to use their pre-submission nrITS sequences. Comments and

suggestions by Dr. Pierre-Arthur Moreau, Prof. Roy Watling, and Dr. Shaun Pennycook

were very helpful for the final version of the manuscript.

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129-133:

Vizzini A. 2008. Revisione Gomphidiaceae-Paxillaceae. 639-648, in: J-C Maire et al. (eds).

Compléments a la Flore des champignons supérieurs du Maroc de G. Malencon et R. Bertault.

Nice, C.E.M.M.

Wilkinson A, Solan M, Taylor AFS, Alexander JJ, Johnson D. 2010. Intraspecific diversity regulates

fungal productivity and respiration. PLOS one 5: e12604.

http://ds.doi.org/10.1371/journal.pone.0012604

MYCOTAXON

http://dx.doi.org/10.5248/119.361

Volume 119, pp. 361-367 January-March 2012

Two new Heteroconium species

and two other forest microfungi newly recorded from China

SHOU-CAI REN?”, JIAN MA? & XIU-GUO ZHANG"

‘Department of Plant Pathology, Shandong Agricultural University, Taian, 271018, China

*Zaozhuang Vocational College, Zaozhuang, 277800, China

*CORRESPONDENCE TO: zhxg@sdau.edu.cn, sdau613@163.com

ABSTRACT — Two new species, Heteroconium annesleae on decaying branches of Anneslea

hainanensis and H. neolitseae on decaying branches of Neolitsea obtusifolia, are described,

illustrated, and compared with closely related taxa. Illustrations are also provided of two

other anamorphic fungi, Xenoheteroconium bicolor and Craspedodidymum fimbriatum,

which are recorded for the first time from China.

KEY worps — conidial fungi, Hainan Province, systematics

Introduction

Petrak (1949) erected Heteroconium with H. citharexyli Petr. as the type

species. The genus is distinguished by euseptate, fusiform, cylindrical to oblong

conidia that arise in acropetal unbranched chains from solitary conidiophores

with monoblastic, terminal, determinate or percurrent conidiogenous cells

(Petrak 1949, Ellis 1971, Castaneda et al. 1999, Taylor et al. 2001). Castafieda

et al. (2008) provided a key to 16 species of Heteroconium, and 19 species are

currently accepted. Few other genera produce unbranched acropetal chains

of phragmoconidia (which can almost be scolecoconidial). Lobatopedis has

branched conidiophores with lobed basal cells and rhexolytic conidial secession.

Lylea has short inconspicuous conidiophores and symmetrical cylindrical

distoseptate conidia with rounded ends and extremely thick walls. Its primary

conidia may produce lateral conidia that, as in Xenoheteroconium, can develop

from non-apical cells. Pirozynskiella has unbranched acropetal conidial chains

but is a tropical mycoparasite of Asterinaceae.

Xenoheteroconium was established by Bhat et al. (Bhat & Kendrick 1993)

to accommodate X. bicolor. The genus is unique in having a bi-coloured

362 ... Ren, Ma & Zhang

conidiophore, with the darker apical portion developing lateral conidia before

seceding and acting as a propagule.

Craspedodidymum Hol.-Jech. (Holubova-Jechova 1972) is typified by C. elatum

Hol.-Jech., which has branched conidiophores terminating in apically inflated

phialides with large funnel-shaped collarettes, producing pale brown aseptate

conidia in slimy heads.

Decaying wood in the tropical forests of Hainan Province supports a high

level of fungal diversity, and many wood-inhabiting macrofungi and microfungi

have been recently discovered there (Ma et al. 2008, Dai et al. 2009, Zhang

et al. 2009a,b, Dai & Li 2010). During continuing investigations of saprobic

microfungi, numerous conidial species were collected on decaying branches,

among which two have morphological characteristics of Heteroconium. They

differ from any described species and are proposed as new to science. Two other

fungi are recorded for the first time from China. The specimens are deposited

in HSAUP (Herbarium of the Department of Plant Pathology, Shandong

Agricultural University) and HMAS (Mycological Herbarium, Institute of

Microbiology, Chinese Academy of Sciences).

A » B Cc D

Fic. 1. Heteroconium annesleae.

A-B. Conidiophores, conidiogenous cells and conidia. C-D. Conidia.

Heteroconium spp. nov. (China) ... 363

Heteroconium annesleae S.C. Ren & X.G. Zhang, sp. nov. Fic. 1

MycoBank MB563628

Differs from Heteroconium decorosum in producing much larger (95-300 x 9.5-15 um)

and differently shaped conidia.

Type: China. Hainan Province: tropical forest of Bawangling, on decaying branches

of Anneslea hainanensis (Kobuski) Hu (Theaceae), 15 May 2011, Sh.C. Ren (holotype,

HSAUP H8492; isotype HMAS 146160).

ETyMOLoey: in reference to the host genus.

COLONIES on natural substratum effuse, hairy, dark brown. Mycelium partly

superficial, partlyimmersed, composed ofseptate, smooth, pale brown, branched,

2-5 um wide hyphae. CONIDIOPHORES macronematous, mononematous,

solitary, erect, unbranched, cylindrical, straight, smooth, brown, 3-5-septate,

55-65 x 4.5-8.5 um. CONIDIOGENOUS CELLS integrated, terminal, monoblastic,

cylindrical, tapered to a truncate apex, determinate, smooth, brown. Conidial

secession schizolytic. ConipIA acrogenous, holoblastic, blastocatenate,

obclavate, tapering gradually towards the apex, base truncate, smooth, 9-24-

euseptate, brown, apical cells pale brown, 95-300 x 9.5-15 um.

Notes: Heteroconium annesleae is similar to H. decorosum R.F. Castafieda et

al. (Castafieda et al. 1999) in conidial morphology, but differs from the latter

in conidial shape, size, and number of septa. In H. decorosum the conidia are

navicular, broadly fusiform to obclavate, 20-30 x 3-5 um, and 3-6-septate.

Heteroconium neolitseae S.C. Ren & X.G. Zhang, sp. nov. Fic. 2

MycoBank MB563630

Differs from Heteroconium tropicale by longer (80-95 um) conidia with more septa (9-

16) and from H. arundicum by narrower (4-6 um) conidia with more septa.

Type: China. Hainan Province: tropical forest of Bawangling, on decaying branches of

Neolitsea obtusifolia Merr. (Neolitsea). 8 May 2011, Sh.C. Ren, (holotype HSAUP H8375;

isotype HMAS 146161).

ETryMoLoey: in reference to the host genus.

COLONIES on natural substratum effuse, hairy, dark brown. Mycelium partly

superficial, partly immersed, composed of branched, septate, smooth, pale

brown, 2-4 um wide hyphae. CoNIDIOPHORES macronematous, mononematous,

solitary, erect, unbranched, straight, smooth, brown, 9-13-septate, 80-95 x 4.5-6.5

um. CONIDIOGENOUS CELLS integrated, terminal, monoblastic, cylindrical,

tapered to a truncate apex, determinate, smooth, brown. Conidial secession

schizolytic. Conrp1a acrogenous, holoblastic, blastocatenate, cylindrical or

fusiform, base truncate, smooth, 9-16-euseptate, with a wide black band at

each septum, brown, unevenly pigmented, pale brown ends, 50-85 x 4-6 um.

Notes: Heteroconium neolitseae is similar to H. tropicale R.F. Castahieda & W.B.

Kendr. (Castafieda & Kendrick 1990) and H. arundicum Chowdhry (Chowdhry

364 ... Ren, Ma & Zhang

woz

Fic. 2. Heteroconium neolitseae.

A-B. Conidiophores, conidiogenous cells and conidia. C. Conidia.

1980) in producing smooth cylindrical to fusiform conidia. However, H. tropicale

is distinguished by conidia that are shorter (26-52 um) and have fewer (3-6)

septa and H. arundicum differs in conidia that are wider (8-12 um) and with

fewer (1-10) septa. In addition, H. neolitseae has a black band at each septum of

conidia, a feature not found in either H. tropicale or H. arundicum.

Xenoheteroconium bicolor Bhat, W.B. Kendr. & Nag Raj, Mycotaxon 49: 81 (1993)

Fic. 3

COLONIES on natural substratum effuse, dark brown, velvety. Mycelium

mostly immersed in the substratum, composed of branched, septate, pale

Heteroconium spp. nov. (China) ... 365

umlgz

Fic. 3. Xenoheteroconium bicolor.

Conidiophores, primary conidia in branched chains (indicated by white arrows)

and secondary conidia (bases indicated by black arrows).

brown to brown, smooth-walled hyphae. CoNIDIOPHORES macronematous,

mononematous, simple, unbranched, erect, straight or slightly flexuous,

cylindrical, smooth, lower part pale brown, upper part brown, with a sharp line

of demarcation between the two levels of pigment, 5-7-septate, 160-195 x 3-4

um, broadening to 5-6 um wide at the lobed base, regenerating percurrently

from broken ends. CONIDIOGENOUS CELLS monoblastic, integrated, terminal

and intercalary in the upper part of the conidiophore axis and in primary

conidia, brown, cylindrical or conical. Conrp1a holoblastic, phragmosporous,

dry, brown; primary conidia fusiform or obclavate, straight, 2-4-euseptate,

slightly constricted at the septa, 15-37 x 3.5-4.5 um; secondary conidia

cylindrical or fusiform, truncate at the base, tapering toward the tip, straight,

5-7-euseptate, slightly constricted at the septa, 35-80 x 3-4 um.

SPECIMEN EXAMINED: CHINA. Hainan Province, tropical forest of Bawangling, on

decaying branches of unidentified plant, 8 May 2011, Sh.C. Ren, HSAUP H8345; HMAS

146162.

366 ... Ren, Ma & Zhang

Notes: Only one species is currently accepted in Xenoheteroconium. The

morphology of our specimen overlaps with the type description by Bhat &

Kendrick (1993).

Craspedodidymum fimbriatum Bhat & W.B. Kendr., Mycotaxon 49: 33 (1993) Fra. 4

COLONIES gregarious, effuse, velvety, dark brown to black. Mycelium

mostly immersed in the substratum, composed of septate, smooth, pale brown,

branched hyphae. CONIDIOPHORES mononematous, fasciculate in groups, erect,

straight or flexuous, unbranched, 4—8-septate, smooth, brown, paler towards

the apex, percurrently regenerating, 135-210 x 4-6 um. CONIDIOGENOUS

CELLS monophialidic, terminal, integrated, cylindrical to clavate, slightly

inflated and rounded at the apex, which bears an inconspicuous collarette,

pale brown to brown. Conip1A blastic-phialidic, globose, non-septate, thick-

walled, pale brown, 19-22 um diam., with numerous acellular, fibrillose, curved

appendages forming a pile or coat on the surface; accumulating in colourless

Fic. 4. Craspedodidymum fimbriatum. A-B. Conidiophores and conidiogenous cells.

C-D. Conidiophores and conidia. E. Conidia.

Heteroconium spp. nov. (China) ... 367

slimy masses at the apex of the conidiophore, or sometimes adhering to the

sides of conidiogenous cells after secession.

SPECIMEN EXAMINED: CHINA. Hainan Province, tropical forest of Bawangling, on

decaying branches of unidentified plant, 8 May 2011, Sh.C. Ren, HSAUP H8405; HMAS

146164.

Notes: Craspedodidymum fimbriatum is unique in possessing a cylindrical to

clavate conidiogenous cell, which is inflated and rounded at the apex and with

an inconspicuous collarette, and slimy globose aseptate conidia with a densely

fibrillose surface. The size ranges of conidia and conidiophores in our specimen

overlap with those of the type specimen described by Bhat & Kendrick (1993),

and other features of this taxon also match those of the original species.

Acknowledgments

The authors express gratitude to Dr Eric H.C. McKenzie and Dr WB. Kendrick for

serving as pre-submission reviewers and for their valuable comments and suggestions.

This project was supported by the National Natural Science Foundation of China (Nos.

31093440, 30499340, 30770015) and the Ministry of Science and Technology of the

People’s Republic of China (Nos. 2006FY120100, 2006FY110500-5).

Literature cited

Bhat DJ, Kendrick WB. 1993. Twenty-five new conidial fungi from the Western Ghats and the

Andaman Islands (India). Mycotaxon 49: 19-90.

Castaneda RF, Kendrick B. 1990. Conidial fungi from Cuba: II. Univ. Waterloo Biol. Ser. 33: 1-62.

Castafieda RE Saikawa M, Guarro J. 1999. A new species of Heteroconium from a tropical rainforest.

Mycotaxon 71: 295-300.

Castaneda RF, Iturriaga T, Heredia Abarca G, Minter DW, Gené J, Stadler M, Saikawa M, Silvera-

Simon 2008. Notes on Heteroconium and a new species from Venezuela. Mycotaxon 105:

175-184.

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey,

England. 608 p.

Dai YC, Li HJ. 2010. Notes on Hydnochaete (Hymenochaetales) with a seta-less new species

discovered in China. Mycotaxon 111: 481-487. http://dx.doi.org/10.5248/111.481

Dai YC, Cui BK, Yuan HS. 2009. Trichaptum (Basidiomycota, Polyporaceae) from China with a

description of three new species. Mycol. Prog. 8: 281-287.

http://dx.doi.org/10.1007/s11557-009-0598-0.

Holubova-Jechova V. 1972. Craspedodidymum, a new genus of phialosporous hyphomycetes. Ceska

Mykologie 26: 70-73.

Ma J, Zhang K, Zhang XG. 2008. Taxonomic studies of Corynespora from Hainan, China.

Mycotaxon 104: 153-157.

Petrak F .1949. Neue Hyphomyzeten-Gattungen aus Ekuador. Sydowia 3: 259-266.

Taylor JE, Crous PW, Palm ME. 2001. Foliar and stem fungal pathogens of Proteaceae in Hawaii.

Mycotaxon 78: 449-490.

Zhang K, Fu HB, Zhang XG. 2009a. Taxonomic studies of Corynespora from Hainan, China.

Mycotaxon 109: 85-93. http://dx.doi.org/10.5248/109.85

Zhang K, Ma J, Wang Y, Zhang XG. 2009b. Three new species of Piricaudiopsis from southern

China. Mycologia 101: 417-422. http://dx.doi.org/10.3852/08-147.

MYCOTAXON

http://dx.doi.org/10.5248/119.369

Volume 119, pp. 369-372 January-March 2012

Second record of Ramularia hypericicola —

collected in Turkey on a new host

FARUK SELCUK’, ELSAD HUSEYIN™ & ALIS. BULBUL”

‘Department of Biology, Arts and Sciences Faculty, Ahi Evran University,

Kirsehir, 40100, Turkey

*Department of Biology, Sciences Faculty, Gazi University,

Teknik okullar, Ankara, 06500, Turkey

*CORRESPONDENCE TO: elsadhuseyin@hotmail.com

ABSTRACT — Ramularia hypericicola, a rare anamorphic Mycosphaerella species, is reported

and illustrated from Turkey on Hypericum calycinum. The Turkish collection represents the

second report of this fungus, known hitherto only from the type locality.

KEY worpDs — symptoms, Clusiaceae, morphology

Introduction

Most species of Ramularia are phytopathogenic, causing leaf spots,

sometimes chlorosis, or without visible symptoms, occasionally saprobic

or hyperparasitic (Braun 1998). Bremer & Petrak (1947) published the first

records of Ramularia species from Turkey, R. anatolica Bremer & Petr. and

R. helminthiae Bremer & Petr. Subsequent papers about Turkish microfungi,

including Ramularia species, were published by Bremer et al. (1948, 1952a,

1952b) and Petrak (1953, 1957). Additional Turkish records, of Ramularia

cynarae Sacc., R. onobrychidis Allesch., and R. sambucina Sacc., were published

by Karel (1958). The relatively small number (nearly 30 species) of Ramularia

species recorded in subsequent papers (Gébelez 1964) reflects a rather poor

degree of research on microfungi on plants in Turkey. However, during the

past two decades research on micromycetes (including hyphomycetes) in the

country has intensified (Altan & Tamer 1996, Hiiseyinov & Selcuk 1999, Braun

et al. 2000, Hiiseyin & Selcuk 2001, Hiseyinov et al. 2002, Hiiseyin et al. 2003,

2005, Melnik et al. 2004, Erdogdu & Hiiseyin 2007, Selcuk et al. 2009, 2010).

We describe here one recent collection from Turkey of a Ramularia species

previously recorded only from Russia.

370 ... Selcuk, Hiiseyin & Bulbul

~ 4 . < o ~*

a ~>

lbw All _. — |.

“ ~

FiGuRE 1. Ramularia hypericicola on Hypericum calycinum; a, spots on infected host leaves;

b, punctiform grayish white caespituli; c, small fasicle of conidiophores; d, conidia.

Material & methods

The plant material was gathered from Gemlik (Bursa Province). The host plant was

identified by reference to Davis (1967). Sections of the fungus were hand cut using a

razor blade and microscopically examined using a Leica DM LB research microscope.

The fungus was identified following Braun (1998). Author abbreviations follow Index

Fungorum (2011). All examined specimens are deposited in the herbarium of Ahi

Evran University, Arts and Sciences Faculty, Department of Biology, Kirsehir Province,

Turkey.

Results

After careful comparison with Ramularia spp. described on hosts of

Hypericaceae based on Braun (1998) the recently collected material on

Hypericum calycinum from ‘Turkey proved to be clearly identical with

R. hypericicola. The following description and illustration are based on the

Turkish material.

Ramularia hypericicola U.Braun, Monogr. Cercosporella, Ramularia 2: 170, 1998.

FIG. 1

On both surfaces of leaves causing subcircular to irregular, occasionally

angular and vein-limited, scattered sometimes confluent spots, covers more

than 50% of the surface of some leaflet, firstly 0.5-1 mm across, then up to 10

mm diam. Spots on upper surface dirty brownish, dull, lover surface clearly

A new host for Ramularia hypericicola (Turkey) ... 371

brown, without distinct margin. Caespituli hypophyllous, punctiform, grayish

white. Mycelium internal, colourless, hyphae septate, branched, formed

stromatic, colourless to yellowish-ochraceous, subglobose or slightly lengthened

hyphal aggregations in leaflet tissues, 15-70(-75) um diam. Conidiophores in

small to moderately large fascicles, loose to usually dense, emerging through

stomata or erumpent through the cuticle, erect, simple, straight, subcylindrical

to geniculate-sinuous, continuous to sparingly septate, hyaline or subhyaline,

smooth, 6-65(-70) x 2.5-5 um. Conidiogenous cells terminal, colourless,

enteroblastic. Conidiogenous scars thickened, darkened. Conidia colourless,

catenate, chains occasionally branched, produced enteroblastically, single

conidia smooth to faintly rough, ellipsoid-ovoid, subcylindrical, fusiform,

10-30(-35) x 4-5(-6) um, unicellular or with a single median septa, not

narrower at the septa, ends rounded or somewhat attenuated.

SPECIMEN EXAMINED —- TURKEY, Bursa PROVINCE, GEMLIK DISTRICT, Haydariye

village, near Selale, alt. 700 m, on living leaves of Hypericum calycinum L. (Clusiaceae),

in hornbeam-beech (Carpineto-Fagetum) forests, 18 Nov 2010, coll. E. Hiiseyin G-06.

Discussion

In 1956, the Russian mycologist M.K. Khokhryakov collected a Hypericum

sp. with Ramularia-type sporulation in the Russian Far East, which he deposited

in LEP as Ramularia sp. Braun (1998: 170) described R. hypericicola as a new

species based on this material. Ramularia hypericicola is undoubtedly a rare

anamorph of Mycosphaerella. The present record from Turkey represents the

second collection of this species and the first on a known host, Hypericum

calycinum.

Acknowledgments

The authors would like to extend a sincere gratitude to Uwe Braun (Martin-

Luther- Universitat, Halle/Saale, Germany) for linguistic help and confirmation of our

identification. We also thank Dr. Olga Vinnere Pettersson (Uppsala, Sweden) and Dr.

Tatiana Andrianova (Kiev, Ukraine) for critically reading the manuscript and serving

as presubmission reviewers. Especially we grateful to Dr. Shaun Pennycook (Auckland,

New Zealand) for detailed linguistic help and nomenclatural review of the manuscript.

Literature cited

Altan Y, Tamer AU. 1996. The parasitic fungi occurring on some endemic plants in Turkey and

their damaging effects. 398-401, in: Plant life in South-West and Central Asia. Ege University

Pres, Izmir, Turkey.

Braun U. 1998. A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic

hyphomycetes). Vol. 2. IHW-Verlag. 493 p.

Braun U, Melnik V, Hiiseyinov E, Selcuk F. 2000. Mycopappus alni on species of Betula and Pyrus

from Turkey. Mikologia i Fitopatologia 34(6): 1-2.

Bremer H, Petrak F. 1947. Neue Kleinpilze aus der Tiirkei. Sydowia, 1(1 - 3): 248 - 263.

372 ... Selcuk, Hiiseyin & Bulbul

Bremer H, Ismen H, Karel G, Ozkan M. 1948. Beitrage zur Kenntnis der parasitischen Pilze der

Tuirkei. III. Revue de la Faculté des Sciences de l‘Université d‘Istanbul, Ser. B, 13(1): 1-53.

Bremer H, Karel G, Bryikoglu K, Goksel N, Petrak F. 1952a. Beitrage zur Kenntnis der parasitischen

Pilze der Tiirkei. VI. Revue de la Faculté des Sciences de |‘Université d‘ Istanbul, Ser. B, 17(3):

259-276.

Bremer H, Karel G, Biyikoglu K, Géksel N, Petrak F. 1952b. Beitrage zur Kenntnis der parasitischen

Pilze der Tiirkei. VII. Revue de la Faculté des Sciences de ‘Université d‘ Istanbul, Ser. B, 17(4):

277-288.

Davis PH. (ed.). 1967. Flora of Turkey and East Aegean Islands. Vol. 2. Edinburgh University

Press.

Erdogdu M, Hiiseyin E. 2007. The world’s second record of Hyalodictyum colchicum reported from

Turkey. Mycotaxon 99: 245-250.

Gébelez M. 1964. La mycoflore de Turquie. I. Mycopathologia et Mycologia Applicata 23(1):

47-67.

Hiiseyin E, Selcuk F. 2001. New and poorly known genera of microfungi for Turkey. Turkish Journal

of Botany 25(6): 437-438.

Hiiseyin E, Selcuk FE, Gaffaroglu M. 2003. Some materials on mitosporic fungi from Turkey. I.

Hyphomycetes. [Kai kurie duomenys apie Turkijos mitosporinius grybus. I. Hyphomycetes].

Botanica Lithuanica 9(2): 151-160.

Hiiseyin E, Selcuk F, Sahin A. 2005. The world’s second record of Neoheteroceras flageolotii reported

from Turkey. Mycotaxon 94: 241-244.

Hiiseyinov E., Melnik V., Selcuk E 2002. Ceratophorum helicosporum - a new for mycoflora of

Turkey genus and species of Hyphomycetes (Dematiaceae). Mikologia i Fitopatologia 36(3):

11-13.

Hiiseyinov E., Selcuk F. 1999. New records of phytopathogenic microfungi for Turkey. Plant Disease

Research 14(2): 175-176.

Karel GA. 1958. A preliminary list of plant diseases in Turkey. Ayyildiz Matbaas1.Ankara.

Melnik V, Hiiseyin E, Selcuk EF. 2004. Contribution to the studying of micromycetes in several Black

Sea provinces of Turkey. Novitates Systematicae Plantarum non Vascularum. Nauka. Petropolis

37: 133-148.

Petrak FE. 1953. Neue Beitrage zur Pilzflora der Tiirkei. Sydowia 7: 14-44.

Petrak F. 1957. Beitrage zur Tiirkishen Pilzflora. Sydowia 10: 101-111.

Selcuk FE, Erdogdu M, Akgil H, Huseyin E. 2009. The genus Septoria Sacc. in Turkey. Mycopath

7(1): 21-28.

Selcuk F, Erdogdu M, Hiiseyin E, Gdcmen E. 2010. Tiirkiyeden Rapor Edilmis Piknidial, Aservular

ve Stromatal Yapilarla Eseysiz Ureyen Mikrofunguslar (The microfungi which growth pycnidial,

acervular and stromata structures, reported from Turkey). Ekoloji 2010 Sempozyumu. Aksaray

Universitesi Fen — Edebiyat Fakiiltesi Biyoloji Béliimii ve Tiirkiye Tabiatin1 Koruma Dernei.

Ozetler. 5-7 Mayis 2010, Aksaray. S. 186.

MYCOTAXON

http://dx.doi.org/10.5248/119.373

Volume 119, pp. 373-380 January-March 2012

Lichenological notes 4:

a revision of Acarospora gallica (Acarosporaceae)

KERRY KNUDSEN * & JANA KOCOURKOVA?

"The Herbarium, Department of Botany and Plant Sciences, University of California,

Riverside, CA 92521-0124, U.S.A.

*Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences,

Prague, Kamycka 129, Praha 6 - Suchdol, CZ-165 21, Czech Republic

* CORRESPONDENCE TO: 'Knudsen@ucr.edu*,’ kocourkovaj@fzp.czu.cz

ABSTRACT — Acarospora gallica from Europe and Asia is revised and a neotype is designated.

Acarospora hungarica is proposed as a synonym of A. gallica. The anamorphic stage is

reported for the first time from the species.

Key worps — Acarospora fuscata, Acarospora janae, Czech Republic

Introduction

The study of the genus Acarospora in Europe rests on the strong foundation

of the great Swedish lichenologist A.H. Magnusson, whose primary work is his

classical monograph (Magnusson 1929). Nonetheless, some of the species he

described are known only from the type specimens or today are poorly known

by lichenologists in general. Acarospora gallica is among those poorly known

species, mainly because of confusion with A. fuscata, which it can superficially

resemble. It also may be a rare or infrequent species in many countries. In

this paper we revise A. gallica as part of our study of Acarosporaceae in the

Czech Republic. We hope this paper will lead to easier and more confident

determinations of the species and to new discoveries.

Materials & methods

Specimens were studied from GZU, PRA, UPS, and the Hb. Mycologicum J.

Kocourkova & K. Knudsen (Hb. Myc. JK & KK). Specimens were examined using

standard microscopical techniques with an Olympus SZX 7 stereomicroscope or an

Olympus BX 51 fitted with Nomarski differential interference contrast. Hand-made

sections were studied in water and 10% KOH [K]. Amyloid reactions were tested in

Lugol's iodine [I] with and without pretreatment with K. Ascospores were measured in

374 ... Knudsen & Kocourkova

water with an accuracy of 0.5 um. Macrophotographs were taken with a digital camera

Olympus DP72 mounted with QuickPhoto Camera 2.3 on the stereomicroscope. The

illustrations were prepared using Adobe Photoshop.

Taxonomy

Acarospora gallica H. Magn., Kungl. Svenska Vetensk.-Acad. Handl., Ser. 3, 7(4):

282.1929: PLATES 1-2

TYPE: France. Puy-de-Dome: Arlane, pres d.Ambert, 1900, Breviere (Hb. B. de Lesdain,

Holotype, n.v., probably lost). Provence-Alpes-Céte d’Azur: Var, La Courtines prés

Ollivules. Sur rochers volcaniques. May, 1923, de Crozals (UPS, Neotype designated

here).

= Acarospora hungarica H. Magn., Kungl. Svenska Vetensk.-

Acad. Handl., Ser. 3, 7(4): 284. 1929, syn. nov.

Type: Hungary. Saro; Eperjes, 1868, Lokja (Holotype, W!).

Thallus arising from an endosubstatic hypothallus, sometimes with well-

developed rhizohyphae or expanding through vegetative division. Thallus

indeterminate, of dispersed or contiguous areoles, or dispersed or crowded

squamules, 0.2-0.6(-1.5) mm diam., < 0.2-0.5 mm thick. Areoles angular

to irregular or round, broadly attached, sometimes with a single apothecium

the thallus becoming reduced to a pseudothalline margin forming verrucae.

Areoles becoming squamulose and lobulate in well-developed specimens,

with a distinct lower surface, the elongated mycelial base broad (gomphate) or

forming a stipe. Upper surface pale to dark brown, epruinose, matt. Lacking

either an epinecral layer or syncortex of polysaccharides. Lower surface pale

brown or white in well-developed areoles, or darkened by substrate interaction.

Cortex relatively thin, 20-35 um thick, of paraplectenchymatous cells mostly

2-4 um diam.; upper layer brown to 10 um thick, lower layer hyaline. Algal

layer 50-100 um, continuous, sometimes with edges of stratum uneven, algal

cells mostly 10 um diam. Medulla up to 100 um thick, of intricate thick-walled

hyphae, mostly 3-4 um thick, continuous with attaching hyphae.

Apothecia 1-12 per areole or squamule, sometimes in agglomerations; disc

usually smooth, usually reddish when wetted, epruinose, 0.1-0.6 mm diam.,

but sometimes scabrid with ontogenic remnants of thalline plectenchyma

or merging with other apothecia especially when in agglomerations. Exciple

indistinct to 10-15 um wide. Hymenium 75-100(-130) um tall, hyaline, I+ red,

blue or greenish, epihymenium 10-20 um thick, reddish-brown. Paraphyses

1.5-2.0 um diam., not branching, apices slightly expanded to 3 um, sometimes

wider. Asci 50-80 x 15-30, 100-200 ascospores per ascus. Ascospores hyaline,

simple, mostly (2.0-)4.0-5.0(-6.5) x 1.5-2.0 um, usually narrowly ellipsoid,

but shorter ascospores broadly ellipsoid. Subhymenium usually 10-30(-50)

um tall, usually I+ blue. Hypothecium narrow, to 15 um wide, continuous with

exciple.

Acarospora gallica, neotypified ... 375

> Re

ey “ed: oe

PLaTE 1. Acarospora gallica (Knudsen 12447). A, Thallus. Angular, squamulose, and lobate

areoles with apothecia. B, Single and confluent apothecia. Some areoles with fungus Lichenothelia

convexa. Scales = A = 1 mm; B = 0.5 mm.

376 ... Knudsen & Kocourkova

Pycnidia usually visible as red dots, globose, to 100 um diam., sometimes

rare or absent. Conidia hyaline, simple, mostly 2.5 x 1 um.

CHEMISTRY— Gyrophoric/lecanoric acid, cortex and/or medulla KC+

reddish. Other spot tests negative.

ECOLOGY & DISTRIBUTION— In sunny, open locations on diabase, mica-

schist, sandstone, serpentine, shale, silicate-rich and volcanic rock, mostly at

low (100-300 m) elevations in Asia (Siberia, Turkey) and Europe (Austria,

Bulgaria, Czech Republic, Denmark, France, Germany, Hungary, Italy, Spain,

Slovakia, Switzerland, Ukraine) (Magnusson 1929, 1936; John & Nimis 1998).

Golubkova's reports of Acarospora gallica from the former Soviet Union appear

to be based on Magnusson’ reports, indicating that the species may be rare in

the Russian Federation (Golubkova 1988). The most well developed specimens

were collected in central Europe at low elevations. The highest elevation was

1820 m in Italy, collected by H. Mayrhofer (GZU).

SELECTED SPECIMENS EXAMINED: Austria: TIROL, on a roof of mica-schist, 900 m,

Aug. 7, 1927, A.H. Magnusson 10954 (UPS). Bulgaria: RHopopE, inter Be¢kovo et

Staminaka, 250-330 m, ad rupes gneissaceas ad vicum publicam, Aug. 9, 1923, J. Suza

s.n. (PRM 578430). Czech Republic: CENTRAL BOHEMIA, Ceské sttedohori mountains,

Vernerice, a phonolit scree above Bobri soutéska pass, 420-480 m, Sept. 26, 1995, Z.

Palice s.n. (PRA); Kiivoklatsko Highland, distr. Rakovnik, between Nezabudice

and Roztoky villages, Nezabudické skaly Nature Reserve, above Berounka River,

50°1'26.101°N 13°50°41.882”E, 390 m, on steep SW-facing slope, on schist outcrop in

oak thin scree forest, Feb. 16, 2002, J. Kocourkova 1142 (PRM 896203); Hracholusky,

Certova skala Nature Reserve, 320 m, at top of steep slope of rock outcrops, on spilite,

June 28, 1996, J. Horakova 2141 (PRM 891402); ibid., 290 m, on steep slope of rocks, on

spilite, July 14, 1997, J. Kocourkova 3436 (PRM 906656); Prazska plosina plateau, distr.

Praha-zapad, Maslovice, Vétrusicka rokle Nature Reserve, 200 m, on SSE-facing rocky

slopes, on rich shale, July 16, 2002, J. Kocourkova 3310 & V. Orthova (PRM 906851);

Prazska plosina plateau, Praha, Mala Ohrada, Albrechtiv hill at Prokopsky brook, 300

m, on diabase rock, Sept. 23, 1999, J. Kocourkova 6747 (PRM 909496); Praha, Motol,

natural monument “Kalvarie, Rock with cross’, W-facing diabase crest with the cross

on top, 50°03’96”N, 14°19°65”E, 320 m, on low diabase outcrop, Apr. 3, 2007, Z. Palice

11396 (PRA); Praha, Pitkovice, Pitkovicka stran, 50°01’26”N, 14°34’21”E, 276 m, on

shale, Sept. 21, 2010, J. Kocourkova 7755 & K. Knudsen 12447 (Hb. Myc. JK & KK), K.

Knudsen 12457 & J. Kocourkova (Hb. Myc. JK & KK, UCR). East BoHEMIA, the Labe

valley, Chvaletice, sedimentation basin NNW of the power station, a dumped stone-

terrace, ca 50°02’20"N, 15°26'40”E, 220 m, on schist, Oct. 27, 2002, Z. Palice 7956 &

Z. Soldan (PRA). SouTH Moravia, in valle fluminis Jihlavka, prope pag. HrubSice, ad

rupe arenacea, 230 m, 1919, J. Suza s.n. (UPS); Ad rupe arenacea prope Hrubiice, ca

230 m, Aug. 5, 1919, J. Suza s.n. (PRM 578429); Distr. Trebi¢, Mohelno, S-SSW-facing

slopes above the valley of Jihlava river, a terrace with well-lit pine-forest, 49°06'26.5°N,

16°11°08” E, 290 m, on small serpentine stone semi-immersed in the ground, Mar. 31,

2011, Z. Palice 14204 (PRA). West Boner, Cesky les, distr. Cheb, Zelezna hirka,

49°59'29.5°N, 12°26'29.5”E, 575 m, on eutrophicated iron-eich phyllitic boulders,

June 24, 2011, J. Halda, Z. Palice & P. Uhlik 14298 (PRA); NorTH BOHEMIA: Protected

Landscape Area Kokofrinsko, former quarry, 300 m, Nov. 27, 1997, Z. Palice s.n. (PRA).

Acarospora gallica, neotypified ... 377

“A, <P soe

PLATE 2. Acarospora gallica (Knudsen 12447).

Areoles. Stipe and well developed rhizohypha. Scale = 1 mm.

Denmark: JYLLAND, SCHLESWIG-HOLSTEI, Insel Alsen, Norburg, Sept. 11, 1913, C.RE.

Erichsen s.n. (UPS); France: PROVENCE-ALPES-COTE D’AzuR, Alpes-Maritimes, Nice,

Peira Cava, on sandstone, Apr., 1927, J. Suzas.n. (UPS). GALLIA MERID., Nice, Peira Cava,

ad saxa arenacea, ca 1500 m, (sepes maritim.), Apr., 1927, Suza s.n. (PRM 578431). Italy:

ARCIPELAGO TOSCANO, Prov. Livorno, Isola di Capraia, 120 m, Apr. 23, 1988, Nimis et

al. s.n. (GZU). NorDL. APENNIN, Prov. Pistoria: Umgebung von Abetone, Val di Luce,

Alpe Tre Potenze, 1820 m, Oct. 27, 1978, H. Mayrhofer s.n. (GZU). Russia: PRIMORYE,

Oriente Extremo, Ditio Ussuriensis, Distr. Vladivostok, in peninsula Bassargin, Sept. 6,

1927, A. Oxner s.n. (UPS). Slovakia: VIHORLAT, Chonkovee, andesit, 1930, J. Nadvornik

s.n. (PRM 783230). Spain: CATALANIA: Vora Tosca, on volcanic rock, Sept. 1934, J.

Maheu s.n. (UPS). Switzerland: GRAUBUNDEN, Engadin, Ardez, 1400 m, Aug. 4, 1927,

A.H. Magnusson 11178 (UPS). Ukraine: Uzhorod, balvany u feky, 1929, J. Nadvornik

s.n. (PRM 783229).

Discussion

Our description of Acarospora gallica differs from Magnusson (1929) in

understanding that the thallus is variable, from areolate to squamulose in a

continuous range of developmental variation. This is probably influenced by

microhabitat conditions. It is often a solitary pioneer found in sunny open sites

on sharp-edged volcanic rocks, serpentine stones, diabase, shale, and even

on toxic substrates in the abandoned sedimentation basins of Chvaletice and

Bukovina in East Bohemia (Palice & Soldan 2004). These populations consist

378 ... Knudsen & Kocourkova

usually of small broadly attached areoles, often with a single apothecium and

the thallus reduced to a pseudothalline margin. On smooth substrates in

microhabitats with more moisture and possibly nitrogen, particularly in central

Europe, the areoles are larger, usually with multiple apothecia, and are often

lobulate and squamulose, with a broad elongated mycelial base (gomphate) or

a narrow stipe.

Specimens of Acarospora gallica with several apothecia per areole correspond

to Magnusson’ original concept of the species (see drawing in Magnusson 1936).

Acarospora gallica var. vinealis H. Magn. has its type locality of Svaty Jur (St.

Georgen) in Slovakia. The name was applied to three specimens with an even,

more contiguous thallus, with areoles with multiple apothecia (Magnusson

1929). The type material of this variety was not available for study, but the

description matches some specimens collected by the second author from Czech

Republic. We observed that areoles with multiple apothecia were usually in the

process of vegetative division or the apothecia eventually merged into a large

irregularly shaped disc. Magnusson (1929) inconsistently referred thalli with

predominately dispersed areoles with a single apothecium (sometimes with the

thallus reduced to a pseudothalline margin) to A. gallica var. devastata (Eitner)

H. Magn. Actually areoles with a single apothecium dominate in the specimens

we examined, possibly because they are easier to distinguish from A. fuscata.

We do not recognize Magnusson’s two varieties as infraspecific categories but

only as typological abstractions of a normal variation. Acarospora hungarica was

described from a single specimen with well-developed < 1 mm wide squamules

(Magnusson 1929). We treat A. hungarica as a synonym of A. gallica.

Acarospora gallica is easily confused with A. fuscata (Ach.) Th. Fr., which

it can resemble in color and which also produces gyrophoric/lecanoric acids.

Acarospora fuscata has a black lower surface and is broadly attached, usually

forming a contiguous areolate crust. It is the most common species throughout

the range of A. gallica. Macromorphological characters are diagnostic for

separating the two species. Acarospora gallica is easily distinguished from

A. fuscata either by its lobulate squamules with pale underside or by

predominance of small areoles with a single apothecium. Poorly developed

specimens of A. gallica may be almost impossible to determine and may be

reported as A. fuscata (as in Palice & Soldan 2004). Squamulose A. gallica

specimens with lower sides darkened by substrate interactions can be

distinguished from A. fuscata by the presence of a well-developed mycelial base

or stipe, lobulate margins, and usually multiple apothecia per areole, sometimes

growing in agglomerations. Careful examination of the thallus, may also reveal

some pale lower surfaces.

Though A. gallica is widespread in Europe and Asia, collections are sparse,

making its distribution, abundance, and conservation status uncertain for each

Acarospora gallica, neotypified ... 379

country where it occurs. It may be undercollected or collections may have

been misdetermined as the ubiquitous A. fuscata. For instance, it was known

in the Czech Republic from several early 20" century collections (Magnusson

1929; Suza 1928, 1931, 1947) but reported only once by Vézda (1998). But our

study has established its presence in eleven localities in Bohemia and Moravia.

Several vouchers were originally identified as A. fuscata or other species. It is

frequent in Czech Republic and not threatened.

Magnusson (1929, 1930) reported A. gallica var. devastata from North

America. We do not recognize A. gallica as occurring in North America and

all the specimens from North America identified by Magnusson belong to

A. janae K. Knudsen (Lumbsch et al. 2011; Knudsen et al. 2011). Acarospora

janae especially differs from A. gallica in forming small areoles or verrucae that

never become squamulose.

Galloway (2007) reported A. gallica from New Zealand, but we are skeptical

of his determination based on the current known distribution of the species.

His taxon may be conspecific with an undescribed taxon from South America

collected by Adam Flakus that we are currently studying.

Another photograph of Acarospora gallica is available on Italic, the

Information System of Italian Lichens (Nimis 2003).

Lichenothelia convexa Henssen is the only lichenicolous fungus we have

collected on Acarospora gallica (Kocourkova & Knudsen 2011). At PRM some

specimens we observed were infected by Lichenostigma gracile Calat. et al.

Acknowledgements

We thank our reviewers, James C. Lendemer (NY) and Adam Flakus (Poland). We

thank the curators of PRA, GZU, UPS, and especially FrantiSek Bouda (PRM) for his

assistance during our visit and company at lunch. We thank Zdenék Palice (PRA) for

supplying us with his recent collections for this paper as well as Mikhail Zhurbenko for

translating N. S. Golubkova. The work of Jana Kocourkova was supported financially by

the grant “Environmental aspects of sustainable development of society”42900/1312/4

23114 from the Faculty of Environmental Sciences, Czech University of Life Sciences

Prague.

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Academy of Sciences of the U.S.S.R. (‘ Nauka’), Leningrad. 136 p.

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380 ... Knudsen & Kocourkova

Kocourkova J, Knudsen K. 2011. Lichenological Notes 2: Lichenothelia convexa, a poorly known

rock-inhabiting and lichenicolous fungus. Mycotaxon 115: 345-351.

http://dx.doi.org/10.5248/115.345

Lumbsch HT, Ahti T, Altermann S$, Amo De Paz G, Aptroot A, Arup U, Barcenas Pefia A, Bawingan

PA, Benatti MN, Betancourt L, Bjork CR, Boonpragob K, Brand M, Bungartz F, Caceres MES,

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MY COTAXON

http://dx.doi.org/10.5248/119.381

Volume 119, pp. 381-384 January-March 2012

The genus Xylaria in the south of China — 3.

X. atroglobosa sp. nov.

Hai-x1A MA’, LARISSA VASILYEVA? & Yu Li?*

' Institute of Tropical Bioscience and Biotechnology, Chinese Academy of

Tropical Agricultural Sciences, Haikou 571101, China

? Institute of Mycology, Jilin Agricultural University, Changchun 130118, China

> Institute of Biology & Soil Science, Far East Branch of the Russian Academy of Sciences,

Vladivostok 690022, Russia

* CORRESPONDENCE TO: yuli966@126.com

Asstract—Xylaria atroglobosa (Xylariales, Xylariaceae) is described from China as a new

species. It is characterized by its semi-globose stromata and ascospores with an appendage on

one end. Photographs of stromata and microstructures are provided.

KEY worDs—Ascomycota, pyrenomycetous fungi, taxonomy

Introduction

The southern provinces of China are characterized by a high diversity of

Xylaria species, and many species known only from southeastern Asia have

been found in southern China. Xylaria badia Pat., described from the Gulf of

Tonkin area in Vietnam (Patouillard 1891), has been reported from Taiwan (Ju

& Rogers 1999), Thailand (Thienhirun & Whalley 2004, Okane et al. 2008),

Malaysia (Whalley & Whalley 2007), and China’s Yunnan Province (Ma 2011).

Xylaria bambusicola Y.M. Ju & J.D. Rogers, described from Taiwan (Ju &

Rogers 1999), is found in Thailand (Okane et al. 2008) and Yunnan Province

(Ma 2011). Xylaria brunneovinosa Y.M. Ju & H.M. Hsieh is only known from

Taiwan (Ju & Hsieh 2007) and Yunnan Province (Ma 2011); X. copelandii

Henn., restricted to Calamus spp. and described from Philippines (Hennings

1908), has been reported from China's Guangdong Province (Ma 2011); and X.

papulis Lloyd was described from China (Lloyd 1921) and later found in Papua

New Guinea (Van der Gucht 1995), Philippines, Taiwan (Ju & Rogers 1999),

and ‘Thailand (Okane et al. 2008). A much longer list, including two recently

described new species, X. choui H.X. Maet al. and X. ficicola H.X. Ma et al. (Ma

et al. 2011a,b), would demonstrate the very unique species composition of the

382 ... Ma, Vasilyeva & Li

genus in southeastern Asia. This paper contributes to our knowledge of Xylaria

in the south of China.

Materials & methods

Collection, preservation, and identification methods follow Ju and Rogers (1999).

Microscopic features and measurements were made from slide preparations mounted

in water and Melzer’s reagent. The photographs of the asci, ascal apical ring, and

ascospores were taken by using a VHX-600E microscope of the Keyence Corporation.

The photographs of stromatal surface were taken with a ZSA30w microscope and

S70 Canon camera. The studied specimen is deposited at the Herbarium of Kunming

Institute of Botany, Chinese Academy of Sciences (HKAS).

Taxonomy

Xylaria atroglobosa H.X. Ma, Lar.N. Vassiljeva & Yu Li, sp. nov. PL. 1-6

MycoBank MB 563468

A X. fraseri structura stromatarum et magnitudine ascosporarum, a X. atrosphaerica

stromatis et ascosporis majoribus, ascosporae appendicula rotundata, hyalina, noncellulari

praeditae, differt.

Type: China, Yunnan Province, Xishuangbanna Tropical Botanical Garden, on rotten

wood, 9 Aug 1991, Peigui Liu (Holotype, HKAS 24003).

Erymo toy: Refers to the color and shape of the stroma.

STROMA hemispherical to depressed-spherical, attached to substrate with

narrow connective, 3-6 mm high x 0.6-1.2 cm diam., externally black,

internally white, woody, texture hard, surface smooth and perithecial mounds

inconspicuous; perithecia embedded in stromata, spherical, 0.5-0.8 mm diam.;

ostioles papillate. Asci 8-spored, cylindrical, long-stipitate, the spore-bearing

part 150-170 um long, with apical ring bluing in Melzer’s iodine reagent,

urn-shaped, 4.5-5 x 3.5-5 um. Ascospores brown, unicellular, ellipsoid-

inequilateral to crescent-shaped to navicular, with broadly to narrowly rounded

ends, bearing a round hyaline noncellular appendage up to 6-7 x 3-4 um at the

one end, smooth, (24-)24.5-27(-29) x 7.5-9 um, with oblique germ slit much

less than spore-length.

ComMeEnts- Xylaria atroglobosa is similar to X. fraseri M.A. Whalley et al.

(Whalley et al. 2000) in stromatal morphology and ascospore size, although

its ascospores are slightly shorter. However, the stromata of X. atroglobosa

lack white scales or a peeling layer at the surface, whereas stromata of

X. fraseri are overlaid with a white to yellowish white peeling layer cracked

into 0.2-0.4 mm broad scales. Furthermore, the ascospores of X. fraseri lack

an appendage and their germ slit is straight.

In stromatal morphology, the new species somewhat resembles X. atro-

sphaerica (Cooke & Massee) Callan & J.D. Rogers, which can be differentiated

Xylaria atroglobosa sp. nov. (China) ... 383

Fics. 1-6. Xylaria atroglobosa: 1—Stroma. 2—Stromatal surface. 3—Ascospores. 4—Germ

slit. 5—Ascospores bearing appendage. 6—Ascal apical ring. Scale bars: 1 = 2 mm, 2 = 0.25

um, 3 = 24 um, 4 = 13.5 um, 5 = 16.5 um, 6 = 8 um

by its smaller stromata (Callan & Rogers 1990) and smaller ascospores (18-20

x 6-7 um) that lack an appendage.

Acknowledgments

We greatly appreciate Dr. Adriana Hladki (Fundacién Miguel Lillo), Dr. Xiuguo

Zhang (Shandong Agriculture University) and Dr. A.J.S. Whalley (Liverpool John

384 ... Ma, Vasilyeva & Li

Moores University) for reviewing the presubmitted manuscript and giving helpful

comments and suggestions. This study was supported by the National Natural Science

Foundation of China (grant no. 30770005). We are also grateful to the Ministry of

Agriculture of China for funding the field study in the project entitled “The Project of

System Construction of Modern Agricultural Technology.’ We thank Dr. Pei-gui Liu

(Kunming Institute of Botany, Chinese Academy of Sciences) who collected the fungus.

We thank Dr. Hua Peng and Dr. Li-song Wang (Kunming Institute of Botany, Chinese

Academy of Sciences), who lent us the specimen.

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Malaysian Fungal Diversity. University of Malaya and Ministry of Natural Resources Malaysia.

Whalley MA, Ju YM, Rogers JD, Whalley AJS. 2000. New xylariaceous fungi from Malaysia.

Mycotaxon 74: 135-140.

MY COTAXON

http://dx.doi.org/10.5248/119.385

Volume 119, pp. 385-389 January-March 2012

A new name for a Coltricia (Basidiomycota) from India

JULIANO M. BALTAZAR* & ROSA MARA B. DA SILVEIRA

Programa de Pés-Graduacao em Botanica, Departamento de Botanica,

Universidade Federal do Rio Grande do Sul,

Av. Bento Goncalves 9500, CEP 91501-970, Porto Alegre, RS, Brazil

* CORRESPONDENCE TO: baltazar.jm@hotmail.com

ABSTRACT — Examination of the isotype of a Coltricia species described from India confirms

it as a good morphological species. A new name, Coltricia indica, is proposed to replace its

two previous illegitimate names. The species is characterized by very small, centrally stipitate

basidiomes with shiny pilei when dried, lobed to serrate and ciliate margins, and oblong-

ellipsoid, more or less thick-walled basidiospores.

Key worps — Coltricia parvula, Coltricia pusilla, Hymenochaetales, Hymenochaetaceae,

polypore

Introduction

Sharma & Wright (1992) described a new species Coltricia pusilla based

on two collections of a small polypore found in two forests in Khasi Hills,

Meghalaya State, India. However, their binomial was a later homonym of a

name previously used by Imazeki & Kobayashi (1966) for a new species from

Japan.

When J.E. Wright later became aware of Imazeki & Kobayashi’s work, he

proposed Coltricia parvula as a nom. nov. for the Indian species (Wright 1997).

Unfortunately, this was also a later homonym of a combination previously

published by Murrill (1904).

Our goal was to confirm the identity of Sharma & Wright's species and

provide a legitimate name for it.

Materials & methods

The isotype of Coltricia pusilla J.R. Sharma & J.E. Wright was studied during a visit

of the senior author to BAFC. Basidiomes were cut by hand for microscopic study,

and sections were mounted in 3% KOH, water and Melzer’s reagent (IKI). Herbaria

acronyms follow Thiers (2011).

386 ... Baltazar & Silveira

Taxonomy

Coltricia indica Baltazar, nom. nov. PLATE 1

MycoBank MB 563260

= Coltricia pusilla J.R. Sharma & J.E. Wright, Bull. Bot. Surv. India 31:

182, 1992 [“1989”], nom. illeg., non Imazeki & Kobayasi 1966.

= Coltricia parvula J.R. Sharma & J.E. Wright, in Wright, Mycotaxon

61: 419, 1997, nom. illeg., non (Klotzsch) Murrill 1904.

EryMo_oey: the new epithet refers to the type locality in India.

ORIGINAL DESCRIPTION (from Sharma & Wright 1992 with our additions within

square brackets) — “FRuITBODy annual, centrally stipitate, coriaceous soft

when fresh, hard and rigid on drying. PrLeus infundibuliform or depressed at

centre, circular, 5-8 mm diam., 1-1.5 mm thick, finely depressed velutinate [to

glabrous when dry], deep golden brown to cinnamon brown, shiny to glossy,

narrowly concentrically zonate. MARGIN sharp, deflexed when dry, ciliate [lobed

to serrate]. PORE SURFACE rusty brown, with a narrow sterile margin. PORES

angular, 2-4 [2-3] per mm, entire, becoming uneven near stipe with age. TUBES

< 1 mm deep, light golden brown to ochraceous brown. CONTEXT < 0.5 mm

thick, rusty brown, fibrous. STIPE equal or tapering upwards, usually swollen

at base, < 2 cm x 1-1.5 mm, finely tomentose to more velutinate towards base,

light rusty brown to dark brown [with a soft, loose discoid or globose base

enclosing some substrate].

“HYPHAL SYSTEM monomitic, generative hyphae with simple septa, branched

at broad [acute] angles, golden brown, 4-7 um wide. Spores oblong-ellipsoid

[to ellipsoid, rarely with a tapering apex becoming almost navicular], smooth,

slightly [to distinctly] thick-walled, [with a distinct apiculus,] golden brown

[in KOH, lighter in water], inamyloid, [nondextrinoid,] 5-6 x 3-4.2 um [(5-)

5.5-6(-7) x 3-4 um]?

SPECIMEN EXAMINED: INDIA. MEGHALAyYA, Khasi hills, Raliang forest, c. 1900 m [the

protologue citation of “6500 m” is apparently an error for 6500 ft; the highest altitude in

Meghalaya State is 1961 m (= 6434 ft), in the Khasi Hills], 25.[X.1986, on ground, leg.

J.R. Sharma 60108 (BAFC isotype).

DISTRIBUTION: known only from two forests in the Khasi Hills, Meghalaya

State, India.

Discussion

Coltricia indica is characterized by its very small, centrally stipitate

basidiomes and a circular infundibuliform pileus with a lobed to serrated and

ciliate margin. The stipe base is also noteworthy in that the isotype basidiomes

have enlarged bases with soft and loose tissue enclosing a piece of substrate or

forming a discus.

This species is very similar to C. minor Y.C. Dai, which also has very small

basidiomes (< 0.5 cm long) and basidiospores almost identical in shape and

Coltricia summa nom. nov. (India) ... 387

8000008 |

OOOOH

PLATE 1. Coltricia indica (isotype).

A. Upper (left) and lateral (right) views of the pileus. B. Drawing of two basidiomes.

C. Basidiospores. Scale bars = 0.5 cm (A-B), 10 um (C).

size. However, they differ both macroscopically and ecologically. Coltricia

minor has laterally stipitate basidiomes, fan-shaped to spathulate pilei, non-

ciliate margins, and regular stipe bases. Furthermore, C. minor was collected on

angiosperm wood and is thus far known only from Hunan Province (China),

which has a subtropical climate. Coltricia indica, on the other hand, was found

growing on the ground of coniferous forests in India in a temperate climate due

the altitude (Sharma & Wright 1992, Dai et al. 2010).

Coltricia barbata Ryvarden & de Meijer and C. velutina Baltazar & Gibertoni,

known from Brazilian Atlantic Forest areas, also have ciliate margins and

small basidiomes — pilei measuring < 1.8 cm and < 2.8 cm diam., respectively

(Meijer 2006, Baltazar et al. 2010). However, C. barbata has basidiospores

388 ... Baltazar & Silveira

that are globose to subglobose and 5-6 x 5-6 um, compared to subglobose to

broadly ellipsoid and 5.5-6.5(-7) x (4-)4.5-5.5(-6) um in C. velutina. Both

species also have smaller pores: 4-9 per mm in C. barbata and (4-)5-7 per mm

in C. velutina.

Within Hymenochaetaceae Imazeki & Toki, stipitate poroid representatives

with smooth basidiospores have been placed mainly in Coltricia Gray or

Phylloporia Murrill. Traditionally Phylloporia species have a duplex pileal

context with a black line below the tomentum, while Coltricia species have a

homogeneous context and usually larger basidiospores. Ecological aspects

were considered important but not decisive in circumscribing the two genera.

However, recent studies have shown that some Coltricia species [including the

type, Coltricia perennis (L.) Murrill] establish ectomycorrhizal associations

(Tedersoo et al. 2007). Furthermore, a phylogenetic analysis by Valenzuela et al.

(2011) supports Coltricia stuckertiana (Speg.) Rajchenb. & J.E. Wright within

the Phylloporia clade. Except for its basidiospores, C. stuckertiana displays

all other features typical of Coltricia. On the other hand, the type species of

Phylloporia (P. parasitica Murrill) has not yet been included in any phylogenetic

analysis. Its placement within the hymenochaetoid clade is needed for a reliable

classification of the two genera, particularly as P parasitica occurs in a very

specialized substrate, i.e., living angiosperm leaves.

Perhaps Coltricia will eventually be restricted to ectomycorrhizal species. If

so, we will only know the ectomycorrhizal status of C. indica by finding more

specimens. For the time being we retain the species in Coltricia based on its

morphology.

Based on its ornamented basidiospores, Coltricia pusilla Imazeki & Kobayashi

is currently treated as Coltriciella pusilla (Imazeki & Kobayashi) Corner (Corner

1991) [= Coltriciella pusilla (Imazeki & Kobayashi) J.E. Wright, comb. superfl.].

Coltricia parvula (Klotzsch) Murrill is accepted as a heterotypic synonym of

Coltricia cinnamomea (Jacq.) Murrill (Murrill 1908, Ryvarden 1976).

Acknowledgments

The authors would like to thank Dr. Sergio P. Gorjén (CIEFAP, Argentina) and

Dr. Gerardo L. Robledo (UNC, Argentina) for critically reviewing the manuscript.

MSc. Larissa Trierveiler Pereira made the drawings. The authors also thank Dra. Andrea

I. Romero (curator of BAFC-Fungi) and Lic. Susana Pereira (technician at BAFC-

Fungi) for kindly supporting JMB during his visit at BAFC. The ‘Conselho Nacional de

Pesquisa (CNPq - Brazil) provides a Ph.D. scholarship for JMB.

Literature cited

Baltazar JM, Ryvarden L, Gibertoni TB. 2010. The genus Coltricia in Brazil: new records and two

new species. Mycologia 102: 1253-1262. http://dx.doi.org/10.3852/09-227

Corner EJH. 1991. Ad Polyporaceas VII: The xanthochroic polypores. Beih. Nova Hedwigia

101-1175.

Coltricia summa nom. nov. (India) ... 389

Dai YC, Yuan H-S, Cui B-K. 2010. Coltricia (Basidiomycota, Hymenochaetaceae) in China. Sydowia

G2 141-2),

Imazeki R, Kobayashi Y. 1966. Notes on the genus Coltricia S.F. Gray. Trans. Mycol. Soc. Japan

7: 42-44,

Meijer AAR de. 2006. Preliminary list of the macromycetes from the Brazilian state of Parana. Bol.

Mus. Bot. Munic. 68: 1-55.

Murrill WA. 1904. The Polyporaceae of North America—VII. The genera Hexagona, Grifola,

Romellia, Coltricia and Coltriciella. Bull. Torrey Bot. Club 31: 325-348.

http://dx.doi.org/10.2307/2478798

Murrill WA. 1908. (Agaricales) Polyporaceae (conclusion). N. Amer. Fl. 9: 73-131.

Ryvarden L. 1976. Type-studies in the Polyporaceae 4. Species described by J.F. Klotzsch. Mem.

New York Bot. Gard. 28: 199-207.

Sharma JR, Wright JE. 1992 [“1989”]. A new species of polypores from India. Bull. Bot. Surv. India

31: 182-183.

Tedersoo L, Suvi T, Beaver K, Saar I. 2007. Ectomycorrhizas of Coltricia and Coltriciella

(Hymenochaetales, Basidiomycota) on Caesalpiniaceae, Dipterocarpaceae and Myrtaceae in

Seychelles. Mycol. Progr. 6: 101-107. http://dx.doi.org/10.1007/s11557-007-0530-4

Thiers B. 2011. Index Herbariorum: A global directory of public herbaria and associated staff. New

York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg. org/ih/ [accessed June

2011].

Valenzuela R, Raymundo T, Cifuentes J, Castillo G, Amalfi M, Decock C. 2011. Two undescribed

species of Phylloporia from Mexico based on morphological and phylogenetic evidence. Mycol.

Progr. 10: 341-349. http://dx.doi.org/10.1007/s11557-010-0707-0

Wagner T, Ryvarden L. 2002. Phylogeny and taxonomy of the genus Phylloporia (Hymenochaetales).

Mycol. Progr. 1: 105-116. http://dx.doi.org/ 10.1007/s11557-010-0707-0

Wright JE. 1997. A name change for Coltricia pusilla (Aphyllophorales). Mycotaxon 61: 419.

MY COTAXON

http://dx.doi.org/10.5248/119.391

Volume 119, pp. 391-395 January-March 2012

Tuber in China: T. microspermum and T. microspiculatum spp. nov.

Li FAN’, JIN-ZHONG Cao’, ZHAO-HUI ZHENG’, & YU LI”

' College of Life Science, Capital Normal University,

Xisanhuanbeilu 105, Haidian, Beijing 100048, China

? Institute of Mycology, Jilin Agricultural University, Changchun 130118, China

CORRESPONDENCE TO *: ' fanli@mail.cnu.edu.cn & * yuli966@126.com

ABSTRACT — Two new species, Tuber microspermum and T: microspiculatum, are described

from China. Tuber microspermum is characterized by small asci and spiny reticulate

ascospores; T: microspiculatum can be separated from the other species with spinoreticulate

ornamented ascospores by numerous minute meshes.

Key worps — Ascomycota, Tuberaceae, truffle

Introduction

China is especially rich in species of Tuber. Since the first publication on this

genus describing a new species from China, T. taiyuanense B. Liu (Liu 1985),

many previously described and new species have been published (Wang 1988;

Wang & Li 1991; Wang et al. 1998; Wang & He 2002; Tao et al. 1989; Chen et

al. 2005; Chen & Liu 2007; Fan et al. 2011). Because all Tuber species grow

underground, it can be very difficult to find specimens. Therefore, the number

of species in China is inevitably underestimated. After continuing investigation

of the genus in China, we describe and illustrate two additional new species.

Materials & methods

The materials were collected from the local mushroom market in Kunming, China.

The specimens are deposited in BJTC (Herbarium Biology Department, Capital Normal

University). Macroscopic characters were described from fresh specimens. Microscopic

characters were described from razor-blade sections of fresh specimens mounted in 3%

KOH; sections were stained with Melzer’s reagent, rinsed, and mounted in polyvinyl

lactic glycerol to make permanent slides for archiving with dried specimens. For

scanning electron microscopy (SEM), ascospores were scraped from the dried gleba

onto doubled-sided tape, which was mounted directly on an SEM stub, coated with

gold-palladium, and examined and photographed with a HITACHI S-4800 SEM.

392 ... Fan & al.

Taxonomy

Tuber microspermum L. Fan & J.Z. Cao, sp. nov. Fics. 1-4

MycoBank MB 561881

A yellow brown truffle with small asci and small spinoreticulate ascospores that differs

from Tuber pseudoexcavatum in an ascoma that lacks an excavated base.

Type: China. Yunnan Province, Kunming, from the local mushroom market. 20 Dec.

2010, Jin-zhong Cao 112 (Holotype, BJTC FAN149).

ETYMOLOGY: microspermum (Lat.), referring to the small ascospores.

AscoMaTA 1.5 cm diam., subglobose or slightly lobed, firm, solid, surface

smooth to partially minutely verrucose, glabrous, yellow brown at maturity.

Odor slight. PERIpruM 200-300 um thick, two layers; outer layer 80-120 um

thick, pseudoparenchymatous, composed of small subangular or subglobose

cells mostly 7.5-12.5 um in diam., with thickened and brown walls, darker

towards the outer surface; inner layer composed of intricately interwoven

hyphae, hyaline, thin-walled, branched, septate, 2.5-5 um in diam. GLEBA

brown to dark brown at maturity, marbled with numerous, narrow, branched,

white to white-yellow veins. Asc1 subglobose, ellipsoid or irregular, hyaline,

thick-walled (3-5 um), 45-70 x 40-60 um, mostly with a short stalk, 1-5

spores. Ascospores ellipsoid to broad ellipsoid, dark brown at maturity,

15-25(-27.5) x 12.5-20 um in 2-5-spored asci (30-35 x 22.5-25 um in

l-spored asci) excluding ornamentation; ornamentation distinctly spino-

reticulate, spines 4-6 um high, straight mostly, the meshes generally 3-6 across

the spore width.

ComMMENTS — Tuber microspermum is characterized by the small ascospores

and asci. In ascospore pattern the new species resembles T’ pseudoexcavatum,

which is easily distinguished by a different ascoma that is typically excavated

at the base (Wang et al. 1998). Other closely related Tuber species with spiny-

reticulate spores are T: taiyuanense and T. huidongense (Liu 1985; Wang & He

2002), which can be separated from T: microspermum by their larger ascospores

and asci.

Tuber microspiculatum L. Fan & Yu Li, sp. nov. Fics 5-8

MycoBaAnk MB 561882

Differs from all other Tuber species with spinoreticulate ascospores by an ornamentation

of numerous minute narrow spine-like meshes.

Type: China. Kunming, Yunnan Province, from the local mushroom market. 20 Dec.

2010, Jin-zhong Cao 101 (Holotype, BJTC FAN138).

ETyMOLOoGy: microspiculatum (Lat.), referring to the numerous minute meshes of the

spinoreticulate spore ornamentation.

ASCOMATA 2-2.5 cm in diam., globose or subglobose, firm, solid, with or

without distinct umbilicate depression at the base, surface smooth and

Tuber spp. nov. (China) ... 393

Fics 1-8. Tuber microspermum (BJTC FAN149, holotype): 1. Ascocarp; 2-3. Asci and ascospores

observed under light microscope; 4. Ascospore observed under SEM. Tuber microspiculatum

(BJTC FAN138, holotype): 5. Ascocarps; 6-7. Asci and ascospores observed under light

microscope; 8. Ascospore observed under SEM.

394 ... Fan & al.

glabrous, white yellow, pale yellow or light brown, reddish brown at maturity.

Odor slight, not pungent. PERrpr1um 200-250 um thick, two layers; outer

layer 50-100 um, pseudoparenchymatous, composed of small subangular

or subglobose cells, mostly 7.5-15 um in diam., with slightly thickened and

yellowish-brown walls, darker towards the outer surface; inner layer composed

of intricately interwoven hyphae, hyaline, thin-walled, branched, septate, 2.5-5

um in diam. GLEBA brown to nearly reddish brown at maturity, marbled with

numerous, narrow, branched, white-yellow veins radiating from the base. Asc1

subglobose, ellipsoid or irregular, 60-85 x 55-70 um, sessile or with a short

stalk, (1-)2—-4 spores. Ascosporgs ellipsoid, a few broad ellipsoid to subglobose,

light brown to yellow-brown at maturity, (20-)22.5-35(-40) x (15-)17.5-22.5

um in 2-—4-spored asci and 42.5-45 x 22.5-25 um in 1-spored asci excluding

ornamentation; ornamentation of spines 2.5-4(-5) um high, mostly connected

at bases by an alveolate reticulum, the meshes minute and numerous, generally

10-16 across the spore width, a few of large spores in 1-spored asci sometimes

with 4-8 meshes across the spore width.

Comments — Tuber microspiculatum differs from all related species with spiny-

reticulate ascospores by spore ornamentation with very minute and numerous

meshes, which are so narrow that they can be easily confused with the spines

under light microscope. SEM observation (Fic. 8) is helpful in clarifying the

characteristics.

Species of the T. rufum complex, which appear similar to T. microspiculatum

under the light microscope, can be separated by the presence of isolated spines

on the ascospore surface.

Acknowledgments

The study was supported by the National Natural Science Foundation of China

(Nos. 30770005, 30870008). We are grateful to Prof. Guo-Zhong Lt and Prof. Anthony

Whalley for reviewing the pre-submitted manuscript. We also thank Dr. Cheng-Lin

Hou for critically correcting the manuscript.

Literature cited

Chen J, Liu PG. 2007. Tuber latisporum sp. nov. and related taxa, based on morphology and DNA

sequence data. Mycologia 99: 475-481. http://dx.doi.org/10.3852/mycologia.99.3.475

Chen J, Liu PG, Wang Y. 2005. Tuber umbilicatum, a new species from China, with a key to the

spinose-reticulate spored Tuber species. Mycotaxon 94: 1-6.

Fan L, Cao JZ, Liu YY, Li Y. 2011. Two new species of the genus Tuber from China. Mycotaxon 116:

349-354. http://dx.doi.org/10.5248/116.349

Liu B. 1985. New species and new records of hypogeous fungi from China (I). Acta Mycologica

Sinica 4(2): 84-89.

Tao K, Liu B, Zhang DC. 1989. A new species of the genus Tuber from China. Journal of Shanxi

University (Nat. Sci. Ed.) 12: 215-218.

Tuber spp. nov. (China) ... 395

Wang Y. 1988. First report of study on Tuber species from China. Atti del II Congresso Internazionale

sul Tartufo. Spoleto, Nov 24-27: 45-50.

Wang Y, He XY. 2002. Tuber huidongense sp. nov. from China. Mycotaxon 83:191-194.

Wang Y, Li ZP. 1991. A new species of Tuber from China. Acta Mycologica Sinica 10: 263-265. (in

Chinese).

Wang Y, Moreno G. Riousset LJ, Manjon JL, Riousset G, Fourre G, Di Massimo G, Garcia- Montero

LG, Diez J. 1998. Tuber pseudoexcavatum sp. nov. a new species from China commercialised in

Spain, France and Italy with additional comments on Chinese truffles. Cryptogamie Mycologie

19: 113-120.

MY COTAXON

http://dx.doi.org/10.5248/119.397

Volume 119, pp. 397-403 January-March 2012

The first report of Ceriporia lacerata

(Phanerochaetaceae, Basidiomycota) in Korea

YEONGSEON JANG’, HA EUN CHol’, YOUNG WOON LIM’,

JIN SUNG LEE? & JAE-JIN K1m’*

‘Division of Environmental Science & Ecological Engineering,

College of Life Sciences & Biotechnology, Korea University

5-1 Anam-dong, Seongbuk-gu, Seoul, 136-701, Korea

School of Biological Sciences, Seoul National University, Seoul, 151-747, Korea

National Institute of Biological Resources, Environmental Research Complex,

Incheon, 404-708, Korea

*CORRESPONDENCE TO: jae-jinkim@korea.ac.kr

AxBsTRACT— An unrecorded Ceriporia species was collected during the field survey in Korea

University Experimental Forests. Based on morphological characteristics and 28S, 18S, and

internal transcribed spacer rDNA sequence analyses, the species was identified as Ceriporia

lacerata, new to South Korea. A key to the Korean Ceriporia species is provided.

Key worps— basidiomycete, polypore, taxonomy

Introduction

Ceriporia Donk is a genus of wood rotting fungi. It causes a white rot and

includes species with resupinate basidiocarps with white, red, purple, orange,

pink, or green pore surfaces, a monomitic hyphal system, simple septate or

rarely clamped generative hyphae, lack of cystidia or other sterile hymenial

elements, and subglobose to cylindric or allantoid basidiospores (Gilbertson

& Ryvarden 1986).

In Korea, previously three Ceriporia species have been reported (Lee & Jung

2005): Ceriporia purpurea (Fr.) Donk, C. reticulata (Hoffm.) Domanski, and

C. viridans (Berk. & Broome) Donk. During our studies on diversity of

indigenous fungi in Korea University Experimental Forests, two specimens of

an unreported Ceriporia species were found. They were first identified from

morphological characters, and sequence analyses of 28S, 18S and internal

transcribed spacer (ITS) rDNA region supported the identification. A detailed

description of the species and a key to the Korean Ceriporia species are provided

below.

398 ... Jang & al.

Materials & methods

Collections

The basidiocarps were collected in 2010 from one of the Korea University

Experimental Forests located in Goesan-gun, Chungcheongbuk-do (36°46'13"N

127°58'57-59"E). They were dried completely overnight using an air drier at 40°C and

deposited at the National Biological Resources Center (KB).

Microscopic observation

Macro- and microscopic basidiocarp features were noted from the collected specimens.

Measurements and drawings were made from slide preparations mounted in 3% KOH

(Largent et al. 1977) using an Olympus BX51 light microscope. Average dimensions

were determined from > 20 measurements of each character. The characteristics of the

four Korean Ceriporia listed in TABLE 1 are based on our observations of C. lacerata

and descriptions of C. purpurea, C. reticulata, and C. viridans from previous studies

(Jung 1994, Lim et al. 2000, Lee et al. 2002). A key to Korean Ceriporia species was

constructed.

TABLE 1. Morphological comparison of four Korean Ceriporia species.

@ieiisieieries C. lacerata C. purpurea C. reticulata C. viridans

(this paper) (Lim et al. 2000) (Jung 1994) (Lee et al. 2002)

Ration resupinate, resupinate, resupinate, resupinate,

effused, confluent effused, confluent ceraceous adnate

white, buff to pale to dark whitish to cream to sordid

PORE SURFACE : A

ochraceous brownish purple ochraceous pinkish brown

Pores (per mm) 2-5 4-5 2-4 3-5

BASIDIA (tm) 11-16.5 x 3.5-5.5 17-21 x 5.7-6.3 15-20 x 5-6 11-15 x 4.5-6

BASIDIOSPORES (tm) 3.5-5 x 2-3 5.7-6.5 X 1.8-2.3 6-8 x 2-3 4-5 x 1.5-2

Molecular analysis

Genomic DNAs were extracted from basidiocarp hymenophores using Accuprep

Genomic DNA extraction kit (Bioneer, Korea). PCR reactions were performed using

Accupower PCR premix kit for 28S, 185, and ITS rDNA, using primers LROR/LR3

(Vilgalys & Hester, 1990), EukNS20F/EukNS1750R (Dams et al. 1988), and ITSIF

(Gardes & Bruns 1993) /ITS4 (White et al. 1990), respectively. 18S PCR amplifications

followed Suhara et al. (2003). 28S and ITS rDNA region conditions were as follows:

initial denaturation at 95°C for 7 min, followed by 30 cycles at 95°C for 30 s, 51°C for 30

s, and 72°C for 30 s, concluded by elongation at 72°C for 7 min. DNA sequencing was

performed in the DNA Engine Tetrad 2 Peltier Thermal Cycler (BIO-RAD, USA) using

the ABI BigDye(R) Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems, USA)

with aforementioned primers and an ABI 3730XL DNA Analyzer (Applied Biosystems,

USA). The sequences obtained in this study were deposited under the GenBank accession

no. JN618335, JN618336, and JN641758-JN641761. Maximum parsimony (MP) and

neighbor joining (NJ) analyses of each region followed Jang et al. (2011).

Ceriporia lacerata new to Korea... 399

(B)

C0010

OoOd00

Fic. 1. Ceriporia lacerata. (A) Basidiocarp. (B) Microscopic features: a, basidiospores; b, basidia;

c, generative hyphae. Scale bar = 10um

Taxonomy

Ceriporia lacerata N. Maek., Suhara & R. Kondo, Mycotaxon 86: 342.2003. Fic. 1

Basidiocarp resupinate, effused, confluent, soft when fresh, then fragile,

firmly attached to the substrate; hymenophore poroid, white, buff to ochreous;

dissepiments entire to lacerate; pores angular, 2-5 per mm; margin white.

Hyphal system monomitic; contextual hyphae 3-5.7 um in diameter, smooth,

thin- to slightly thick-walled, simple-septate; tramal hyphae 2.3-4.6 um in

diameter, smooth, thin- to slightly thick-walled, simple-septate, cystidia

lacking; basidia clavate, 11-16.5 x 3.5-5.5 um, with 4 sterigmata; basidiospores

oblong-ellipsoid to ellipsoid, 3.5-5 x 2-3 um (ave. 3.9 + 0.2 x 2.5 + 0.4 um, n=

25), smooth, thin-walled.

SPECIMENS EXAMINED: KOREA, CHUNGCHEONGBUK-DO, Goesan-gun, Korea University

Experimental Forest, 36°46'13"N 127°58'57-59"E, fallen branches of Quercus sp. 07

July 2010, Yeongseon Jang (KB NIBRFG0000114705, GenBank JN618336, JN641758,

JN641760; KB NIBRFG0000114712, GenBank JN618335, JN641759, JN641761).

ECOLOGY & DISTRIBUTION — China, Korea, and Japan on gymnosperms and

angiosperms.

REMARKS — The characters generally match those of the holotype. However,

basidiospore sizes were more variable in the Korean specimens. Suhara et al.

(2003) cited 4.25-5 x 2.5-2.75 um (ave. 4.5 + 0.3 x 2.6 + 0.1 um, n = 20) for the

holotype basidiospores, whereas those from our specimens measured 3.5-5 x

2-3 um (ave. 3.9 + 0.2 x 2.5 + 0.4 um, n = 25).

Phylogeny

MP and NJ analyses of the 28S, 18S, and ITS rDNA regions were conducted

to confirm the morphological identification. The aligned dataset of 11 sequences

400 ... Jang & al.

Ceriporia lacerata NIBRFG0000114705

62/66 | Ceriporia lacerata NIBRFG00001 14712

seartoo Ceriporia lacerata (AY858357)

Ceriporia lacerata (AB566280)

Ceriporia lacerata (FJ471540)

Ceriporia purpurea (AF287852)

Ceriporia tarda (GQ470632)

Ceriporia viridans (AF347109)

Ceriporia reticulata (EU118614)

Ceriporia viridans (AF393049)

Ceriporiopsis aneirina (FJ496704)

—_————= 10 substitution/site

Ceriporia lacerata NIBRFG0000114705

Ceriporia facerata NIBRFG0000114712

Ceriporia lacerata (AB084585)

Ceriporia spissa (AB084588)

Ceriporia reticulata (AB084587)

Ceriporia tarda (AB084591)

Ceriporia purpurea (AB084586)

Ceriporia viridans (AB084592)

Ceriporiopsis aneirina (AB084589)

Ceriporiopsis pannocincta (AB084590)

10 substitution/site

(ss) Ceriporia lacerata NIBRFG00001 14705

Ceriporia lacerata NIBRFG0000114712

Ceriporia lacerata (DQ912694)

400/100

Ceriporia lacerata (AB091675)

Ceriporia lacerata (HM008936)

Ceriporia spissa (GU594154)

Ceriporia spissa (GU594155)

Ceriporia reticulata (EU118613)

Ceriporiopsis aneirina (AY219362)

—— 10 substitution/site

Fic. 2. 28S, 188, and ITS rDNA region maximum parsimonious trees of Ceriporia lacerata and

allied species. The trees were rooted by Ceriporiopsis species. Bootstrap values = 50% are shown

above branches (MP bootstrap proportions/NJ bootstrap proportions). Specimens found in this

study are in bold. GenBank accession numbers in parentheses.

of 28S rDNA comprised 504 characters, of which 435 sites were constant, 24

sites variable, and 45 sites parsimony informative; tree length was 90 steps

with consistency index (CI) = 0.8222 and retention index (RI) = 0.8841. The

18S rDNA dataset of 12 sequences comprised 1645 characters, of which 1578

were constant, 41 variable, and 26 parsimony informative; tree length was 79

steps with CI = 0.8608 and RI = 0.8226. The ITS rDNA dataset of 9 sequences

comprised 390 characters with 281 constant, 44 variable, and 65 parsimony

informative sites; tree length was 138 steps with CI = 0.9855 and RI = 0.9802. The

MP and NJ analyses of all three regions clustered our sequences with previously

reported sequences of Ceriporia lacerata with high bootstrap supports (Fic. 2).

The Korean C. lacerata (KB NIBRFG0000114705, KB NIBRFG0000114712)

showed 100% sequence similarity to a Japanese specimen (GenBank

AB084585) in 18S rDNA region according to a BLAST search of GenBank. ITS

rDNA region also showed 99.8% similarity (one out of 560 positions showed

nucleotide difference) to a Japanese specimen (GenBank AB091675). No 28S

Ceriporia lacerata new to Korea... 401

rDNA sequences of a Japanese specimen were available, but our specimens had

100% similarity to C. lacerata strains of Korea and Japan.

Discussion

Ceriporia lacerata was first described as a new species from Japan in 2003

(Suhara et al. 2003) and three years later reported from China (Cui et al. 2006,

Dai 2012). In Korea, C. lacerata strains have been isolated from playground

wood (Kim et al. 2005), CCA-treated wood (Kim et al. 2007), decayed pine

tree logs (Kim et al. 2009), and decayed bamboo (Kim et al. 2011). They

were also isolated from Mt. Bukhan (Seoul), Mt. Halla (Jeju), Mt. Joryeong

(Chungcheongbuk-do), Mt. Mani (Incheon), Mt. Yongmun (Gyeonggi-do),

and Yangpyeong Korea University Experimental Forest (Gyeonggi-do) in

2008-10 during the Korean indigenous species research project (unpublished

data) and basidiocarps were found in Goesan-gun, Chungcheongbuk-do, in

2010. Ceriporia lacerata occurs on both angiosperm and gymnosperm wood

and is widely distributed in China (Dai 2012). Our observation confirms

C. lacerata within northeast Asia, and it is reasonable to infer that it inhabits

other Asian countries in the 30-45°N latitude.

Morphologically, Ceriporia lacerata is similar to C. alachuana (Murrill)

Hallenb. and C. ferruginicincta (Murrill) Ryvarden by having a whitish to

ochreous pore surface and oblong to ellipsoid basidiospores. However, the other

two species differ in having pores with entire dissepiments (3-8 per mm in

C. alachuana and 6-8 per mm in C. ferruginicincta; Suhara et al. 2003),

contrasting with the entire to lacerate dissepiments (2-5 per mm) of

C. lacerata.

The 28S and ITS rDNA phylogenetic trees had similar topologies with

Ceriporia lacerata forming a clade distinct from that of the other Ceriporia

species (Fic. 2). On the other hand, all the analyzed Ceriporia species were

monophyletic in 18S rDNA tree, as previously suggested by Wu et al. (2010),

with C. spissa (Schwein.) Rajchenb. sister to C. lacerata. Both species occur

on hardwoods and conifers, but C. spissa has bright orange basidiocarps

(Gilbertson & Ryvarden 1986). The phylogenetic conflict represented here is

probably the result of limited taxon sampling. More molecular data derived

from other species are needed for the clear elucidation.

Since Kondo et al. (1999) reported that Ceriporia lacerata had the ability

to degrade polychlorinated dibenzo-p-dioxines (PCDDs) and polychlorinated

dibenzo-furans (PCDFs), its industrial potential has been investigated in

pretreatment of lignocellulosic biomass for efficient hydrolysis and biofuel

production (Lee et al. 2007) and PAHs degradation (Lee et al. 2010). Remarkably,

Long et al. (2010) reported that C. lacerata had the highest decolorization ability

for synthetic dyes (Congo Red, Orange II, and Reactive Blue 4) among the

402 ... Jang & al.

white rot fungal species tested in solid media. However, information regarding

this species is still limited and the abilities of this species should be investigated

further.

Key to the species of Ceriporia in Korea

1. Pore surface pale to dark brownish purple;

(basidiospores allantoid, 5-7 um long) ............ 0.0... eee ee eee C. purpurea

1 APOE SuTtACe WITS LO DEO Wale en, cena ten Neg Po, Ao as Pencil ans dla acd dle a tle atthe caehsiidlg nda 2

2. Pore surface cream to sordid pinkish-brown;

(basidiospores cylindric to allantoid, 4-5 um long)................. C. viridans

2POre Surtace white to-OchTace ous cs HG aes lee x he Ye Ee eG ote eee 3

3. Basidiospores allantoid, 6-8 x 2-3 UM ........ eee eee eee eee C. reticulata

3. Basidiospores oblong-ellipsoid to ellipsoid, 3.5-5 x 2-3 um............ C. lacerata

Acknowledgements

This work was carried out with the support of ‘Forest Science & Technology Projects

(Project No. $121010L110000) provided by Korea Forest Service. This work was also

supported by the Korean indigenous species research project from the National Institute

of Biological Resources (NIBR), Ministry of Environment, Republic of Korea, and by a

Korea University Grant. We are grateful to Dr. Yu-Cheng Dai and Dr. Kentaro Hosaka

for their valuable suggestions on the manuscript.

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Dai YC. 2012. Polypore diversity in China with an annotated checklist of Chinese polypores.

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Dams E, Hendriks L, Van de Peer Y, Neefs JM, Smits G, Vandenbempt I, De Wachter R. 1988.

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to the identification of mycorrhizae and rusts. Mol. Ecol. 2: 113-118.

http:/dx.doi.org/10.1111/j.1365-294X.1993.tb00005.x

Gilbertson RL, Ryvarden L. 1986. North American polypores, Vol. 1. Fungiflora, Oslo.

Jang YS, Choi HE, Lim YW, Lee JS, Kim J-J. 2011. The first report of Antrodia sitchensis (Polyporaceae,

Basidiomycota) in Korea. Mycobiology 39: 226-229.

http:/dx.doi.org/10.5941/MYCO.2011.39.3.226

Jung HS. 1994. Floral studies on Korean wood-rotting fungi (II) - on the flora of the Aphyllophorales

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Kim G-H, Lim YW, Song Y-S, Kim J-J. 2005. Decay fungi from playground wood products in

service using LSU rDNA sequence analysis. Holzforschung 59: 459-466.

http:/dx.doi.org/10.1515/HE2005.076

Kim G-H, Lim YW, Choi Y-S, Kim M-J, Kim J-J. 2009. Primary and secondary decay fungi on

exposed pine tree logs in the forest. Holzforschung 63: 633-638.

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Kim J-J, Kang S-M, Choi Y-S, Kim G-H. 2007. Microfungi potentially disfiguring CCA-treated

wood. Int. Biodeter. Biodegr. 60: 197-201. http:/dx.doi.org/10.1016/j.ibiod.2007.05.002

Kim J-J, Lee S-S, Ra J-B, Lee H, Huh N, Kim G-H. 2011. Fungi associated with bamboo and their

decay capabilities. Holzforschung 61: 271-275. http:/dx.doi.org/10.1515/HE.2011.004

Kondo R, Mori T, Yoshimitu M, Hirose M, Wakao K, Matufuji Y, Sakai K. 1999. 10th Haikibutugakkai

Youshishuu: 877-879. [In Japanese]

Largent DL, Johnson D, Watling R. 1977. How to identify mushrooms to genus, HI. Microscopic

features. Mad River Press, Eureka, CA, USA.

Lee H, Choi Y-S, Kim M-J, Huh N-Y, Kim G-H, Lim YW, Kang S-M, Cho S-T, Kim J-J. 2010.

Degrading ability of oligocyclic aromates by Phanerochaete sordida selected via screening of

white rot fungi. Folia Microbiol. 55: 447-453. http:/dx.doi.org/10.1007/s12223-010-0075-9

Lee JS, Jung HS. 2005. List of recorded Korean Aphyllophorales. Kor. J. Mycol. 33: 38-53.

Lee JS, Kim KM, Jung HS. 2002. The Aphyllophorales of the Kyeryoungsan National Park.

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Lee J-W, Gwak K-S, Park J-Y, Park M-J, Choi D-H, Kwon M, Choi I-G. 2007. Biological pretreatment

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Lim YW, Kim YH, Jung HS. 2000. The Aphyllophorales of Mungyong Saejae. Mycobiology

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Abstr. B027, p. 194, in: Proc. International Meeting of the Federation of Korean Microbiological

Societies.

Suhara H, Maekawa N, Kaneko S, Hattori T, Sakai K, Kondo R. 2003. A new species, Ceriporia

lacerata, isolated from white-rotted wood, Mycotaxon 86: 335-347.

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MY COTAXON

http://dx.doi.org/10.5248/119.405

Volume 119, pp. 405-411 January-March 2012

Marasmioid and gymnopoid fungi of the Republic of Korea. 5.

Marasmius sect. Hygrometrici

VLADIMiR ANTONIN’, RHIM RyYOO?,

KANG-HYEON Ka? & HYEON-DONG SHIN?

' Moravian Museum, Dept. of Botany, Zelny trh 6, CZ - 659 37 Brno, Czech Republic

? Korea Forest Research Institute, Department of Forest Resources Utilization,

Seoul 130-712, Republic of Korea

> Korea University, Division of Environmental Science and Ecological Engineering,

Seoul 136-701, Republic of Korea

* CO-CORRESPONDENCE TO: | vantonin@mzm.cz &* hdshin@korea.ac.kr

ABSTRACT — Two species of Marasmius sect. Hygrometrici from South Korea are published

in this paper. Descriptions based on morphological characteristics and illustrations are

provided. Marasmius crescentiae is a new record for the studied region, and “M. junipericola”

represents a provisional new species.

Key worps — Fungi, Agaricales, Marasmiaceae, taxonomy, DNA studies

Introduction

This new paper in a series on marasmioid and gymnopoid fungi collected

in the Republic of Korea (South Korea, Antonin et al. 2009, 2010a,b,c, 2011)

covers species of sect. Hygrometrici. A provisional new taxon, “Marasmius

junipericola” is described based on its very distinct macro- and microscopic

characteristics. However, we do not formally publish a new name because there

was too little material for molecular sampling. Marasmius crescentiae is recorded

as new to Korea. We provide ITS and LSU sequences of three specimens of

M. crescentiae.

Materials & methods

Macroscopic descriptions of collected specimens are based on fresh basidiocarps

collected by the first author. Colour abbreviations follow Kornerup & Wanscher (1983),

herbarium abbreviations are according to Holmgren & Holmgren (1998). Authors of

fungal names are cited according to the International Plant Names Index Authors website

(http://www.ipni.org/ipni/authorsearchpage.do). Microscopic features are described

A406 ... Antonin & al.

from dried material mounted in H,O, KOH, Melzer’s reagent, and Congo Red using

an Olympus BX-50 light microscope with a magnification of 1000x. Abbreviations:

L = number of entire lamellae; | = number of lamellulae tiers between each pair of entire

lamellae; E = quotient of length and width in any one basidiospore; Q = mean of

basidiospore E-values. Specimens are preserved in the herbarium of the Moravian

Museum, Brno, Czech Republic (BRNM).

DNA extraction, PCR amplification, DNA sequencing, and phylogeny tree

construction methodologies followed Antonin et al. (2010a). New GenBank accession

numbers are listed in TABLE 1. The Bayesian-based ITS phylogeny used the general

time reversible (GTR) model with gamma-distributed substitution rates (Ronquist &

Huelsenbeck 2003). Markov chains were run for 1,000,000 generations, saving one tree

every 100th generation and discarding the first 1000 trees. This MCMC tree was used

to compute a 50% majority rule consensus of the remaining trees to obtain estimates

for the posterior probabilities (PPs) of the groups. The dataset includes M. crescentiae

and M. micraster (both from sect. Hygrometrici), with the type species of Marasmius,

M. rotula, selected as outgroup.

TABLE 1. Marasmius specimens sequenced for this study.

SPECIES SPECIMEN ID ACCESSION NUMBER

Results

Marasmius crescentiae Murrill, North American Fl. 9: 259, 1915. Fics 1-2

BASIDIOCARPS single. PILEUS 3-5 mm broad, broadly conical to convex with

slightly depressed centre and straight margin, sometimes with slightly depressed

centre (when old) centre rugulose, sulcate otherwise, margin crenulate, surface

tomentose, light brown to brown (+ 6-7D-E6-7, 7E6), paler (brownish orange,

+ 6B-C5-6) at margin. LAMELLAE distant, L = 9-15,1=0(-1), shortly to broadly

adnate, slightly intervenose when old, pale cream coloured, concolorous or

with coloured (+ pileus colour, especially in external part), pubescent edge.

STIPE 20-65 mm long, filiform, insititious, smooth, glabrous, concolorous with

lamellae at apex, (reddish) black-brown (8E-F7) otherwise.

BASIDIOSPORES 6.0-9.0 x 3.0-4.5 um, average = 7.6 x 3.7 um, E = 1.6-3.0, Q

= 1.8-2.6, ellipsoid-fusoid, fusoid or sublacrimoid, thin-walled. Basrp1a 17-26

x 7.0-9.0 um, 4-spored, (broadly) clavate. BASIDIOLEsS 12-23 x 4.0-9.0 um,

clavate, fusoid. CHEILOCYSTIDIA of two types: [1] (10-)15-20(-23) x (4.5-)

6.0-13 um, broom cells of the Rotalis-type, broadly clavate, pyriform, thin-

walled, often with brown walls in KOH; [2] 15-30 x 4.5-9.0 um, lageniform or

fusoid, rostrate, smooth or with scattered projections at upper part of a base,

Marasmius sect. Hygrometrici in Korea ... 407

Fic. 1. Marasmius crescentiae.

Basidiocarps (Hongcheon, Nae-myeon, Sambong-ri, V. Antonin 08.26, BRNM 718666).

obtuse to subacute, thin-walled. PLEUROcystTip14 absent or rare, similar to

cheilocystidia of the type 2, 17-35 x 4.5-9.0 um, fusoid or lageniform, rostrate,

thin-walled. TRAMA HYPHAE + cylindrical, thin- to slightly thick-walled, hyaline,

up to 12 um wide. PILEIPELLIS a hymeniderm composed of (8.0—)10-21 x

7.0-14(-17) um, clavate, pyriform or (sub) vesiculose cells of the Rotalis-type,

thin-walledatbase, thin-toslightlythick-walledatapex;diverticulaup to 1.50.75

um, numerous; thick-walled parts brown in KOH. Piteocystip1a (12-)15-25

(-30) x 4.0-8.0 um, fusoid or lageniform, rostrate, thin- to slightly thick-walled

(especially in central part), sometimes with projections in upper basal part; with

chestnut brown wall in thick-walled parts. STIPITIPELLIs a cutis of cylindrical,

parallel, slightly thick-walled, diverticulate, up to 5.0 um wide hyphae with

brown walls in KOH. CauLocystip1a absent. CLAMP CONNECTIONS present

in all tissues.

CHEMICAL REACTIONS — No part of basidiocarps amyloid or dextrinoid.

EcoLocy — On fallen leaves of Acer, Quercus acutissima, Juglans, and other

broadleaved trees, dead plant stems.

SPECIMENS EXAMINED — REPUBLIC OF KOREA, Hongcheon, Nae-myeon, Eulsudong

valley, 37°41'11"N 128°29'35"E, alt. c. 670 m, 26 July 2007 leg. V. Antonin (07.131) and

R. Ryoo (BRNM 714783); Deogyusan National Park, Cheon-yeon falls, 24 Aug. 2007 leg.

R. Ryoo KG 164 (BRNM 714678); Hongcheon, Nae-myeon, Sambong-ri, 26 June 2008

A408 ... Antonin & al.

Fic. 2. Marasmius crescentiae.

a. Pileipellis cells, b. basidiospores, c. pileocystidia,

d. cheilocystidia (type 2) and pleurocystidia, e. cheilocystidia (type 1). Scale bar = 20 um.

leg. V. Antonin (08.25, 08.26) and R. Ryoo (BRNM 718665, 718666); Wonju, Chiaksan

National Park, between N.P. office and Guryongsa, 37°24'17"N 128°03'00"E, alt.

310-360 m, 19 July 2009 leg. V. Antonin (09.135) and R. Ryoo (BRNM 718790).

REMARKS — The South Korean collections of M. crescentiae are characterized

by small basidiocarps, a sulcate light brown to brownish orange pileus, rather

large basidiospores, tetrasporic basidia, cheilocystidia of two types, absent or

scattered pleurocystidia, present pileocystidia, absent caulocystidia, and present

clamp connections. Although Singer (1976) observed smaller basidiospores

(6.5 x 2.7 um), Desjardin & Horak (1997) cited larger basidiospores (7.5-8.5 x

3.5-4.0 um).

Very similar species are M. micraster Petch, with slightly larger basidiospores

(Desjardin & Horak 1997: 9-11(-12) x (3.5-)4-5(-5.5) um; Pegler 1986:

8.5-11 x 4.0-5.5 um) and 18-30 um long basidia (Singer 1976), and M. exustus

Marasmius sect. Hygrometrici in Korea ... 409

Berk. & M.A. Curtis, with smaller (6.5-8.0 x 4-5 um) basidiospores (Desjardin

& Horak 1997). Other species differ either by macroscopic characters (e.g.,

M. corbariensis (Roum.) Sacc.; Antonin & Noordeloos 2010) or by developed

caulocystidia (e.g., M. minutus Peck; Antonin & Noordeloos 2010).

Three sequences obtained from collections VA 07.131, KG 164 and VA 09.135

(BRNM 714783, 714678 and 718790; TABLE 1) of Marasmius crescentiae showed

a significant ITS rDNA similarity (Fic. 3). Among LSU sequences trimmed by

1167 nucleotide characters, the three sequences showed no more than five base

substitutions with each other.

0.54 , Marasmius crescentiae JN003840

1.00 || M. crescentiae FJ936142

M. crescentiae FJ936141

M. micraster FJ431258

M. micraster FJ431260

M. micraster FJ431261

M. micraster FJ431259

M. rotula DQ182506

0.1

Fic. 3. Phylogenetic tree for Marasmius sect. Hygrometrici based on the ITS region (ITS1+5.8S

rDNA+ITS2). Bayesian analysis running 1,000,000 generations represent mean branch lengths of

a 50% majority-rule consensus tree calculated from 1000 trees revealed. Scale bar = number of

nucleotide substitutions per site. Bold face = Species sequenced for this study.

“Marasmius junipericola” ad int. Fic. 4

BASIDIOCARPS single. PILEUS 1-1.5 mm broad, low convex to convex with

obtusely applanate then slightly depressed centre, + smooth when young, then

slightly crenulate towards margin, covered with small brown granulose + spiny

scales, entirely brown (7E7) when young, later brown only at centre and whitish

towards margin. LAMELLAE distant, L = 8-10 , 1 = 0-1, free or with small tooth,

white, with concolorous, slightly pubescent edge. Stipe 4-8 mm long, filiform,

insititious, entirely finely strigose-pubescent, dark brown (6-7E-F7), paler at

apex.

BASIDIOSPORES (7.0-)8.0-9.5 x (4.2-)4.5-5.0 um, average = 8.5 x 4.7 um,

E = 1.5-1.9, Q = 1.8, broadly ellipsoid, fusoid-ellipsoid, hyaline, thin-walled.

Basip1a 18-24 x 5.5-10 um, 4-spored, clavate. BASIDIOLES up to 22 x 5.0-9.0

um, clavate, fusoid, subcylindrical. CHErLocystip1A of two types, [1] 15-21

x 8.0-12 um, broom cells of the Rotalis-type, broadly clavate, thin-walled;

[2] 23-35 x 5.0-9.0 um, lageniform or fusoid, rostrate, obtuse, thin-walled.

410... Antonin & al.

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Fic. 4. “Marasmius junipericola”.

a. Pileipellis cells, b. pileocystidia, c. cheilocystidia (type 2),

d. cheilocystidia (type 1), e. basidiospores. Scale bar = 20 um.

PLEUROCYSTIDIA rare, similar to cheilocystidia of type 2. TRAMA AND CONTEXT

HYPHAE + cylindrical, branched, thin- to slightly thick-walled, smooth or

minutely incrusted, up to 8.0 um wide. PILEIPELLIS a hymeniderm composed

of 16-28 x 9.0-16 um, (broadly) clavate, vesiculose cells of the Rotalis-type,

entirely thin-walled or with slightly thick-walled apex; mixed with scattered

cells with larger diverticula or diverticulate hyphae. PILEocystTip1A 21-25 x

5.0-6.5 um, + fusoid, rostrate, thin-walled. STIPITIPELLIs a cutis of cylindrical,

parallel, distinctly thick-walled (walls up to 2.0 um thick), mostly smooth, less

frequently with scattered diverticula. CauLocystip1a 20-40 x 3.0-6.0 um,

adpressed to erect, cylindrical, narrowly clavate, subfusoid, mostly slightly

thick-walled. CLAMP CONNECTIONS present in all tissues.

CHEMICAL REACTIONS — No part of basidiocarps amyloid or dextrinoid.

EcoLocy — On fallen needles of Juniperus chinensis var. kaizuka.

Marasmius sect. Hygrometrici in Korea... 411

SPECIMEN EXAMINED — REPUBLIC OF KOREA, Sobaeksan National Park, Sobaeksan

Youth Hostel area, 16 July 2007 leg. J.G. Han (Antonin 07.59, BRNM 714677).

REMARKS — “Marasmius junipericola” is similar to M. crescentiae but differs

by the presence of well-developed caulocystidia. Although it represents a

distinct taxon, the collection was too small (two minute basidiocarps) for a

valid description of a new species. For similar taxa, see the discussion on

M. crescentiae.

Acknowledgments

We gratefully acknowledge Jan Holec (National Museum, Prague, Czech Republic)

and Michal TomSsovsky (Mendel University, Brno, Czech Republic) for critically

reviewing this manuscript. The collecting trip to the Republic of Korea and the studies of

the collected material by the first author were supported by the Czech Science Foundation

(No. 206/07/J003). The other authors were supported by the Korea Research Foundation

Grant funded by the Korean Government (KRF-2006-F00001), and research grant by

Korea Forest Institute (FP 0801-2010-01).

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Antonin V, Ryoo R, Shin HD. 2009. Marasmioid and gymnopoid fungi of the Republic of Korea.

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429-440. http://dx.doi.org/10.5248/108.429

Antonin V, Ryoo R, Shin HD. 2010a. Marasmioid and gymnopoid fungi of the Republic of Korea.

2. Marasmius sect. Globulares. Persoonia 24: 49-59.

http://dx.doi.org/10.3767/003158510X496107

Antonin V, Ryoo R, Shin HD. 2010b. Marasmioid and gymnopoid fungi of the Republic of Korea.

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MY COTAXON

http://dx.doi.org/10.5248/119.413

Volume 119, pp. 413-417 January-March 2012

A new hairy species of Leptogium (Collemataceae) from China

Hua-Jig Liu’ & SHUAI GUAN

Key Lab of Microbial Diversity Research and Application of Hebei Province,

College of Life Sciences, Hebei University, Baoding, 071002, China

* CORRESPONDENCE TO: liuhuajie@foxmail.com

ABSTRACT — This paper describes Leptogium weii sp. nov., a new hairy species from Sichuan,

southwestern China. Leptogium weii has dense white tomentum on apothecial stalks and

thalline margins. It closely resembles L. burnetiae in external morphology but differs in

having globose isidia on the upper surface and dense tomentum on the apothecial stalk and

thalline margin. A key is given to hairy species with tomentose apothecia.

Key worps — lichen, Ascomycota, Peltigerales, corticolous

Introduction

Thirty-two Leptogium species have previously been reported in China (Wei

1991, Aptroot 2002, Wang et al. 2010). Here we describe a characteristically

hairy new species, Leptogium weii.

Material & methods

The type specimen was collected from Sichuan Province in southwestern China,

and deposited in Herbarium Mycologicum Academiae Sinicae-Lichenes (HMAS-L). A

dissecting microscope (Motic SMZ-140) and a light microscope (Motic B2) were used

for the morphological and anatomical studies.

Taxonomy

Leptogium weii H.J. Liu & S. Guan, sp. nov. FIGURE |

MycoBank MB 519891

Differt a Leptogio burnetiae isidiis globosis, et marginibus apotheciorum dense

hirsutissimis; asci interdum quadrospori.

Type: China. Sichuan Province, Mt. Gonggashan, Zimeishan (29°37'N 101°53’E), alt.

4000 m, on branch of Rhododendron sp. (Ericaceae), 29/VII/1982, Xian-Ye Wang, Xie

Xiao & Bin Li 9082 (Holotype, HMAS-L 031713).

Erymo.oey: The species is named in honour of Professor Jiang-Chun Wei.

414 ... Liu & Guan

THALLUS foliose, 6-8 cm in diam. Lobes broadly rounded, 10-20 mm wide,

apices entire, undulate; UPPER SURFACE blue gray, usually shiny, glabrous,

without wrinkles, isidiate; isrp1a globular, laminal to marginal, dense, darker

than upper surface; LOWER SURFACE pale to medium gray, smooth, with a dense

covering of long-celled, white hairs, to 0.2 mm long, + naked near margin.

APOTHECIA + dense, laminal to submarginal, 1.0-2.5 mm in diam., stipitate;

pisc dark red brown, concave and finally plane, glabrous, dull; staLk always

with dense, suberect to erect, white tomentum; THALLINE MARGIN smooth,

paler than upper surface, non-isidiate, with similar tomentum on stalks,

sometimes glabrous when old.

THALLUS 90-110 um thick when moist. UpPpER CORTEX paraplecten-

chymatous, consisting of a single layer of globular, isodiametrical cells, 4-6 um

thick; LOWER CORTEX paraplectenchymatous, commonly consisting of a single

layer of ellipsoidal thin-walled cells, 8-10 um thick, or 40-50 um thick when

below apothecia and consisting of 3-4 layers of cells; MEDULLA 80-100 um

thick, composed of loosely interwoven hyphae running in various directions;

PHOTOBIONT Nostoc in chains, + globose, 5-7 um in diam., + concentrated near

both cortices.

APOTHECIA 300-400 um thick (when moist); THALLINE EXCIPLE 100-180

um wide; PROPER EXCIPLE not seen; EPIHYMENIUM brown, 10-15 um high;

HYPOTHECIUM yellowish, 40-50 um high. HyMENIUM in its central part

160-210 um high; paRAPHYSES simple, 1-1.5 um wide, at apices often thickened

(2-3 um); Asci clavate, 125-150 x 20-30 um, commonly 8-spored, sometimes

4-spored. Ascosporss fusiform to ellipsoidal, apices obtuse or sometimes acute,

25-35 x 12-17 um, transversely septate to submuriform, with 3-5 transverse

and 0-1 longitudinal septa, colourless. CONtDIOMATA not observed.

HABITAT AND DISTRIBUTION: Corticolous on branches of Rhododendron sp.,

in the alpine region of southwestern China.

REMARKS -— Leptogium weii is characterized by an upper surface with globular

isidia and without wrinkles, long hairs on the underside, white tomentum

on apothecial stalks and thalline margins, and (4—)8-spored ascus. It closely

resembles L. burnetiae C.W. Dodge in external morphology. But the latter

always has coralloid isidia, 8-spored ascus, and the apothecium is often isidiate

on thalline margin (Jorgensen & Nash 2004, Aragon et al. 2004, 2005, Wang et

Fic. 1. Leptogium weii (holotype). A. Thallus with apothecia. B. Apothecia with tomentose stalk

(note the globular isidia on upper surface). C. Thallus cross-section, showing corticated surfaces

and long hairs on lower surface. D. Apothecium cross-section. E. Thalline margin (note the

dense tomentum on the surface). F. Vertical section of the central part of apothecia (1 stands for

hymenium, 2: subhymenium, 3: medulla, and 4: corticated lower surface). G. An ascus with 8

spores. H. An ascus with 4 spores. I. Spores. Scale bars: A = 1 cm; B = 1 mm; C, E-I = 10 um;

D = 100 um.

Leptogium weii sp. nov. (China) ... 415

416 ... Liu & Guan

al. 2010). Leptogium trichophoroides P.M. Jorg. & A.K. Wallace and L. askotense

D.D. Awasthi also have tomentose stalks and thalline exciples, and long hairs

on the underside as observed in L. weii. But L. trichophoroides has wrinkles

on a browned upper surface without isidia and smaller spores (18-22 x 9-12

um) (Jorgensen 1997). Leptogium askotense differs from the new species in

lacking isidia, having wrinkles on the upper surface and longer spores (30-50

x 10-17 um) (Jorgensen 1997). Leptogium sphaerosporum P.M. Jorg. & Olley is

differentiated by phyllidia (without tomentum on apothecial stalk and isidia)

on the upper surface and sphaeroid spores (Jorgensen & Olley 2010), while

L. saturninum is distinguished by a darker upper surface, sessile apothecia with

a glabrous thalline exciple and euparaplectenchymatous proper exciple (Sierk,

1964, Jorgensen & Nash 2004).

The hairy species in Leptogium have been summarized by Jorgensen (1997).

Here we present a key to 12 hairy Leptogium species with tomentum on

apothecial stalks or thalline margins derived from keys by Jorgensen (1997),

Jorgensen & Kashiwadani (2008), and Jorgensen & Olley (2010).

Key to hairy Leptogium species with tomentose apothecial stalks

or thalline margins

1 Hains On both sides-or tallUsy 0.05 6 at ela a ead a, L. australe

Hairsonkon lower surtace. of fhalliiss: fe. assed. atthe atin deals daa tanede edna 2

2 Wpper-suriace-distiict ih Wrinkled mit RitoF Pot ee he ek eee 3

Upper surface smooth or slightly rough, not wrinkled ...................0. 2

3 UiFicll UiseISTOTAL ers 20 sleva RL vacate Parnstee tanec, Mews My teva ect twn Sets ed L. papillosum

Ubyal ers Gt stl bate a be sete aes cad Nee ee a ge Be age sie laght Bin deg ee d's Gennes +

4 Proper exciple distinct, euparaplectenchymatous ............. 0.0.00. eee eee 5

PLODEERC IIE ANSEL Wh.ny Fuk ee FURAN TUE eh hee REE RAAT 2 L. acutisporum

5 Thallus normally bluish; apothecia sessile, margin wrinkled ...... L. arisanense

Thallus normally dark brownish or greenish ............... 0.0. cece 6

6 Apotnectaywit hist marotnal Maitse.s 5 bret nS eencset mation mats oe eee ta aoe ee fi

Apothecia without stiff marginal hairs, sometimes softly hairy on the stalk

Settle attetlent eats lnt a ve tus bah ua eee bien eneatlush eematiseercarntsteneata sranceas ak L. askotense

7 Apothecia stalked, with white hairs when young ............ L. trichophoroides

Apothecia sessile or short stalked, usually with blackish hairs ............... 8

8 Lobes strap=sshaped so. Saw. Se Be en eh ew ee soe aaah eeo ee es L. loriforme

MODES SHOGMOR DICH AT ae ui, 9% cia'g % ain'y Ue eig oh hls Oh ee ORGS Oy L. trichophorum

9 MANS AST Later eee Fe Sage a Sie a Pale apalie Nae ie sea lal te eat dla aban dats bi aleeata tease 10

WANS AGE ASWATATS wurde. eceaeths haceeathy Sere aey eee en een he Ee Ee Rhy ok 11

10 __Isidia cylindrical coralloid; apothecia rare, often isidiateon margin L. burnetiae

Isidia usually granular; apothecia abundant, not isidiate on margin ..... L. weii

Leptogium weii sp. nov. (China) ... 417

ld, ~ ‘Apothecia prominently stalked* .. 2..cc2.nencb he eta ce kee eh L. pedicellatum

mpothecia sessile or Short stalkers ..tone. sate hiu eho Bis wha Rie wate Bix a's L. menziesii

Acknowledgements

This paper was supported by the National Natural Science Foundation of China

(31093440, 31000239), Polar Science Foundation of China (20080205) and Natural

Science Foundation of Hebei University (Y2007-099). The authors are indebted to Prof.

Jiang-Chun Wei and Ms. Hong Deng (HMAS-L), Institute of Microbiology, Chinese

Academy of Sciences for sending the specimens on loan. Thanks go to Prof. Shou- Yu

Guo (Institute of Microbiology, Chinese Academy of Sciences), Prof. Per M. Jorgensen

(Institute of Bergen), and Prof. Jae-Seoun Hur (Korean Lichen Research Institute,

Sunchon National University) for reading and improving the manuscript, and for acting

as presubmission reviewers.

Literature cited

Aptroot A, Sparrius LB, Lai MJ. 2002. New Taiwan macrolichens. Mycotaxon 84: 281-292.

Aragon G, Martinez I, Otalora MAG. 2004. New data on the distribution of Leptogium azureum

(Swartz) Mont. The Lichenologist 36(5): 345-347.

http://dx.doi.org/10.1017/S0024282904014318

Aragon G, Otalora MAG, Martinez I. 2005. New data on the genus Leptogium (lichenized

ascomycetes) in the Iberian Peninsula. Nova Hedwigia 80(1-2): 199-226.

http://dx.doi.org/10.1127/0029-5035/2005/0080-0199

Jorgensen PM. 1997. Further notes on hairy Leptogium species. Symbolae Botanicae Upsalienses

32(1): 113-130.

Jorgensen PM, Kashiwadani H. 2008. Leptogium loriforme P.M. Jorg. & Kashiw., a new hairy species

from Papua New Guinea. The Lichenologist 40(2): 123-125.

http://dx.doi.org/10.1017/S0024282908007524

Jorgensen PM, Nash TH III. 2004. Leptogium. 330-350, in: TH Nash III et al. (eds). Lichen flora

of the Greater Sonoran Desert Region, Vol. 2. Lichens Unlimited, Arizona State University:

Tempe, Arizona.

Jorgensen PM, Olley L. 2010. A new hairy Leptogium from Nepal. The Lichenologist 42(4):

387-389. http://dx.doi.org/10.1017/S0024282909990764

Sierk HA. 1964. The genus Leptogium in North America north of Mexico. The Bryologist 67(3):

245-317. http://dx.doi.org/10.2307/3240893

Wang HY, Ren Q, Li HM, Wang HY, Zhao ZT. 2010. Five lichens of Leptogium new to China.

Mycotaxon 111: 161-166. http://dx.doi.org/10.5248/111.161

Wei JC. 1991. An enumeration of lichens in China. International Academic Publishers: Beijing.

278 p.

MY COTAXON

http://dx.doi.org/10.5248/119.419

Volume 119, pp. 419-429 January-March 2012

Abrachium, a new genus in the Clathraceae,

and Itajahya reassessed

TIARA SOUSA CABRAL', PAULO MARINHO’,

BRUNO TOMIO GOTO? & IURI GOULART BASEIA™?

'Programa de Pés Graduagao em Ciéncias Biol6dgicas,

*Departamento de Biologia Celular e Genética, &

*Departamento de Botanica, Ecologia e Zoologia,

Universidade Federal do Rio Grande do Norte, PO Box 59072-970, Natal, RN, Brazil

* CORRESPONDENCE TO: baseia@cb.ufrn. br

AxsstrRact — Molecular and morphological analyses have elucidated phylogenetic

relationships of two remarkable species in the Phallales: Aseroe floriformis and Phallus roseus.

Genes from ATPase subunit 6 (atp6), the nuclear large subunit ribosomal DNA (nuc-LSU),

and the second largest RNA polymerase II subunit (RPB2) underwent Bayesian and parsimony

molecular analyses. Molecular datasets, combined with morphological characters, support a

new genus (Abrachium for Aseroe floriformis), reassessment of Itajahya, and emendation of

Clathraceae.

Key worps — Phallomycetidae, gasteroid fungi, neotropics, phylogeny, evolution

Introduction

Fischer (1898-99) established the order Phallales, initially composed of two

families, Clathraceae and Phallaceae, to separate representatives with branched

and unbranched basidiomata. Clathraceae, established by Chevallier (1826),

encompasses the “lattice stinkhorns’, fungi that exhibit a peculiar basidioma

featuring tubular or mucilaginous arms either interconnected or intertwined

in a network or comprising a receptacle with vertical arms either connected at

the top or arched in the shape of a star (Miller & Miller 1988). Pegler & Gomez

(1994) included twelve genera within the family (Blumenavia, Linderiella,

Pseudocolus, Anthurus, Aseroe, Lysurus, Clathrus, Ileodictyon, Simblum, Colus,

Neolysurus and Kalchbrennera).

Corda (1842) proposed Phallaceae for species with fruiting bodies

composed of a single, hollow receptacle with a foul-smelling gleba covering

the apex. Morphological characters widely used to separate genera include

420 ... Cabral & al.

the presence of a campanula (head) at the apex of the receptacle in Phallus,

indusium in Dictyophora, and calyptra in Itajahya. Some genera lack a distinct

head and form the mucilaginous gleba directly on the apical receptacle (e.g.,

Mutinus) or at a constriction near the apex (e.g., Staheliomyces). Several

attempts were made to establish a synopsis of the family (Lloyd 1909, Fischer

1933), and Cunningham (1944) included seven genera: Dictyophora, Itajahya,

Phallus, Mutinus, Staheliomyces, Aporophallus, and Floccomutinus. However,

Kreisel (1996) viewed Dictyophora, Aporophallus, Itajahya, Echinophallus, and

Endophallus as subgenera of Phallus sensu lato, using the shape and surface

of the head, pigmentation of the receptacle, and morphology of the volva and

rhizomorphs as primary characteristics. He did not deem the presence of a pore

at the receptacle apex or the morphology of the head, indusium, and calyptra as

sufficient for segregation at the generic level (Kreisel 1996).

Taxonomic studies of Neotropical phalloid species have revealed new

and intriguing species (Baseia et al. 2003, Baseia & Calonge 2005, Fazolino

et al. 2010). The discovery of new species is essential to understanding the

evolutionary path of Phallales clades, since according to known estimates

(Hawksworth 2001, Mueller et al. 2007, Blackwell 2011), most tropical species

remain unknown. Through morphological data and molecular tools the

classification of macrofungi should change in coming years.

The present study aims to evaluate the actual taxonomic position of Aseroe

floriformis and Phallus roseus, two species with distinct morphological features

that have not yet been analyzed by molecular studies.

Materials & methods

Basidioma sampling

Basidiomata were sampled at the Parque Estadual Dunas do Natal (06°22'10" S

and 35°00'28" W) and Estacao Ecoldgica do Seriddé (06°35'02" S and 37°17'02" W),

Rio Grande do Norte state, Brazil. Specimens of Aseroe floriformis and Phallus roseus

were photographed and collected during the rainy seasons in 2008 and 2009. Materials

studied were deposited in the fungal collection of the UFRN Herbarium.

Macroscopic characters were described from fresh specimens. Basidiomata colors

were based on Kornerup & Wanscher (1978). Microscopic characteristics were

determined from temporary mounts, hand-sectioned with a razor blade on the vertical

axis of the basidiomata and mounted in 3% KOH and 1% cotton-blue lactophenol,

respectively. All measurements were made under oil-immersion microscope objective

at 1000x magnification. Spore measurements included the largest and smallest spores

and at least 20 additional randomly selected spores from each specimen.

Molecular techniques

Fresh basidiomata of Aseroe floriformis (UFRN 1481) and Phallus roseus (UFRN

535) were used for CTAB DNA extraction. The CTAB buffer (1.4M NaCl, 100mM Tris-

HCl pH 8.20mM EDTA and 2% CTAB) was added to 0.8¢ of fresh tissue pulverized

Abrachium floriforme gen. & sp. nov. (Brazil) ... 421

with liquid nitrogen and incubated for 30 min. at 65°C. After incubation, a chloroform-

isoamyl (24:1) extraction was performed, followed by isopropanol precipitation. Total

DNA amount was quantified on a NanoDrop-2000 spectrophotometer (NanoDrop

Technologies Inc. Wilmington, DE 1980, United States).

DNA sequences were obtained for three different regions: nuc-LSU-rDNA (LROR/

LR5), ATPase subunit 6 (ATP61/ATP62) and the second largest subunit of RNA

polymerase RPB2 (bRPB2-6F/bRPB2-7R), with previously designed primers (Vilgalys

1990; Liu et al. 1999; Kretzer & Bruns 1999).

All PCR reactions were carried out in a final volume of 50ul, with PCR buffer 10X,

4 mM of MgCl, 25 ng of DNA, 0.3 mM of dNTP Mix, 12.5 pmol of each primer and

0.3ul of Invitrogen® DNA Taq Polymerase (5U/ul). The initial cycling parameter for

atp6 reaction condition consisted of 2 min at 95°C for initial denaturation, 5 cycles of 35

s at 94°C, 55 s at 37°C and Imin at 72°C. This was followed by 30 cycles of 35 s at 94°C,

55 s at 45°C, 1 min at 72°C, and final extension of 5 min at 72°C with final hold of 4°C.

The nuc-LSU-rDNA amplification condition underwent initial denaturation for 2 min

at 95°C, followed by 39 cycles for 1 min at 94°C, 30 s annealing at 45°C, 1 min extension

at 72°C, and a final extension for 5min at 72°C. RPB2 amplifications followed conditions

described by Liu et al. (1999).

Amplicons were verified by ethidium bromide-stained 0.4% agarose gel

electrophoresis. Total amount of amplified DNA was quantified by a NanoDrop-

2000 spectrophotometer. Amplifications with low DNA content were re-amplified by

increasing PCR cycles.

PCR products were purified with Wizard SV Gel and PCR Clean-Up System

(Promega). The sequencing reaction was performed with a DNA sequencing kit

(BigDye Terminator v3.1 Cycle Sequencing Kit) using the same primers employed to

amplify each fragment, and sequencing was conducted an ABI 3730 DNA Analyzer

(Applied Biosystems). Both were carried out at the Center for Human Genome Studies

(University of Sao Paulo, CEGH-USP) and later assembled using DNA Baser v3.0.12

beta.

Phylogenetic analysis

Phylogenetic analyses were performed for three combined DNA datasets under

Bayesian and parsimony criteria. Newly generated sequences for Aseroe floriformis and

Phallus roseus ATP6, nuc-LSU-rDNA and RPB2 and those retrieved from GenBank

(National Center for Biotechnology Information, http://www.ncbi.nlm.nih.gov — TABLE

1) were separately aligned using ClustalX2 (Larkin et al. 2007), with default settings

(alignment files are available at http://purl.org/phylo/treebase/phylows/study/TB2:

$11546). Manual adjustments were made in BioEdit v. 7.0.9.0 (Hall 1999). The three

alignments were then concatenated with Phyutility (Smith & Dunn 2008). Maximum

parsimony analyses were conducted with combined dataset file, in PAUP* (Phylogenetic

Analysis Using Parsimony) version 4b10 (Swofford, 2002). Heuristic search was used with

tree bisection-reconnection (TBR) and MULTrees. Confidence levels were established

through 1000 bootstrap replicates. Bayesian analysis was carried out with combined

dataset in MrBayes v.3.1.2 (Huelsenbeck & Ronquist 2001). Two different runs with

four incrementally heated simultaneous Monte Carlo Markov chains were conducted

over 2 million generations, applying respective substitution models determined for

422 ... Cabral & al.

each partition in MrModeltest v.2.2 (http://www.abc.se/~nylander/). Trees were

sampled every 100 generations to produce 20,000 trees. In order to estimate posterior

probabilities, 25% of the trees were discarded as a burn-in stage, observing when average

standard deviation of split frequency (ASDSF) values dropped below 0.01.

TABLE 1. GenBank accession numbers used for phylogenetic analysis.

GENBANK ACCESSION NUMBER

SPECIES

nucLSU ATP6 RPB2

Anthurus archeri DQ218624 DQ218913 DQ219081

Abrachium floriforme JF968440 JF968438 JF968442

Aseroe rubra DQ218625 DQ218914 DQ219082

Clathrus chrysomycelinus DQ218626 DQ218915 DQ219083

Claustula fischeri K.M. Curtis — — DQ219086

Dictyophora duplicata (Bosc) E. Fisch. DQ218481 DQ218765 DQ219087

D. indusiata (Vent.) Desv. DQ218627 DQ218917 DQ219088

D. multicolor Berk. & Broome DQ218628 DQ218918 DQ219089

Gelopellis sp. DQ218630 DQ218919 DQ219090

Gelopellis sp. DQ218631 DQ218920 DQ219091

Tleodictyon cibarium DQ218633 DQ218922 DQ219093

I. gracile DQ218636 DQ218925 DQ219096

Itajahya rosea JF968441 JF968439 —

POL CRE EM EADS ae DQ218638 DQ218926 DQ219098

Laternea triscapa DQ218640 DQ218928 DQ219099

Lysurus borealis (Burt) Henn. DQ218641 DQ218929 DQ219100

L. mokusin (L.) Fr. DQ218507 DQ218791 DQ219101

Mutinus elegans (Mont.) E. Fisch. AY574643 AY574785 DQ219102

*Phallobata alba G. Cunn. DQ218642 DQ218930 DQ219103

~Phallus costatus (Penz.) Lloyd DQ218513 DQ218797 DQ219104

P. hadriani Vent. DQ218514 DQ218798 DQ219044

P ravenelii Berk. & M.A. Curtis DQ218515 DQ218799 DQ219105

Protubera borealis S. Imai DQ218516 DQ218800 DQ219106

P. canescens DQ218645 DQ218932 DQ219108

P. jamaicensis (Murrill) Zeller DQ218647 DQ218933 DQ219110

P. maracuja Moller DQ218518 DQ218802 DQ219111

P. parvispora Castellano & Beever DQ218648 DQ218934 DQ219112

P. sabulonensis Malloch DQ218649 DQ218935 DQ219113

Simblum sphaerocephalum Schltdl. DQ218521 DQ218806 DQ219115

*Trappea darkeri (Zeller) Castellano DQ218651 DQ218938 DQ219116

Pseudocolus fusiformis AF213128 — —

* Species used as outgroup

Abrachium floriforme gen. & sp. nov. (Brazil) ... 423

Results

Phylogenetics analysis

Bayesian and maximum parsimony analyses produced no significant

differences in tree topology, indicating data consistency to resolve the

phylogeny of the group. The phylogram (Fic. 3) illustrates six well-formed

clades corresponding to the families of Phallales with high support values

(PP = 1, BT = 90-100). Intergeneric relationships in each family are also

shown. It was found that Aseroe floriformis does not cluster with Aseroe rubra,

and occupies a basal position in the Clathraceae, with PP = 1 and BT = 99.

In Phallaceae, Phallus roseus occupies a position between Mutinus elegans and

other Phallus species, including indusiate representatives (Dictyophora), which

cluster into a single clade displaying high levels of reliability (PP = 1, BT = 94).

Taxonomic revision

Based on the morphological and molecular data set, we propose a new genus

to accommodate Aseroe floriformis. The most important morphological feature

of the new genus —the lack of arms— requires an emendation in Clathraceae.

Phylogenetic analyses also support the status change of the genus Itajahya in

Phallaceae.

Clathraceae Chevall. emend. Baseia

EMENDED DESCRIPTION: Basidiomata sessile to shortly stalked, upright

receptacle, spreading, sometimes arched and united arms or a latticed

receptaculum composed of interconnected tubular or gelatinous arms

expanding from a flaccid button, which acts as a volva, or armless sunflower-

shaped receptacle. Gleba located on the central disc of receptacle or inside

and next to the clathrate receptaculum. Peridium three-layered. Basidiospores

narrowly elliptical, covered with mucilaginous layer.

TYPE GENUS: Clathrus P. Micheli ex L.

OTHER GENERA: Abrachium Baseia & T.S. Cabral, Aseroe Labill.,

Blumenavia Moller, Pseudocolus Lloyd, Lysurus Fr., Ileodictyon Tul. & C.

Tul.

The genus Aseroe was proposed by Labillardiére (1800), with Aseroe rubra

Labill. as the type (and only) species. Fischer (1890) subsequently described

A. arachnoidea. Yoshimi & Hongo (1989) added A. coccinea, a name recently

validated by Kasuya (2007). Baseia & Calonge (2005) discovered a new species,

Aseroe floriformis, in northeastern Brazil in sand dunes. Although Index

Fungorum lists 27 names in Aseroe, the latest edition of the Dictionary of

Fungi (Kirk et al. 2008) lists only two: A. floriformis and A. rubra. The genus

has a pantropical distribution, although A. rubra has been reported in Europe

(Spooner 2005). Aseroe is characterized by a cylindrical receptacle and stipe

424 ... Cabral & al.

that emerge from a whitish volva. With the exception of Aseroe floriformis, all

species have apical free arms internally covered by the gleba (Dring 1980).

Abrachium Baseia & T.S. Cabral, gen. nov.

MycoBank MB 561618

Ovo subglobosae, epigaeo, rhizomorphis basalis. Stipite cylindrico, spongioso. Receptaculum

floriformis cum disco sine ramulis, centralis perforatae cum gleba gelatinosa marginata,

foetida. Sporis cylindraceo-bacilliformis, laevis, hyalinae.

TYPE SPECIES: Abrachium floriforme (Baseia & Calonge) Baseia & T.S. Cabral

EtyMo.oecy: Abrachium, due to the absence of arms.

KEY CHARACTERS: Egg subglobose, epigeous with several and basal mycelial

cords, white. Stipe cylindrical, spongy. Receptacle with the same anatomy,

sunflower-shaped, without arms, central disc covered by a gelatinous gleba.

Spores cylindrical to bacilloid, smooth, hyaline.

Abrachium floriforme (Baseia & Calonge) Baseia & T.S. Cabral, comb. nov. = Fic 1

MycoBank MB 561619

= Aseroe floriformis Baseia & Calonge, Mycotaxon 92: 170, 2005.

SPECIMENS EXAMINED: BRAZIL, R10 GRANDE DO NorrzE, Natal, Parque das Dunas,

ad solum arenarius, 3-VII-2004, legit I.G. Baseia & P.P.T. Lacerda, UFRN-Fungi 1851

(holotype); 07-V-2009, legit I.G. Baseia UFRN 1481, GenBank JF968440, JF968438,

JF968442.

Our phylogenetic analyses confirm high support values for Phallales in

accordance with Hosaka et al. (2007). Morphological and molecular analyses

supported the establishment of the genus Abrachium, named for its armless

receptacle (Fic 1). This taxon was originally described in the Brazilian Atlantic

rainforest (Baseia & Calonge 2005), but has since been recorded in northeastern

Brazil (Trierveiler-Pereira & Baseia 2009, 2011, Bezerra et al. 2009).

The clathraceous clade comprises members with a variable morphology of

the basidiomata, consisting of tubular or gelatinous arms either interconnected

or free (Dring 1980). The armless receptacle of Abrachium justifies an

emendation in Clathraceae and seems to be a primitive characteristic of the

clade, as shown in the tree obtained by molecular inference (Fic. 3). Molecular

data indicate that receptacle morphology is a phylogenetically informative

characteristic in Clathraceae. In phylogram, Abrachium represents a basal

clade, which is a sister group of a clade formed by other Clathraceae species.

The latter have a bifurcation leading to two other subgroups. ‘The first subgroup

either has arms attached at the ends [Pseudocolus fusiformis (E. Fisch.) Lloyd],

or free and arched like a star [Aseroe rubra and Anthurus archeri (Berk.) E.

Fisch.]. The second subgroup displays a complex structure, forming a “cage”

type configuration including Clathrus chrysomycelinus Moller, Laternea

triscapa Turpin, Ileodictyon cibarium Tul. & C. Tul., Ileodictyon gracile Berk.,

Abrachium floriforme gen. & sp. nov. (Brazil) ... 425

FiGuRES 1-2. Fresh specimens: 1. Abrachium floriforme. 2. Itajahya rosea. Bar = 10 mm.

and Protubera canescens G. W. Beaton & Malajczuk. Based on these data,

we can infer that the evolution to receptacle bifurcation (forming arms, e.g.,

Aseroe) occurred at least once in the Neotropical region. Presumably, after the

divergence of A. floriforme, a common ancestor has diverged to forms where

the receptacle may bifurcate like arms in one group, and form a cage in the

other (such as Ileodictyon).

Itajahya rosea (Delile) E. Fisch., Ber. Deutsch. Bot. Ges. 47: 294, 1929. Fic. 2

= Phallus roseus Delile, Flore d Egypte: 300, 1813.

SPECIMEN EXAMINED: BRAZIL, R10 GRANDE DO NortTE, Serra Negra do Norte,

Estacao Ecoldgica do Serid6, ad solum arenarius, 23-V-2008, legit T.B.S. Ottoni, UFRN-

Fungi 535; GenBank JF968441, JF968439.

Molecular data analysis of Phallus roseus points to the need for reassessing

its current taxonomic status in the subgenus Itajahya proposed by Malencon

(1984) and Kreisel (1996). Itajahya was established by Mdller (1895) based on I.

galericulata Moller, a taxon discovered in Santa Catarina state, Brazil. The main

feature distinguishing Itajahya from other Phallaceae genera is the presence

of a structure called a calyptra (Fic. 2) located at the apex of the gleba (Méller

1895, Malencon 1953, Ottoni et al. 2010). Molecular data demonstrated that

P. roseus diverged after Mutinus and prior to other species of Phallus (Fic.

3). Since I. rosea does not cluster with any other Phallus species, we propose

426 ... Cabral & al.

Phallobata alba

Trappea darkeri

Abrachium floriforme

Pseudocolus fusiformis

1/99} 1/92

Aseroé rubra

0,73/75

Anthurus archeri

1/61

0.99/68

Clathrus chrysomycelinus

Laternea triscapa

1/92

Protubera canescens

1/99 lleodictyon gracile

1/98

lleodictyon cibarium

1/100

1/97

1/93

1/90

1/74 1/97

1/90

1/99

1/94

Mutinus elegans

Itajahya rosea

1/99 Dictyophora multicolor

1/94 Dictyophora duplicata

Phallus ravenelli

1/99 Phallus costatus

Phallus hadriani

0.99/87

Dictyophora indusiata

Lysurus mokusin

Simblum sphaerocephalum

Lysurus borealis

Protubera parvispora

1/98 1/100

1/100

Kobayasia nipponica

Protubera sabulonensis

1/92

Protubera borealis

Protubera jamaicensis

Protubera maracuja

Gelopellis sp. H4571

Claustula fischeri REB 2326

Gelopellis sp. H4397

0.1

100

Claustula fischeri OSC122661

FiGurE 3. Phylogenetic tree of Phallales obtained through Bayesian analysis of atp6, nuc-LSU

rRNA, and RPB2 sequences. Numbers preceding ‘/’ = Bayesian posterior probability values (PP)

and after ‘/’ = maximum parsimony bootstrap values (BT).

Abrachium floriforme gen. & sp. nov. (Brazil) ... 427

raising Itajahya to generic status. Further collections and molecular studies of

I. galericulata (= Phallus galericulatus (MGller) Kreisel) are also needed in order

to ensure morphological characteristics (mainly the presence of calyptra) agree

with molecular data recognizing the genus Itajahya.

Molecular data also imply that the indusium is not a phylogenetically

informative character in Phallaceae. Malencon (1984) and Kreisel (1996)

stated that species may be closely related with or without this structure, and

that a species may or may not produce it or have a rudimentary form of it.

The phylogram indicates that although Dictyophora multicolor and D. duplicata

form a separate clade, D. indusiata clusters with Phallus hadriani, exhibiting

prior divergence with the other mentioned clade. This suggests that indusiate

species may have emerged independently during several phases of the group's

evolution, thereby characterizing it as a polyphyletic assemblage. The phylogram

also illustrates that Mutinus is a more primitive genus in Phallaceae, as observed

by Cunningham (1944), based on data from simple sporophore morphology

and the absence of complex structures such as campanulate apex, indusium

and calyptra. It is important to note that further molecular studies are needed,

especially with key-taxa that may lead to a change in current classification and

give us a clearer picture of the systematics and evolution of phalloid fungi.

Acknowledgments

We thank CNPq for the Msc. Scholarship and PPBio Semi-Arid Project for

financial support for this study. Sincere thanks are given to Prof José Luiz Bezerra

(Universidade Estadual de Santa Cruz) for encouraging this work, to Prof Vagner

Cortez for presubmission review, and to Prof. Gladstone Alves da Silva (Departamento

de Micologia, Universidade Federal de Pernambuco) for critical comments of molecular

analyses.

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MY COTAXON

http://dx.doi.org/10.5248/119.431

Volume 119, pp. 431-438 January-March 2012

Nomenclatural notes 13. An incorrect neotype designation

and provision for a lectotype and an epitype for Helvella fusca

FIDEL LANDEROS !** & RICHARD P. KorF®

'Facultad de Ciencias Naturales, Universidad Auténoma de Querétaro,

Avenida de las Ciencias s/n, Santiago de Querétaro, Qro., 76270, Mexico

?Departamento de Botanica y Zoologia, Universidad de Guadalajara,

Apartado postal 1-139, Zapopan, Jal., 45101, Mexico

*Plant Pathology Herbarium, Cornell University, Ithaca, NY 14853, USA

CORRESPONDENCE TO *: *landeros@uaq.mx, *info@mycotaxon.com

ABSTRACT — A neotype designation for Helvella fusca proposed in 1997 proves to have been

erroneous because an illustration accompanying the original description was overlooked. Such

illustrations normally have served as the lectotype when all known specimens of the author's

taxon have been lost. The authors of that neotype designation should instead have designated

the illustration as a lectotype and a specimen as an epitype, which is done in this paper.

The so-called neotype specimen is now selected as the epitype specimen. A full description,

illustrations and remarks about synonyms and a non-synonym are also provided.

Kry worps — International Code of Botanical Nomenclature

Helvella fusca was described by Gillet (1879: 9), but unfortunately his description

did not specify any specimen or figure to represent that species, making

typification more difficult. Dissing (1966b) wrote about this species: “I have not

seen any authentic specimen of H. fusca Gillet, whereas the many Bresadola-

collections of H. fusca in the herbarium in Stockholm (S) make it possible to

get a precise concept of Bresadola’s species.” With respect to the type specimen

of H. fusca, Dissing (1966b) wrote only, “not seen.” Later, in their seminal paper

on the northern and northwestern Helvellaceae of North America, Abbott &

Currah (1997) made a minor but (we believe) critical error by designating a

neotype specimen for H. fusca, even though that species does not occur in

North America. They overlooked the excellent illustration in Gillet’s paper

(pl. 4, not cited in the species description but referenced mistakenly in one plate

listing as pl. 14 and correctly in the other as pl. 4). They may have been unaware

that such an illustration, if one upon which it can be shown that the description

432 ... Landeros & Korf

HELVELLEES.

HELVELLA FUSCA. Gillet. 1

Fics. 1-2. 1: Type of Helvella fusca, reproduced from Gillet (1879). 2: Part of Helvella fusca

epitype (Bresadola’s specimen 05 21 1898, S). Scale bar = 1 cm.

or diagnosis validating the name was based, is considered part of the original

material (McNeill et al. 2006: Vienna Code Art. 9 Note 2) and becomes eligible

as a lectotype (sometimes informally referred to an “iconotype”) of the species

name (Fic. 1). An interesting article by Ross (2002) brings into question

whether an illustration accompanying an original description is always part

of the original material and thus eligible as a lectotype when all specimens are

lost or missing. At the XVIII International Botanical Congress in 2011 one

proposal attempting to clarify such situations (Art. 9, Prop E (215); see McNeill

& Turland 2011, Perry 2010) regrettably was rejected by a card vote (McNeill

et al. 2011). We feel that this case should be corrected at a future Congress but

now proceed under the assumption that Gillet’s plate was drawn from living

specimens now lost and that the illustration is part of the original material that

can and should be designated as a lectotype.

Helvella fusca, lecto- and epitypified ... 433

A neotype may only be proposed when ALL original material is lost [Vienna

Code]:

Art. 9.6. A neotype is a specimen or illustration selected to serve as nomenclatural

type if no original material is extant, or as long as it is missing.”

Also pertinent is Art. 9.7:

An epitype is a specimen or illustration selected to serve as an interpretative type

when the holotype, lectotype, or previously designated neotype, or all original

material associated with a validly published name, is demonstrably ambiguous

and cannot be critically identified for purposes of the precise application of the

name of a taxon.

The senior author pointed out this discrepancy to the junior author, and we

have chosen instead to propose as epitype the same specimen designated

as neotype by Abbott & Currah (1997) from the Bresadola collection in the

Naturhistorische Rijksmuseum in Stockholm, Norway —AI Deserto, [Italy],

collected by Bresadola, 05 21 1898 (Fic. 2). This specimen is also illustrated by

Dissing (1966b). Oddly, Abbott & Currah failed to cite (or were unaware of)

Bresadola’s careful description, which clearly cited Gillet’s figure and provided

a superb plate of his own collection (Bresadola 1900: 100, tab. CCXII) (Fie. 3).

That plate and description was reissued (Bresadola 1933: 1174, pl. 1174) with the

headers and footers of the plate in different typeface and numbering and with

a number of emendations of the text —notably eliminating Bresadola’s 1900

acknowledgement that Boudier had informed him that specimens Bresadola

had sent to him were identical with Gillet’s species. The 1933 plate renders the

figures in paler colors throughout.

Since Gillet’s plate does not provide enough information for modern

identification, an epitype is in order. The CopDE requires that if an illustration is

the only surviving element of the original material, that illustration should be

designated as a lectotype, which we do in this paper. The confused information

that already exists on the morphology of this species in the literature can

be resolved with an epitype specimen. Gillet (1879) described H. fusca with

spherical to subspherical ascospores, and he did not mention any other useful

microscopic character. Bresadola (1900) described ascospores as ellipsoidal,

18-20 x 12-13 um (but he did not indicate whether those measurements

correspond to smooth or to verrucose ascospores), and a glabrous, sterile outer

surface of the apothecium. Dissing (1966b) mentioned ascospores 17-18.4-21

x 11-12-13 um, with pustules when young, and the outside of the apothecium

naked or very delicately pubescent. The senior author studied Bresadola’s

specimen 05 21 1898 (now the epitype) and observed that smooth (young)

ascospores were larger, while verrucose (mature) were smaller (Fic. 5a) (see

species description), with the hyphal fascicles of the apothecium absent or < 50

um in length (Fre. 5b), corresponding to a glabrous to subpubescent surface.

434 ... Landeros & Korf

Inasmuch as the incorrect neotypification appeared in Mycotaxon (Abbott

& Currah 1997: 61), we have chosen that same journal in which to rectify their

action and to propose both the new lectotype and new epitype for the name.

The corrected typifications and full description

Helvella fusca Gillet, Champignons de France, Discom.: 9, pl. 4, 1879. FIG. 5

Type: Pl. 4, in Gillet, Champignons de France, Discom., 1879 (Lectotype, designated

here); Bresadola 05.21.1898, Al Deserto, Italy (S) (Epitype. designated here).

= Helvella fusca var. bresadolae Boud., Icon. Mycol. (Paris),

Tome 2: pl. 230, 1910; Tome 4: 121, 1911.

= Helvella fusca var. gyromitroides Chenant., in Pelé & Chenantais,

Bull. Soc. Sci. Nat. Ouest, Sér. 4, 1: 75, 1921.

# Helvella sulcata a fusca Afzel., K. Vetensk.-Acad. Handl. 4: 305, Tab. X, fig. 1, 1783.

APOTHECIUM 6-25 mm diam., 4-25 mm high, irregularly lobed or irregularly

saddle shaped, margin fused to the stipe or rarely free, hymenium pale brown,

tawny, to dark brown when dry, outside glabrous to subpubescent, whitish,

yellowish, to pale brown, ribs branched or unbranched, extending near to

the marginal zone of the excipular surface. Stripe 8-45 mm high, 3-15 mm

broad, equal or slightly wider at the base, whitish or pale brown, glabrous to

subpubescent, lacunose, internally chambered.

Asct 260-320 x 14-15.5 um, pleurorhynchous. AscosPporEs young, smooth:

(17-)18.5-21 x 11-13 um; mature, verrucose: 15.5-18.5 x 11-12.5 um.

PARAPHYSES Clavate, 5-8 um wide at the apex, simple or branched, septate, thin-

walled, yellowish brown, pigmented deposits in the cytoplasm. MEDULLARY

EXCIPULUM hyaline. ECTAL EXCIPULUM yellowish brown, pigmented deposits

usually in the cell wall and in a few cells in the cytoplasm. INNER LAYER OF

THE STIPE hyaline. OUTER LAYER OF THE STIPE yellowish brown, pigmented

deposits usually in the cell wall and in few cells in the cytoplasm. HyPHAL

FASCICLES of the apothecium and stipe up to 50 um long, light brown in mass,

pigmented deposits in the cell walls. None of the pigmented deposits in any of

the structures are visible in cotton blue mounts. All tissues are “rr-.”

REMARKS — Helvella fusca can be confused with H. lacunosa Afzel., because

both have lobed apothecia, a lacunose stipe, glabrous to subpubescent excipular

surface with the outer surface of the stipe similar, but H. fusca has a pale brown

hymenium and grows associated with Populus spp., while H. lacunosa has a

black or greyish hymenium and has a global distribution with no particular

association with Populus. Boudier (1904) first introduced H. fusca var.

bresadolae as a nomen nudum for a plate he planned to issue later. As pointed

out by Korf (1986: 213, footnote; 1988: 213, footnote), Boudier (1907) soon after

abandoned the variety, citing his plate as merely H. fusca. Boudier (1910), who

eventually validly proposed H. fusca var. bresadolae (Fic. 4) based on apothecia

Helvella fusca, lecto- and epitypified ... 435

Fran a » As

FF Seow Fab. COXII

Fics. 3-5. 3: Helvella fusca reproduced from Bresadola (1900, from Boudier’s book deposited in

Cornell University Library). 4: Type of H. fusca var. bresadolae reproduced from Boudier (1910,

Tome 2, pl. 230). 5: Helvella fusca (epitype), a: ascospores, b: ectal excipulum, c: paraphyses and

asci. Scale bar: 5a = 8 um; 5b, 5c = 20 um.

436 ... Landeros & Korf

that Bresadola sent him (but unfortunately not cited in the description), wrote

“Heec varietas typo simillima, differt tantummodo pediculo non albido, sed

pileo concolore.’ Boudier (1911) later expanded the description. Dissing

(1966a) considered this variety as invalid, because this specimen is possibly

part of the Bresadola material that he studied and determined as “in all details

identical?” We also do not recognize the variety, because Gillet did not describe

the stipe color only as white but as “dessous pale, blanc sale ou lég|éremen]t.

fuligineux.” Furthermore, in Helvella stipe coloration varies too greatly within a

species to be considered useful for recognizing varieties. For example the stipe

of H. lacunosa can vary from whitish to grayish to black. Chenantais (Pelé &

Chenantais 1921) proposed H. fusca var. gyromitroides, but the features that

he used to recognize the variety (multilobed, fawn and free apothecial margin,

and lacunose and yellowish stipe) are within the range of variation of the

species. We do not consider H. fusca a synonym of H. sulcata a fusca, because

Afzelius recognized H. sulcata with “sulcis regularibus,’ ie., only longitudinal

ribs on stipe (Afzelius 1783, Tab. X, fig.1; this figure is also in Dissing (1966b,

fig. 27c), while H. fusca and H. lacunosa have longitudinal and transverse ribs

(lacunose stipe) [i-e., in the words of Afzelius: “lacunis sulcisque irregularibus” ].

DNA sequences support H. sulcata as a distinct species (unpublished data),

contrary to the opinions by Dissing (1966b) and Abbott & Currah (1997), who

considered it a synonym of H. lacunosa. Persoon (1801) described H. sulcata

a fusca with “pileo nigro.’ Helvella fusca has more recently been reported from

Europe, North America, and Asia (Dissing 1966b, Ceccaldi 2006, Kempton &

Wells 1970; Kaushal 1991). Abbott & Currah (1997) doubted the existence of

H. fusca in North America because their study indicated that such reports were

probably H. maculata N.S. Weber. On the other hand, Kaushal (1991) recorded

H. fusca from the Himalayas, but as his description mentioned hyphal fascicles

of the apothecium measuring < 105 um long, it is unclear whether that report

corresponds to H. fusca. Based on the above information, we can say that its

distribution outside of Europe remains doubtful.

Acknowledgments

The authors thank John McNeill and Walter Gams, who have served as pre-submission

reviewers. We also wish to acknowledge the efforts of the organizers of the July 2011 VII

Congress of Latin American Mycology for providing an opportunity to meet each other

and to begin a collaboration on studies of Helvella and other operculate discomycetes.

The senior author thanks Laura Guzman-Davalos for her support and direction in his

Doctoral studies and for revision of a preliminary version of this paper, and thanks

Anders Tehler, Curator of the Herbarium S, for the loan of Bresadola’s specimen. Virginia

Ramirez Cruz (Universidad de Guadalajara, Mexico) is acknowledged for her valuable

help inking the drawings. We both appreciate those responsible for the digital library of

the Real Jardin Botanico de Madrid (http://bibdigital.rjb.csic.es/spa/index.php) as well

Helvella fusca, lecto- and epitypified ... 437

as the Internet Archive (http://www.archive.org/) for uploading invaluable literature for

taxonomists on their websites.

Literature cited

Abbott SP, Currah RS. 1997. The Helvellaceae: systematic revision and occurrence in northern and

northwestern North America. Mycotaxon 62: 1-125.

Afzelius A. 1783. Svamp-Slagtet Helvella. K. Vetensk.-Acad. Handl. 4: 299-313.

Boudier [JL]E. 1904. Liste préliminaire des noms des 600 espéces de champignons qui doivent étre

figurés dans les six séries de Boudier, Icones Mycologice. 4 pp.

Boudier [JL]E. 1907. Histoire et classification des Discomycétes d’Europe. Librairie des Sciences

Naturelles. Paris. 223 pp.

Boudier [JL]E. 1910. Icones mycologicze ou iconographie des champignons de France. Tome I],

planches 194-421. Librairie des Sciences Naturelles. Paris.

Boudier [JL]E. 1911. Icones mycologicee ou iconographie des champignons de France. Tome IV,

texte descriptif. Librairie des Sciences Naturelles. Paris. 362 pp.

Bresadola G[J]. 1900 [“1892”]. Fungi tridentini novi, vel nondum delineati, descripti, et iconibus

illustrati, ser. 2, fasc. 14, pp. 83-118, plates 196-217. Tridentini.

Bresadola G[J]. 1932. Icones Mycologicae et Supplementum, [eds. Traverso JB, Catoni G, Fenaroli

L, Trener JB] vol. 24 (Tab. 1151-1200). Medeolani.

Ceccaldi JB. 2006. Helvella fusca, une Helvelle (champignon) récemment observée en Gironde (SW

France) pour la premiere fois. Bull. Soc. Linn. Bourdeaux 141, n. 34(2): 135-139.

Dissing H. 1966a. A revision of collections of the genus Helvella L. ex Fr. emend. Nannf. in the

Boudier Herbarium. Rev. Myc. 31: 189-224.

Dissing H. 1966b. The genus Helvella in Europe with special emphasis on the species found in

Norden. Dansk Botanisk Arkiv 25(1): 1-172.

Gillet CC. 1879. Champignons de France. Les discomycetes, livr. 1, pp. [1]-28. Alencon, Paris.

Kaushal SC. 1991. Systematics of N.W. Himalayan species of Helvella (operculate discomycetes).

In: Khullar SP, Sharma MP (eds.). Himalayan botanical researches. Ashish Publishing House,

New Delhi. Pp. 61-75, pl. I-VII.

Kempton PE, Wells VL. 1970. Studies on the fleshy fungi of Alaska IV. A preliminary account of the

genus Helvella. Mycologia 62(5): 940-959. http://dx.doi.org/10.2307/3757607

Korf RP. 1986. A compendium of acceptable names for species illustrated in volumes 2 and 3 of

Boudier’s Icones Mycologicae [with French translation by Jean-Daniel Pellet], pp. 209-252, in:

van Brummelen J, Korf RP, Clémencon H, Jiilich W, Demoulin V. Icones Mycologicae par Emile

Boudier, Tome V, Liste préliminaire & explication des planches [reprint edition]. Editions

Piantanida, Lausanne.

Korf RP. 1988. A compendium of acceptable names for species illustrated in volumes 2 and 3 of

Boudier’s Icones Mycologicae, Annotated: errata, some necessary changes in citations, and

suggested changes based on recent taxonomic information. Ithaca: Privately printed. 48 pp.

McNeill J, Turland NJ. 2011. Synopsis of proposals on botanical nomenclature - Melbourne 2011.

A review of the proposals concerning the International Code of Botanical Nomenclature

submitted to the XVIII International Botanical Congress. Taxon 60: 243-286.

McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nicolson

DH, Prado J, Silva PC, Skog JE, Wiersema JH, Turland NJ. 2006. International

Code of Botanical Nomenclature (Vienna Code). Regnum Vegetabile 146. 568 pp.

(http://ibot.sav.sk/icbn/main.htm).

438 ... Landeros & Korf

McNeill J, Turland NJ, Monro A, Lepschi BJ. 2011. XVIII International Botanical Congress:

preliminary mail vote and report of Congress action on nomenclature proposals. Taxon 60:

1507-1520.

Pelé P, Chenantais JE. 1921. Contribution a la flore mycologique de la Loire-Inférieure. Bull. Soc.

Sci. Nat. Ouest., Sér. 4, 1: 71-115.

Perry G. 2010. (215-222) Proposals on original material. Taxon 59: 1909-1910.

Persoon CH. 1801. Synopsis methodica fungorum 2: i-ii, 241-706. Henricus Dieterich, Gottingen,

Germany.

Ross R. 2002. Which illustrations are original material? Taxon 51: 523-524.

http://dx.doi.org/10.2307/1554867

MYCOTAXON

http://dx.doi.org/10.5248/119.439

Volume 119, pp. 439-443 January-March 2012

Herpothallon weii, a new lichen from China

Yu-LIANG CHENG’, JING NING’, Hou-PinG Xv’, Lu-Lu ZHANG’,

Hal-YING WANG™ & ZUN-TIAN ZHAO™*

‘College of Life Sciences, Shandong Normal University, Jinan, 250014, P. R. China

*Jinan Institute of Metrological Verification, Jinan, 250002, P. R. China

* CORRESPONDENCE TO: “lichenwhy@yahoo.com.cn, ° ztzhao@sohu.com

ABSTRACT — A new species, Herpothallon weii, is described from China. This species is

characterized by a byssoid whitish green thallus tightly attached to the substratum, pinkish

pseudisidia, and the presence of psoromic acid and unknown substances.

Key worps — biodiversity, Arthoniaceae, Cryptothecia, taxonomy

Introduction

Herpothallon (Arthoniaceae, Arthoniales, Arthoniomycetes, Ascomycota) was

established by Tobler in 1937 and reinstated by Aptroot et al. (2009). The lichen

genus is characterized by a crustose thallus without upper cortex, a byssoid

prothallus, and a trentepohlioid photobiont. Herpothallon species often have

pseudisidia or pseudisidioid structures, calcium oxalate crystals, pigments,

and various lichen substances, but usually lack ascocarps. Cryptothecia species,

which are morphologically similar to Herpothallon species, have an I+ blue

medulla. Of Herpothallon, only six species (H. biacidum, H. flavominutum,

H. granulosum, H. isidiatum, H. philippinum, and H. rubroechinatum) have an

I+ blue medulla (Aptroot et al. 2009, Frisch et al. 2010a,b, Jagadeesh & Sinha

2009, Jagadeesh et al. 2009).

Herpothallon includes 35 species worldwide (Aptroot et al. 2009, Frisch et

al. 2010a,b, Jagadeesh & Sinha 2009, Jagadeesh et al. 2009). In China, three

Herpothallon species, H. echinatum, H. granulare, and H. philippinum have

been reported (Aptroot et al. 2009). During our study of the lichen flora of Mt.

Leigong in Guizhou province, China, a species of Herpothallon new to science

was found.

Materials & methods

The specimens studied were collected from Guizhou, China, and are preserved

in SDNU (Lichen Section of Botanical Herbarium, Shandong Normal University).

440 ... Cheng & al.

The morphological and anatomical characters of the specimens were examined using

a stereo microscope (COIC XTL7045B2) and a polarizing microscope (OLYMPUS

CX41). Lichen substances in the specimen cited were identified using the standardized

thin layer chromatography techniques (TLC) with C system (Orange et al. 2010). Photos

of the thalli were taken using a Olympus SZX12 with DP72.

Taxonomic description

Herpothallon weii Y.L. Cheng & H.Y. Wang, sp. nov. Figs 1-2

MycoBank MB 563539

Differs from Herpothallon albidum by pinkish pseudisidia and lack of pigmentosin D.

Type: China. Guizhou province, Mt. Leigong, on bark, alt. 1000 m, 2 Nov. 2009, Q. Tian

20103366. (Holotype in SDNU).

Erymo.oey: In honour of the 80" birthday (6 Nov. 2011) of our dear teacher, Prof.

Jiang-Chun Wei, who as the ‘father of lichenology in China has made a preeminent

contribution to the development of lichenology and the education of Chinese talent in

this field.

Thallus corticolous, crustose, tightly attached, < 1.5 cm in diam., without

upper cortex, distinctly byssoid, whitish green, with irregular calcium oxalate

crystals. Prothallus distinct, < 0.5 mm wide, white, byssoid, composed of

radiating hyphae. Pseudisidia (also called pseudisidioid structure) numerous,

pinkish, irregularly cushion-shaped, fluffy-felty with erect hyphae, < 1 x 0.5

mm. Photobiont cells single or a few cells aggregated. Apothecia and pycnidia

not seen.

CHEMISTRY — ‘Thallus K+ yellow, C+ yellow, KC+ yellow, P+ yellow, I-,

UV-. Psoromic acid and unknown substances present.

DISTRIBUTION AND SUBSTRATE — Herpothallon weii is known only from the

type locality; on bark.

ADDITIONAL SPECIMEN EXAMINED — CHINA. GUIZHOU PROVINCE, Mt. Leigong, on

bark, alt. 1000 m, 2 Nov. 2009, Q. Tian, 20103174 (SDNU).

CoMMENTS — Other Herpothallon species with pseudisidia and psoromic

acid include H. australasicum, H. echinatum, H. flavominutum, H. globosum,

H. rubroechinatum, and H. albidum. In contrast to H. weii, H. australasicum

has a red thallus with concolorous compact pseudisidia; H. echinatum has

soredioid fragments and concolourous pseudisidia and thallus; H. flavominutum

has a greyish yellow thallus, granular pseudisidia, and lichexanthone,

norlichexanthone, and psoromic acid; H. globosum has a red thallus, globose

pseudisidia, and chiodectonic acid (red pigment); and H. rubroechinatum has

an orange to red thallus, pseudisidia with red hairs, and common pycnidia. The

more closely related H. albidum possesses very similar pseudisidia (also called

pseudisidioid structures) but has white, rather than pinkish, pseudisidia and

possesses pigmentosin D.

Herpothallon weii sp. nov (China) ... 441

Fic. 1. Herpothallon weii (holotype). A. Thallus; B. Prothallus.

Acknowledgements

This research was supported by the City Science Foundation of Guiyang

(20101Zhong05), the National Natural Science Foundation of China (31070010,

31000008), and Natural Science Foundation of Shandong Province (2R2010CQ038).

We thank Prof. A. Aptroot (ABL Herbarium, Soest, The Netherlands) and Prof. Shou-

Yu Guo (Institute of Microbiology, Chinese Academy of Sciences, Beijing, China) for

presubmission reviews.

442 ... Cheng & al.

proe 10.Fe

SsuTyeoy pue

wu cge 1ysTTUNsS

ee ae ee

Ocm1

Fic. 2. Herpothallon weii (holotype).

A. Pseudisidia; B. TLC of H. weii (U: unknown substance; P: psoromic acid).

Herpothallon weii sp. nov (China) ... 443

Literature cited

Aptroot A, Thor G, Licking R, Elix JA, Chaves JL. 2009. The lichen genus Herpothallon reinstated.

Bibliotheca Lichenologica 99: 19-66.

Frisch A, Elix JA, Thor G. 2010a. Herpothallon biacidum, a new lichen species from tropical

Australia. Lichenologist 42(3): 285-289. http://dx.doi.doi/10.1017/S0024282909990697

Frisch A, Thor G, Elix J. 2010b. Herpothallon rubroechinatum (Arthoniaceae), a new species from

tropical and subtropical America. Bryologist 113(1): 144-148.

http://dx.doi.doi/10.1639/0007-2745-113.1.144

Jagadeesh Ram TAM, Sinha GP. 2009. New species and new records of Herpothallon (lichenized

Ascomycota) from India. Mycotaxon 110: 37-42. http://dx.doi.doi/10.5248/110.37

Jagadeesh Ram TAM, Sinha GP, Singh KP. 2009. New species and new records of Cryptothecia and

Herpothallon (Arthoniales) from India. Lichenologist 41(6): 605-613.

http://dx.doi.doi/10.1017/S0024282909008123

Orange A, James PW, White FJ. 2010. Microchemical methods for the identification of lichens. 2nd

edition. London: British Lichen Society.

MY COTAXON

http://dx.doi.org/10.5248/119.445

Volume 119, pp. 445-451 January-March 2012

Four new records of lecideoid lichens from China

Lu-Lu ZHANG’, LI-SONG WANG’,

Hat-YING WANG" & ZUN-TIAN ZHAO

‘College of Life Sciences, Shandong Normal University Jinan, 250014, P. R. China

*Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany,

Chinese Academy of Science, Kunming, 650204, P. R. China

*CORRESPONDENCE TO: *lichenwhy@yahoo.com.cn, °ztzhao@sohu.com

ABSTRACT — Four lecideoid lichen species, Carbonea vorticosa, Lecidea diducens, L.

promiscens, and Lecidella bullata, are reported for the first time from China.

Key worps — Yunnan, Xizang, Lecanoromycetes, taxonomy

Introduction

As once one of the largest lichen genera, Lecidea s. lat. has had an

extraordinarily broad circumscription, with Zahlbruckner (1926) accepting

more than 1350 species. Subsequently, Carbonea, Lecidella, and many other

obviously more natural units have been excluded based on the ascomal structure,

especially the nature of the hamathecial tissues, ascus apical structures, and

exciple (Hertel 1977, 1995; Smith et al. 2009). Lecidea s. str. has now become

a medium-sized genus that lacks algae in the exciple and has a Lecidea-type

ascus (Hertel 1995, Pérez-Ortega & Etayo 2008). Carbonea is characterized by

a “carbonized” exciple and Lecanora-type ascus (Hertel 1983, Nash et al. 2004).

Lecidella is characterized by Lecanora-type ascus and discrete paraphyses (Nash

et al. 2004).

Worldwide, Carbonea includes 20 known species, Lecidea s. str. about 100

species, and Lecidella 79 species (Kirk et al. 2008). In China, these lecideoid

lichens are still poorly known with only two Carbonea, 10 Lecidella, and 14

Lecidea s. str. species previously reported (Wei 1991; Abass & Wu 1998; Aptroot

2002, 2003; Obermayer 2004; Guo 2005; Zhang et al. 2010). During our study

of lecideoid lichens from China, four species new to the country —Carbonea

vorticosa, Lecidea diducens, L. promiscens, and Lecidella bullata— have been

identified.

446 ... Zhang & al.

Materials & methods

The specimens studied were collected from Western China, and are preserved

in SDNU (Lichen Section of Botanical Herbarium, Shandong Normal University),

HKAS (Herbarium of Cryptogams, Kunming Institute of Botany Academia Sinica), or

HMAS-L (Lichen Section, Herbarium of Mycology, Institute of Microbiology, Academia

Sinica). Specimen morphology and anatomy were examined using a stereo microscope

(COIC XTL7045B2) and a polarizing microscope (Olympus CX41). Lichen substances

were identified using standardized thin layer chromatography techniques (TLC) with

C and J system (Orange et al. 2010). Photos of the thalli were taken using a Olympus

SZX12 with DP72.

The new records

Carbonea vorticosa (Florke) Hertel, Mitt. Bot. Staatssamml. Miinchen 19: 442

(1983) Fig. 1A

Thallus crustose, indistinct, endolithic to areolate, thin; prothallus absent to

not obvious; surface grey or white, rough. Apothecia sessile, up to 1.0 mm in

diam.; disc black, plane to a little convex, epruinose; margin black, flexuous in

older apothecia, long persistent; exciple blackish green to dark brown, hyphae

radiating, 3.5-5 um; epihymenium bluish green; hymenium hyaline with

some green, 35-60 um; hypothecium black to brown; asci clavate, 8-spored;

ascospores hyaline, simple, oblong-ellipsoid, 10-13.5 x 3-5 um.

SPOT TESTS — K-, C-, KC-, P-.

SECONDARY METABOLITES — none detected.

SPECIMENS EXAMINED — CHINA. YUNNAN: DEQIN couNTY, Mt. Baimaxueshan,

alt.4800 m, on rock, 13 Jul. 1981, L.S. Wang 12786 (HKAS); Lijtane city, Mt. Laojun,

alt. 4000m, on rock, 7 Nov. 2009, Y.L. Cheng 20100337-1, 20100289 (SDNU).

CoMMENTS — Carbonea vorticosa is closely related to C. capsulata, which

also produces small apothecia, a blue epithecium, and a brown hypothecium.

However, C. capsulata has a more conspicuously blue hymenium and broader

(5-10 um diam.) excipular hyphae.

Reported from Antarctica, Asia, Australasia, Europe, and North and South

America (Hertel 1991, Galloway 2007, Nash et al. 2004, Smith et al. 2009),

C. vorticosa is new to China.

Lecidea diducens Nyl., Flora 48: 148 (1865) Fic. 1B

Thallus lacking or very indistinct, grey, not continuous; prothallus indistinct;

medulla I+ deeply violet. Apothecia black, sessile, constricted at the base, up to

1.4 mm in diam.; disc black, flat to slightly convex, dull to shiny, epruinose;

margin well developed, persistent; exciple with a blackish to dark brown rim and

paler to hyaline interior, C+ red, K-; epihymenium blackish green; hymenium

45-50 um, very pale green to hyaline; paraphyses simple, occasionally branched

and anastomosing; subhymenium somewhat more intensely green than the

Lecideoid lichens new to China... 447

CO PHF E.

rae LS Ba

; am * “y

aA 4 ‘

oe od al

nO & ’

Oe Se 2

Fic. 1 A. Carbonea vorticosa (L.S. Wang 12786, HKAS);

B. Lecidea diducens (H.Y. Wang 20100297, SDNU).

448 ... Zhang & al.

hymenium, 10-30 um thick; hypothecium pale brown to brown; asci clavate,

8-spored; ascospores simple, oblong-ellipsoid, 7.5-11 x 3.5-4 um.

SPOT TESTS — K-, C-, KC-, P-.

SECONDARY METABOLITES — 2’-O-methylanziaic acid.

SPECIMENS EXAMINED — CHINA. YUNNAN: LIJIANG COUNTY, Mt. Laojun, alt. 4000 m,

on rock, 7 Nov. 2009, H.Y. Wang 20100332, 20100297 (SDNU).

ComMENTS — Lecidea diducens is morphologically similar to L. auriculata,

which lacks confluentic acid in the exciple and so is C-.

Probably cosmopolitan, L. diducens has been reported from Europe, North

and South America, Asia, and Australasia (Hertel 1991, 2006, Hertel & Andreev

2003, Nash et al. 2004, Smith et al. 2009). New to China.

Lecidea promiscens Nyl., Flora 55: 358 (1872) Fic. 2A

Thallus grey, thin, not continuous; prothallus indistinct; medulla white, I+

deeply violet. Apothecia black, sessile, 0.8-1.8 mm in diam., disc black, flat

to slightly convex, epruinose; margin black, well developed, persistent, often

undulate; exciple with a thin blackish rim and hyaline to light brown interior;

epihymenium greenish black to black; hymenium hyaline to pale green,

50-62 um tall, I+ blue; paraphyses simple, occasionally branched, anastomosing;

subhymenium hyaline to light brown; hypothecium brown to dark brown;

asci clavate, 8-spored; ascospores hyaline, simple, oblong to oblong-ellipsoid,

7.5-12 x 4—4.5 um.

SPOT TESTS — K-, C-, KC-, P-.

SECONDARY METABOLITES — confluentic acid.

SPECIMEN EXAMINED — CHINA. YUNNAN: DEQIN county, Meilishi village, Suola

Yakou, alt. 4700 m, on rock, 30 Aug. 2009, L.S. Wang 00-19783 (HKAS).

ComMENTS — Lecidea promiscens is very similar to L. auriculata but has broader

and larger spores. It also has a less prominent exciple (in L. auriculata the exciple

extends quite far under the hypothecium), and a darker hypothecium.

Reported from Europe, North and South America, Asia, and Australasia

(Thomson 1997, Hertel 1991, 2006, Nash et al. 2004), L. promiscens is new to

China.

Lecidella bullata Korb., Parerga Lichenol.: 200 (1861) Fig. 2B

Thallus crustose, aerolate or bullate, moderately thick, grey-white to yellow-

white, without soredia or isidia; prothallus absent. Apothecia sessile to semi-

immersed, up to 1.5 mm in diam., disc black, flat to slightly convex, greyish

pruinose to epruinose; margin black, at first prominent, later excluded; exciple

greenish black outside and hyaline interior; epihymenium blue-green to

black-green; hymenium hyaline, 40-55 um tall, I+ blue; paraphyses simple;

subhymenium and hypothecium hyaline, with crystals; asci Lecanora-type,

Lecideoid lichens new to China... 449

4 ne SEPA ‘a

Fic. 2 A. Lecidea promiscens (L.S. Wang 00-19783, HKAS);

B. Lecidella bullata (J.C. Wei X024916, HMAS-L).

450 ... Zhang & al.

clavate, 8-spored; ascospores hyaline, simple, ellipsoid to broad-ellipsoid,

7-12 X 5-6 um.

SPOT TESTS — K+ yellow, C-, KC+ yellow, P+ yellow.

SECONDARY METABOLITES — atranorin, xanthones, + psoromic acid, + zeorin.

SPECIMENS EXAMINED — CHINA. YUNNAN: DEQIN couNTY, Mt. Baimaxueshan, alt.

4500 m, on rock, 25 May 1985, L.S. Wang 8909(HKAS); X1zane: Mt. Qomolangma,

Zhongrongbu, alt. 5550 m, on rock, 28 May 1966, J.C. Wei X024916 (HMAS-L).

Comments — Lecidella bullata is very similar to L. stigmatea, which also has

a colorless internal excipulum and subhymenium. However, L. stigmatea has a

very thin, rimulose or scattered verrucose thallus and no psoromic acid.

Reported from the Arctic, Europe, Asia, and North America (Inoue 1997,

Thomson 1997), L. bullata is new to China.

Acknowledgements

The project was financially supported by the National Natural Science Foundation

of China (31170187, 31070010, 31000008). The authors would like to thank the keeper

of the HMAS-L, Ms Deng Hong for assistance during this study. The authors thank

Dr. A. Aptroot (ABL Herbarium, Soest, The Netherlands) and Prof. Shou-Yu Guo (Key

Laboratory of Systematic Mycology & Lichenology, Institute of Microbiology, Chinese

Academy of Sciences, Beijing, China) for presubmission reviews.

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Zhang LL, Wang HY, Sun LY, Zhao ZT. 2010. Four lichens of the genus Lecidea from China.

Mycotaxon 112: 445-450. http://dx.doi.org/10.5248/112.445

MY COTAXON

http://dx.doi.org/10.5248/119.453

Volume 119, pp. 453-457 January-March 2012

Puccinia, Uromyces, and Xenodochus species new to Turkey

SANLI KABAKTEPE & ZELIHA BAHCECIOGLU

Department of Biology, Faculty of Science and Art, Inonu University, TR 44280, Malatya, Turkey

*CORRESPONDENCE TO: Zeliha. bahcecioglu@inonu.edu.tr

ABSTRACT — The genus Xenodochus and nine rust taxa —Xenodochus carbonarius, Puccinia

hysteriiformis, PB. majoricensis, P. pedunculata, P. stachydis, P. tendae, Uromyces colchici,

U. kochiae, U. veratri— are reported for the first time from Turkey. The morphological and

microscopical features of these fungi are described.

Key worps — Anatolia, Ardahan, Kars, Pucciniales, Uredinales

Introduction

The rust genus Xenodochus, with only seven species worldwide, differs from

Phragmidium and Frommeella by teliospores having one germ pore in the apical

cell and two or more germ pores in all other cells. Puccinia, the largest genus of

Uredinales with about 3000 to 4000 species, has been recorded in all land areas

except the polar regions.

In the last 10 years reports of new taxa and records of rust fungi have greatly

increased in Turkey (Bahcecioglu & Gjzerum 2003, 2004; Bahcecioglu & Yildiz

2005; Kabaktepe & Bahcecioglu 2005, 2006; Bahcecioglu et al. 2005, 2006,

2009; Kirbag 2003, 2004; Kirbag et al. 2011). Although approximately 350 rust

species representing 26 genera have been recorded from Turkey, this number

is far less than the total number of potential host plants, and further studies are

required on rust fungi in Turkey.

This paper presents rust species collected from Kars and Ardahan provinces

in northeast Turkey.

Materials & methods

Materials were collected during 2006-09 from Kars and Ardahan Provinces in Turkey.

The host specimens were prepared according to established herbarium techniques.

Spores were scraped from dried host specimens and mounted in lactophenol. The

preparations were examined with an Olympus CX31 light microscope and micrographs

taken with a Olympus E-330 camera. Analysis LS Starterwas software used to measure

454 ... Kabaktepe & Bahcecioglu

at least 30 spores for each spore state. Host names follow Davis (1965-85) and Davis et

al. (1988). Specimens are deposited in the Inonu University Herbarium (INU), Turkey.

Taxonomy

Xenodochus carbonarius Schltdl., Linnaea 1: 237 (1826)

Telia amphigenous, pulvinate, 1-2 mm diam, black. Teliospores elongate-

cylindrical, often curved, 6-15 celled, rounded at the both ends, strongly

constricted, smooth but subverruculose at the apex, 97-250 x 24-28 um, each

cell with 2 opposite superior pores, the uppermost with one apical pore with

a small hyaline papilla, wall brown in basal cells often nearly colorless, 2 um

thick, pedicel short, colorless, persistent.

Known on Sanguisorba officinalis from Asia, Europe, and North America;

X. carbonarius is a new rust species and Xenodochus a new genus for Turkey.

SPECIMENS EXAMINED - On Sanguisorba officinalis L. (Rosaceae): TURKEY: ARDAHAN,

30 km from Cildir to Hanak, 1950 m, 10.07.2006, Sanli Kabaktepe 4109 (INU 8534); 20

km Cildir to Ardahan, 1880 m, 23.09.2006, Sanli Kabaktepe 4732 (INU 8535).

Puccinia hysteriiformis Peck, Bot. Gaz. 6: 276 (1881)

Telia amphigenous, scattered, covered by the epidermis, < 1 mm diam,

brown. Teliospores oblong or oblong-clavate, 40-45 x 19-23 um, constricted

at the septum, pores usually indistinct, pore of upper cell apical, pore in lower

cell near the septum, wall 1.5-2.5 um, at apex < 4-9 um, pale brown, smooth,

pedicel < 106 um, persistent, hyaline.

Known from Europe, North America, and South America; P hysteriiformis

is a new rust species for Turkey.

SPECIMEN EXAMINED - On Minuartia recurva (L.) Schinz & Thell. (Caryophyllaceae).

TURKEY: Kars, Duman Mountain, 2000 m, 08.07.2006, Sanli Kabaktepe 4023 (INU

8537).

Puccinia majoricensis Maire, Bull. Soc. Myc. Fr. 21: 221 (1905)

Telia hypophyllous on stem and petioles, < 1 mm diam, scattered or in groups,

dark brown. Teliospores ellipsoid, oblong, 38-59 x 14-17 um, constricted at

the septum, pore of upper cell apical, wall 1-1.5 um, at apex up to 5-8 um,

yellowish brown, at apex pale brown, smooth, pedicel < 90 um, persistent, pale

yellow or hyaline.

Known from Asia and Europe, P majoricensis is a new rust species for

Turkey.

SPECIMENS EXAMINED — On Teucrium chamaedrys L. (Lamiaceae). TURKEY: ARDAHAN,

Posof, Dogrular village, 1600 m, 28. 08. 2008, Sanli Kabaktepe 6551(INU 8538); 13-15

km from Ardahan to Gole, 1850 m, Sanli Kabaktepe 6836 (INU 8539).

REMARKS —Puccinia annularis (F. Strauss) G. Winter on Teucrium chamaedrys

and T! polium L. (Bahcecioglu & Yildiz 2005; Kabaktepe & Bahcecioglu 2006;

Rust fungi new to Turkey ... 455

Bahcecioglu et al. 2006) and P constricta (Lagerh.) Bubak on Teucrium orientale

L. (Bahcecioglu & Gjzerum 2004) have also been reported from Turkey. Puccinia

majoricensis differs from P. annularis by dark colored teliospores, and differs

from P. constricta by a bigger and hyaline pedicel.

Puccinia pedunculata J.Schrot., Jber. schles. Ges. vaterl. Kultur 52: 117 (1875)

Uredinia hypophyllous, covered by the epidermis, roundest, scattered,

reddish brown. Urediniospores ellipsoid, ovoid, 28-35 x 20-24 um, wall 2.5-3

um thick, pale brown, aculeate, 5-6 scattered pores. Telia hypophyllous, on

stems, covered by the epidermis, dark brown. Teliospores ellipsoid, oblong or

oblong-clavate, the upper cell broader, rounded or slightly attenuate, 41-58 x

20-26 um, slightly or not constricted at the septum, pore of upper cell apical,

wall smooth, chestnut brown, 1.5-2.5 um, at apex 5-9 um, pedicel < 80 um,

persistent, brownish.

Known from Europe, P. pedunculata is a new rust species for Turkey.

SPECIMEN EXAMINED - On Rumex scutatus L. (Polygonaceae). TURKEY: Kars,

Sarikamis, Catak village, 1750 m, 24.08.2009, Sanli Kabaktepe 6934 (INU 8540).

REMARKS — Puccinia acetosae (Schumach.) Korn. on Rumex scutatus,

R. tuberosus L., and R. acetosella L. have been reported from Turkey. (Bahcecioglu

& Yildiz 2005; Bahcecioglu et al. 2006; Tamer 1978). Puccinia pedunculata

differs from P. acetosae by not being constricted at the septum and lacking a

papilla on the upper teliospore cell.

Puccinia stachydis DC., Fl. franc. 2: 595 (1805)

Uredinia amphigenous, rounded, scattered, brown. Urediniospores ellipsoid,

globoid, 25-28 x 24-26 um, wall 2-3 um thick, brown, aculeate with 2-3 pores.

Telia similar to uredinia but dark colored. Teliospores ellipsoid or oblong,

37-47 x 19-23 um, constricted at the septum, pore of upper cell apical, wall

smooth, chestnut brown, < 6-11 um thick at the apex, pedicel as long as spores,

persistent, hyaline.

Known from Europe, P. stachydis is a new rust species for Turkey.

SPECIMENS EXAMINED - On: Stachys annua L. (Lamiaceae). TURKEY: ARDAHAN,

16-18 km Haskoy to Ardahan, 1850 m, 17.09.2008, Sanli Kabaktepe 6675 (INU 8541).

Kars, Sarikamis, Catak village, 1750 m, 24.08.2009, Sanli Kabaktepe 6935 (INU 8542).;

Sarikamis, 2100-2500 m, 25.08.2009, Sanli Kabaktepe 6996 (INU 8543).

REMARKS —Puccinia hariotii Lagerh. on Stachys spp. (Henderson 1957) and

P. betonicae (Alb. & Schwein.) DC. on S. iberica M. Bieb. and S. citrina Boiss.

& Heldr. (Bahcecioglu & Yildiz 2005; Bahcecioglu & et al. 2006) have been

reported from Turkey. Puccinia stachydis differs from P. hariotii by constriction

at the septum and a persistent pedicel and from P. betonicae by pale colored

teliospores and a persistent pedicel.

456 ... Kabaktepe & Bahcecioglu

Puccinia tendae Gaum., Rev. Mycol. 18: 184 (1953)

Uredinia hypophyllous, rounded, scattered, pale brown. Urediniospores

ellipsoid, globoid, 13-28 x 18-26 um, wall 1.5 um thick, brown, echinulate with

2-3 equatorial pores. Telia similar to uredinia but dark colored. Teliospores

ellipsoid or oblong, 35-50 x 19-22 um, constricted at the septum, pore of

upper cell apical, wall smooth, pale chestnut brown, 2-3 um thick, at apex up

to 9 um thick, pedicel < 100 um, persistent, hyaline.

Known from Europe, P. tendae is a new rust species for Turkey.

SPECIMEN EXAMINED - On Thymus fedtschenkoi Ronniger (Lamiaceae). TURKEY:

Kars, 8 km from Kagizman to Tuzluca, 1140 m, 12. 06. 2007, Sanli Kabaktepe 4989

(INU 8544).

REMARKS —Puccinia serpylli Lindr. on Thymus fallax Fisch. & C.A. Mey.,

T. fedtschenkoi, and T: kotschyanus Boiss. & Hohen. (Henderson 1959, 1964;

Bahcecioglu & Gjzrum 2004; Bahcecioglu et al. 2006) has been reported

from Turkey. Puccinia tendae differs from P serpylli by narrow, dark colored

teliospores.

Uromyces colchici Massee, Grevillea 21: 6 (1892)

Telia amphigenous, scattered, rather large, elliptical, sometimes circinate,

up to 2 mm long, covered for some time by the epidermis, brown. Teliospores

subgloboid to ovoid, rounded above with a flat hyaline papilla, 28-40 x 20-28

um, smooth, pale brown, wall 2.5-3.5 um thick, pedicel deciduous, hyaline.

Known from Europe, U. colchici is a new rust species for Turkey.

SPECIMEN EXAMINED - On Colchicum speciosum Steven (Colchicaceae). TURKEY:

ARDAHAN, Bulbilen Mountain, 2400 m, 18.07.2007, Sanli Kabaktepe 5423 (INU 8545).

Uromyces kochiae Syd. & P. Syd., Annls mycol. 10: 215 (1912)

Uredinia amphigenous, small, scattered, black. Urediniospores globoid,

subgloboid, 21-25 x 21-24 um, wall 1.5-2.5 um thick, yellowish brown,

echinulate with 6-9 pores. Telia similar to uredinia. Teliospores globoid,

ellipsoid, 18-23 x 16-18 um, wall smooth, brown, at apex dark colored, up to

3-5 um thick at apex, pedicel < 100 um, persistent, hyaline.

Known from Asia and Europe, U. kochiae is a new rust species for Turkey.

SPECIMENS EXAMINED — On Kochia prostrata (L.) Schrad. (Chenopodiaceae). TURKEY:

Kars, 20 km from Digor to Igdir, 1500 m, 07.07.2006, Sanli Kabaktepe 3989 (INU 8546).;

Kagizman, Esenkir village, 1220 m, 01.07.2008, Sanli Kabaktepe 6033 (INU 8547).

Uromyces veratri (DC.) J. Schrét., Jber. schles. Ges. vaterl. Kultur 49: 10 (1871)

Uredinia hypophyllous, rounded, scattered, chestnut brown. Urediniospores

ellipsoid, 22-26 x 19-20 um, wall 1-2 um thick, pale brown, aculeate with 1-3

equatorial pores. Telia amphigenous, scattered, pulverulent, up to 2 mm long,

Rust fungi new to Turkey ... 457

covered by the epidermis, blackish brown. Teliospores ellipsoid, ovoid, oblong,

22-30 x 14-18 um, at apex with a flat hyaline papilla, wall smooth, brown,

< 7 um thick at apex, pedicel deciduous, brownish.

Known from Asia and Europe, U. veratri is a new rust species for Turkey.

SPECIMEN EXAMINED — On Veratrum album L. (Melanthiaceae). TURKEY: ARDAHAN,

Posof, Ilgar mountain pass, 2540 m, 26.08.2009, Sanli Kabaktepe 7023 (INU 8548).

Acknowledgments

We are indebted to TUBITAK (Project no. TBAG-106T260) for financial support. We

also thank Dr. Ahmet Asan and Dr. Sevda Kirbag for presubmission expert reviews.

Literature cited

Bahcecioglu Z, Gjzrum HB. 2003. New and rare rust fungi (Uredinales) from Anatolia (Turkey).

Mycotaxon 85: 165-173.

Bahcecioglu Z, Gjarum HB. 2004. New and rare rust fungi (Uredinales) from Anatolia (Turkey)

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23-44.

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Province, Turkey. Turkish J. Bot. 30: 419-434.

Bahcecioglu Z, Berndt R, Kabaktepe S. 2009. Puccinia ardahanensis sp nov., a new rust fungus from

Turkey. Sydowia 61(2): 209-213.

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(supplement). Edinburgh University Press, Edinburgh.

Henderson DM. 1957. Uredinales from Asia Minor. Notes Roy. Bot. Gard. Edinburgh 22:

195-200.

Henderson DM. 1959. Uredinales from S.W. Asia. Notes Roy. Bot. Gard. Edinburgh 23: 71-83

Henderson DM. 1964. Uredinales from S.W. Asia III. Notes Roy. Bot. Gard. Edinburgh 25:

197-277.

Kabaktepe S, Bahcecioglu Z. 2005. Seven rust species recorded as new to Turkey. Mycotaxon 91:

393-396.

Kabaktepe S, Bahcecioglu Z. 2006. Microfungi identified from the flora of Ordu Province in Turkey.

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445-448.

Kirbag S, Aime MC, Kursat M. 2011. A new Puccinia on Thymelaea from Turkey. Mycotaxon 115:

501-504. http://dx.doi.org/10.5248/115.501

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MY COTAXON

http://dx.doi.org/10.5248/119.459

Volume 119, pp. 459-465 January-March 2012

Pseudobaeospora deckeri sp. nov.—

a new agaric from central California

CHRISTIAN SCHWARZ

The Brat Pack, 301 Van Ness Avenue, Santa Cruz, CA, 95060, USA

CORRESPONDENCE TO: cfs.myko@gmail.com

ABSTRACT — Pseudobaeospora deckeri is described from the central California coast. This

new species is characterized by fruitbodies with deep purple to lilac tones, a pileipellis with

irregularly shaped erect elements, an undifferentiated subpellis, absence of cheilocystidia,

and strong alkaline-virescent reactions in the pileus subpellis and stipe context.

Key worps — Agaricales, biodiversity, fungal taxonomy, Coast Redwood, Monterey Cypress

Introduction

The genus Pseudobaeospora Singer has recently undergone a generic revision,

spurred by the recognition and description of many new taxa (Bas 2002, 2003).

Following this revisitation, novel Pseudobaeospora species were discovered

in Europe and California (Adam¢cik & Bas 2002, Desjardin 2004, Vellinga

2009). Two of the more recently described species are from the central coast of

California: P stevensii Desjardin (Desjardin 2004), from cypress and redwood

habitats in Marin and San Mateo counties, and P. aphana Vellinga (Vellinga

2009), from Marin County. Considered with Tricholoma microsporum (Ellis)

Sacc. (not yet formally assigned to Pseudobaeospora), these species represent

the second and third Pseudobaeospora species reported from California and the

fifth and sixth species reported from North America.

Following a cold precipitation event in March of 2009, many redwood-

associated macrofungi (Hygrocybe spp., Entoloma spp.) fruited on the University

of California Santa Cruz (UCSC) campus. While surveying this assemblage, I

encountered a small purple agaric for which the generic disposition was not

immediately obvious. While investigating the fruitbodies closely, I collected a

range of representative specimens in January 2010, when the same mycelium

produced over forty fruitbodies. In December 2010, while surveying macrofungi

of the California coast with Erin Page Blanchard and Noah Siegel, I collected

460 ... Schwarz

specimens from two additional, relatively distant, sites on the central California

coast. A striking member of the Central California mycobiota, P deckeri

represents the fourth described species of Pseudobaeospora in California and

the seventh in North America.

Materials & methods

The macromorphology was described from fresh material. Micromorphological

features were observed from both fresh and dried material. Standardized color notations

were made using Kornerup & Wanscher (1984). Specimens were sectioned using a Nikon

SMZ-10A dissecting microscope and mounted in 3% KOH with and without staining

by aqueous Congo Red or Melzer’s reagent. Micromorphological features were observed

with a Nikon Eclipse E800 light microscope. Lamellae and lamellulae are denoted as L

and l, respectively. Spore measurements are given with the arithmetic means denoted as

L,,, x W,,., DNA was extracted with Sigma Extraction Kit- REDExtract-N-Amp Plant

PCR using standard procedures; ITS1F and ITS4 primers were used for PCR reactions,

and cleanup was performed with Exosap. Sequencing reactions used Big Dye v3.1 at

1/8 strength of manufacturer’s instructions, on a 3730 Sequencer (Applied Biosystems,

Foster City, California). The resulting sequence was edited and assembled with

Sequencher 4.2.2 (Gene Codes Corp., Ann Arbor, Michigan) and deposited in Genbank

(JF898319).

Taxonomy

Pseudobaeospora deckeri C.F. Schwarz, sp. nov. FIGs 1-2

MycoBank MB 561249

Similar to Pseudobaeospora pyrifera but pileipellis with erect hyphae, cap and stipe with

slightly different colors and KOH reactions, and without cheilocystidia.

Ho.otype: USA. CALIFORNIA: SANTA CRUZ COUNTY, Santa Cruz, UC Santa Cruz,

north side of Engineering 2 Building, 12 Jan 2010, CS 12Jan2010-1 (Holotype UCSC

7451; Genbank JF898319).

ErymMo.Loecy: Named in honor of Lee Decker, instructor of biology and ocean science,

whose passion has inspired generations of students.

FruiTsopies collybioid, entirely rubbery-tough to fleshy and _ slightly

cartilaginous when fresh, brittle in age, but rather persistent. PILEUs 10-27 mm

wide, broadly convex and circular to plano-convex or uplifted and irregularly

wavy; margin of young specimens distinctly but narrowly involute and in

some specimens distinctly ribbed; very finely pruinose, evenly or in zones

or blotches, especially near margin, in age nearly glabrous; deep royal purple

(16E-F8) when young and moist, fading to dull brownish or grayish purple

(14F4-6), sometimes in age with areas of obscure orange-brown tones (5D5),

at margin paler to nearly white. Flesh thin, pallid lilac to darker purple just

above lamellae. LAMELLAE L = 22-38(50), 1 = 3-5, subdistant to fairly close,

adnate to finely and obscurely sinuate, ventricose in age, in some specimens

intervenose and transvenose, when young dull gray (8B1-2) to dull lilac (13E3),

Pseudobaeospora deckeri sp. nov. (USA) ... 461

FIG 1. Pseudobaeospora deckeri.

Basidiocarps. NS 14Jan2010. Photo by Noah Siegel.

A Qae@

FIG 2. Pseudobaeospora deckeri.

A. Spores. B. Caulocystidia. C. Pileipellis. Bars: A = 3.5 um, B-D = 30 um.

462 ... Schwarz

in age distinctly ochre brownish (6C-D8); margin often thick, even to slightly

irregular. Stipe (13-)29-50(-85) x 2-4.5 mm, equal or nearly so, at apex

with belts or zones of fine, pale squamules, sometimes quite dense, with a fine

pruina below these squamules over the upper quarter of the stipe, especially

visible in young specimens, very dark purple (15F8) to royal purple (16D7-8)

more or less remaining so at apex, but lower portions becoming duller purple

to reddish-brown (11F7, 12D6, 12F4), at base usually conspicuously strigose,

this tomentum white to lilac or purplish; base slightly rooting and bound to

particles of substrate; flesh solid, dull, grayish purple, cartilaginous. ODOR

indistinct, although dried specimens wetted with alcohol emitted a ‘mushroomy’

odor (like Agaricus bisporus). TASTE weakly but distinctly peppery or acrid.

MACROCHEMICAL REACTIONS instantly dark blue-green in 3% KOH, this

reaction obscured by the dark color of the fruitbodies and thus best detected by

swabbing the treated fruitbody on white paper, or immersing a thin section in

KOH. Spore Deposit white.

BASIDIOSPORES L x W = (2.8)3.6-3.8(4.8) x (2.8)3.3-3.5(3.8) um. Q = (1.0)

1.05-1.1(1.3). A total of 40 spores from two fruitbodies were examined

microscopically: i) (n = 20; from stipe apex, in Melzer’s reagent) 2.8-3.8

x 2.8-4(4.8) um, Le, x Was = 3.6 x 3.3 um, Q = 1.0-1.3, Oe = 1.1, ii) @=20,

from stipe apex, in Melzer’s reagent) 2.9-4.3 x 2.9-3.8 um, L,,, x W,,, = 3.7 x

3.5 pm, Q,,, = 1.05. Globose to subglobose or broadly ellipsoid, some with

visible internal droplet; with conspicuous hilar appendage. Morphology and

chemical characters of spores variable within a well defined range: immature

spores smooth, thin or thick-walled and inconsistently dextrinoid, more

mature spores dextrinoid within 5 minutes, thick-walled and smooth. BAsIDIA

24-30 x 3.5-5 um, clamped at base but obscurely so, tetrasporic, infrequently

bisporic, protruding only slightly from lamellar face. LAMELLAR TRAMA regular,

composed of inflated elements 43-80 x 5-28 um, barrel shaped and tapered at

both ends, clamped. Walls irregularly and coarsely thickened. CystTip1A absent,

but irregularly shaped basidioles at the lamellar margin were inconsistently

present. PILEIPELLIs fundamentally a cutis of equal to slightly irregularly inflated

hyphae 4.3-8.6 um in diam., but with many erect elements, thus appearing as

a trichoderm, remaining purplish in KOH; erect elements 16-58 x 3.6-6.7

uum, cylindrical to cylindrical-flexuous, often irregularly swollen, constricted

or knobbed, often once or twice septate and clamped, often slightly entangled

or clustered, more scattered away from disc, not encrusted. Subpellis of tightly

interwoven hyphae, not or obscurely differentiated from pileal trama, distinctly

green in KOH. PILEus TRAMa parallel, made up of narrow, non-encrusted

clamped hyphae. STIPITIPELLIS at apex with a dense coating of clustered

caulocystidia over a layer of narrow hyphae 2.8-5 um in diam., attenuated over

upper part of stipe. CAULOCYSTIDIA 29-40 x 2.9-4.8 um, cylindrical-flexuous

Pseudobaeospora deckeri sp. nov. (USA) ... 463

to strangulate or irregularly swollen and constricted or knobby, often once-

septate and clamped; green in KOH (3%). Stipe TRama of inflated hyphae

< 8.6-14 um in diam., pseudoparenchymatous in cross-section, strongly green

in KOH. CLAMP CONNECTIONS abundant in all tissues.

ECOLOGY & DISTRIBUTION — Subcaespitose or scattered, terrestrial. Fruiting

at the type locality amongst woodchip mulch in a landscaped area under coast

redwood (Sequoia sempervirens). Currently known from three locations: the

type locality (University of California Santa Cruz Campus, Santa Cruz Co.),

Redwood Camp (Monterey Co.), and Skyline Community College (San Mateo

Co.). At the southernmost known locality (Santa Cruz Co.) fruiting near

huckleberry (Vaccinium ovatum), red alder (Alnus rubra), and coast redwood,

while at the northernmost known locality (San Mateo Co.) fruiting in deep

needle duff of Monterey cypress (Hesperocyparis macrocarpa). Fruiting dates

include records from early December, January, and March.

ADDITIONAL COLLECTIONS EXAMINED: USA. CALIFORNIA: SANTA CRUZ County, UC

Santa Cruz, north side of Engineering 2 Building, 5 Mar 2009, CS 5Mar2009-7 (type

mycelium); SAN MATEO Counry, San Bruno, Skyline Community College, 14 Jan 2010,

NS 14Jan2010-3; MONTEREY County, Big Sur, UC Big Creek Reserve, Redwood Camp,

13 Dec 2010, CS 13Dec2010-5.

Discussion

Pseudobaeospora deckeri is a small agaric that occurs throughout central

California but probably has a wider distribution northwards. It is distinguished

by its deep purple to violet pileus with a pale pinkish bloom, white spores,

strong green KOH reactions, pileipellis with upright elements, frequent clamp

connections, and lack of cheilocystidia.

Because ofits deeply colored fruitbodies and KOH reactions, Pseudobaeospora

deckeri shows affinity with P pyrifera and P jamonii in Bas infrageneric Pyrifera

and Frieslandica groups (Bas 2003). In Voto’s worldwide key, P. deckeri could

appear in either sect. Anistoderma Voto or sect. Pseudobaeospora, depending

on interpretation of the pileipellis (Voto 2009). Pseudobaeospora deckeri can

be distinguished from other members of the genus by its distinctly purple

coloration combined with its alkaline-virescent reactions, a pileipellis that is an

irregular cutis with abundant erect elements, abundant clamp connections, and

a lack of a distinctly cellular subpellis and cheilocystidia.

In age, the fruitbodies resemble those of P. stevensii, with is sympatric at

two of the known sites. Side-by-side comparison of such sympatric fruitbodies

showed that P. stevensii possesses a ‘yellower’-brown stipe with a coarser,

paler, and generally more scurfy apex and a paler, more strongly strigose base,

completely lacks violaceous tones, and shows an olive-green KOH reaction that

is easily visible against the paler brown flesh. This contrasts with P. deckeri, in

which the bluish-green color change in KOH is usually obscured by the purplish

464 ... Schwarz

flesh, and thus seeming blackish-vinaceous or blackish-olivaceous unless a thin

section is made to reveal a solidly bright green color. Micromorphologically,

P. stevensii has a cellular subpellis with no upright elements, cheilocystidia, and

slightly more elongate spores with a higher average Q value.

Although photographs of P. pyrifera Bas & L.G. Krieglst. from Europe

show a macroscopically similar entity, P pyrifera has short broadly clavate

cheilocystidia, a pileipellis of short, strongly inflated hyphae in chains, and a

different KOH reaction in the stipe trama. Another European taxon, P. jamonii

Bas et al., has a paler pileus, nearly free lamellae, and a pileipellis with scattered,

broadly clavate erect elements amongst strongly inflated chains of hyphae (Bas

et al. 2002). Also macroscopically similar is P dichroa Bas, which has intense

violet tones to the lamellae and stipe but a much paler pileus, only occasionally

(then only slightly) intervenose lamellae, and a reddish-purple KOH reaction

that eventually becomes yellowish-green. Pseudobaeospora euganea Voto

has a browner pileus, a pileipellis with only scattered erect elements and

a cellular subpellis, and significantly longer spores (Q = 1.14-1.45) (Voto

2009). Pseudobaeospora cyanea Arnolds et al. is a European species with a

more intensely bluish-purple pileus, paler lamellae, a stipe with much less

violet coloration, encrusted elements in the pileipellis, and significantly larger

(4.0-5.5 um x 3.3-4.0 um) spores (Arnolds et al. 2003).

At least four other members of this genus (some undescribed) occur

sympatrically, but all can be distinguished by their differently colored

fruitbodies. Pseudobaeospora aphana produces much paler fruitbodies and

smells distinctly fishy (Vellinga 2009); microscopically it is distinguished by the

presence of cheilocystidia and a cellular subpellis. Species not known to occur

in California include “Pseudobaeospora pillodii” sensu Redhead, P. murrillii E.

Horak, Tricholoma microsporum, and Agaricus fuscolilacinus Peck, all of which

lack KOH reactions and differ in other microscopic and macroscopic characters

(Desjardin 2004).

The ITS sequence obtained from the type collection was aligned against

that of Pseudobaeospora pyrifera; this comparison showed 93% identities

(E-value = 0.0), clearly demonstrating generic placement but distinct species—

level difference. Unfortunately, there are currently too few Pseudobaeospora

sequences in GenBank to perform broader phylogenetic analyses.

Acknowledgements

Very special thanks to Alexandra Grote for her sequencing work and to Drs. Else

Vellinga and Dennis Desjardin for their reviews, as well as encouragement, advice, and

relevant literature. Thanks also to Dr. Greg Gilbert for providing support and facilities,

Dimitar Bojantchev for his review of the manuscript and suggestion of the generic

disposition of this taxon, and Drs. Roy Halling and Greg Mueller for their thoughts

regarding generic disposition.

Pseudobaeospora deckeri sp. nov. (USA) ... 465

Literature cited

Adam¢ik S, Bas C. 2002. Pseudobaeospora mutabilis, a new species discovered in Slovakia.

Mycotaxon 84:272-275.

Arauzo S. 2011. Estudios en el género Pseudobaeospora. Revista Micolégica Errotari 8: 135-158.

Arnolds E, Tabarés M, Rocabruna A. 2003. Pseudobaeospora cyanea, a new agaric species from

Catalonia. Rev Catal Micol 25:65-70.

Bas C. 2002. A reconnaissance of the genus Pseudobaeospora in Europe I. Persoonia 18:115-122.

Bas C. 2003. A reconnaissance of the genus Pseudobaeospora in Europe II. Persoonia 18:163-199.

Bas, C, Lalli G, Lonati G. 2002. Pseudobaeospora jamonii, una nuova specie in Italia. Micol Veget

Medit 17:31-35.

Desjardin DE. 2004. A new species of Pseudobaeospora from California. Mycotaxon 90:69-76.

Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3rd Ed. Eyre Methuen, London.

202 Ps

Vellinga EC. 2009. Pseudobaeospora aphana, a new species from California. Mycologia 101:

243-246. http://dx.doi.org/10.3852/07-204

Voto P. 2009. Proposta di una sistematica infragenerica del Genere Pseudobaeospora, fondata su

basi morfologiche, e note su due specie non ancora descritte. Riv. Micol. 452: 291-311.

MY COTAXON

http://dx.doi.org/10.5248/119.467

Volume 119, pp. 467-476 January-March 2012

A new species of Volvariella from India

GUNASEKARAN SENTHILARASU’, RAHUL SHARMA? & SANJAY K. SINGH?

National Facility for Culture Collection of Fungi, MACS’ Agharkar Research Institute,

G. G. Agarkar road, Pune-411 004, India

CORRESPONDENCE TO: ’ senthilarasug@rediffmail.com, * rahulxsharma@yahoo.co.in,

° singhsksingh@rediffmail.com

ABSTRACT— Volvariella sathei sp. nov., collected from Agharkar Research Institute Campus,

Pune, is described, illustrated, and compared with morphologically closely related taxa of

Volvariella and Volvopluteus. Diagnostic characters of the new species are the large white

fruitbodies, relatively small basidiospores, and cylindric-clavate pleuro- and cheilocystidia.

Neighbour-Joining analysis based on ITS sequences confirmed that it is distinct from its

closest relative, V. nivea and other morphologically similar Volvariella species. A key to the

Indian species of Volvariella and Volvopluteus is provided.

Key worps—Agaricales, Basidiomycetes, macrofungi, taxonomy

Introduction

Volvariella Speg. (Agaricales) is well represented in India, and considerable

work has been carried out by Pathak et al. (1978) and Pradeep et al. (1998), who

have studied the genus and described and/or reported several species for the

country. Agaric checklists compiled by Manjula (1983), Natarajan et al. (2005),

and Kumaresan (unpublished list) cite 22 species of Volvariella including

Volvopluteus Vizzini et al. as reported from India. A white agaric from Pune

is here described as a new Volvariella species based on its morphological

and molecular characters. Its phylogenetic position has been determined by

ITS1+5.8S rDNA+ITS2 sequence analyses.

Materials & methods

Thin handmade sections were made from dried specimens, revived in 10% KOH, and

stained in 2% Phloxine. The microscopical characters were studied using an Olympus

U-CMAD3 microscope. Approximately 30 basidiospores from sections were measured;

the range (extreme values in parentheses) precedes mean spore measurements (in

parentheses). Camera lucida diagrams were made using Nikon E200 attached with

Nikon Y-IDT prism. Colour terminology follows Kornerup & Wanscher (1978). The

type specimen is deposited at Ajrekar Mycological Herbarium (AMH), MACS’ Agharkar

Research Institute, Pune, India.

468 ... Senthilarasu, Sharma & Singh

TABLE 1. Sequences used in the phylogenetic analyses.

SPECIES VOUCHER No. GENBANK No.

Volvariella bombycina AJ244 HM562212.1

V. caesiotincta MA54717 HM562211.1

V. dunensis SCM3513 JF415140.1

V. hypopithys TO AV137 HM246492.1

V. lepiotospora AJ155 HM562214.1

V. nivea GDGM25489 FJ749127.1

V. nullicystidiata SP393639 EU920671.1

V. pusilla AJ51(LOU) JF415137.1

V. pusilla TO AV139 HM246494.1

V. sathei AMH 9436 JN792550

V. strangulata TO AV141 HM246493.1

V. surrecta AJ55 HM562213.1

V. taylorii AJ54 HM562210.1

V. terrea LUG11010 JF415141.1

V. volvacea TO AV143 HM246500.1

V. volvacea ATCC MYA-4696* HQ999973.1

Volvariella sp. LOU18924 JF415139.1

Volvopluteus earlei (= V. acystidiata) TO HG1973 HM246499.1

Vp. earlei (= V. cookei) TO AV133 HM246496.1

Vp. earlei (= V. earlei) TO AV134 HM246497.1

Vp. earlei (= V. media) TO HG2001 HM246498.1

Vp. gloiocephalus (= V. gloiocephala) TO AV136 HM246495.1

Schizophyllum commune IFM 56967* AB566277.1

*Strain no.

DNA isolation methodology followed Aljanabi & Martinez (1997) with slight

modifications. ~300 mg of the holotype gills was washed thoroughly with 99% ethanol

and water. DNA was extracted using high salt extraction buffer (NaCl, Tris. HCl and

EDTA) and addition of one volume of phenol:chloroform:isoamyl alcohol (25:24:1)

and finally precipitated with isopropanol, washed with 70% ethanol, and suspended

in Tris-EDTA buffer. The protocol outlined by White et al. (1990) was followed for

polymerase chain reaction in Eppendorf Master cycler (Eppendorf, Germany) with

universal primers ITS5 and ITS4. Sequencing was performed using BigDye Terminator

Cycle Sequencing Kit (Applied Biosystem, UK) as per manufacturer's instructions.

The cleaned PCR product was run on ABI 3100 automated DNA sequencer. The raw

sequences obtained from ABI sequencer were manually corrected for inconsistencies

with the help of Chromas lite software. The DNA sequences obtained in Fasta format

were subjected to BLASTn (http://www.ncbi.nlm.nih.gov) analysis.

For phylogenetic analysis, additional related sequences obtained from NCBI DNA

sequence database (TABLE 1) were aligned by CLUSTAL W to generate a phylogenetic

tree using MEGA v.5.0 software (Neighbour-joining using Kimura-2 parameter model).

The outgroup comprised Schizophyllum commune and Volvopluteus spp. (recently

transferred from Volvariella).

Volvariella sathei sp. nov. (India) ... 469

PraTE 1. Volvariella sathei: Basidiomata under natural conditions in Agharkar Research Institute

campus. A. Surface view of basidiome. B. Gill view of basidiome. Photo Senthilarasu G.

470 ... Senthilarasu, Sharma & Singh

Taxonomy

Volvariella sathei Senthil., Rahul Sharma & S.K. Singh, sp. nov. PLaTEs 1-2

MycoBank MB 561949

Differs from Volvariella nivea in smaller subglobose to ovoid basidiospores and smaller

pleuro- and cheilocystidia.

Type: India, Maharashtra State, Pune, Agharkar Research Institute Campus (18°52'N

73°83’E), on ground (soil), grass field, 03.08.2009, coll. G. Senthilarasu (Holotype AMH

9436; GenBank JN792550).

Erymo.ocy: The epithet honors Dr. A.V. Sathe, a leading mycologist in western India.

PILEus 8-9 cm diam., convex; surface white to yellowish white (4A2), fibrillose,

shiny with fine radial striations in the margin; MARGIN thin, eroded; LAMELLAE

free, remote, white becoming sordid, < 10 mm wide, crowded with lamellulae

of different lengths, margin concolorous; StT1PE 80-105 x 9-13 mm, cylindric,

distinctly compressed, tapering towards apex, solid; surface white to cream,

glabrous; PILEAL CONTEXT 7 mm thick near stipe, white, unchanging on

bruising; VoLva free from the stipe, white (4A2), irregularly lobed, thick; ODOR

fungus like; TasTE not recorded.

BASIDIOSPORES (5.5)6-6.5 x (4.5)5-5.5, (5.97+0.26 x 5.05+0.26) um, Q= 1.1,

subglobose to ovoid, hyaline with a stramineous thickened wall, smooth;

BASIDIA 20-36 x 5.5-11 um, cylindric clavate to clavate, tetrasporic; STERIGMATA

short (< 2 um), slender; LAMELLA EDGE heteromorphous; CHEILOCYSTIDIA

scattered, 20-62 x 8-33 um, cylindric-clavate with broadly rounded apex,

thin-walled, easily collapsing; PLEUROCysTIDIA scattered, 27-72 x 15-34 um,

cylindric-clavate, mucronate to broadly rounded apex, thin-walled; LAMELLA

TRAMA convergent, consisting of thin walled, hyaline hyphae 5-18 um diam.;

SUBHYMENIUM little developed up to 20 um wide, pseudoparenchymatous;

PILEAL SURFACE a regular cutis consisting of radially repent to semi erect hyphae,

thin-walled, 5-30 um diam.; PILEAL CONTEXT consisting of tightly interwoven

hyphae, 5-30 um diam., thin-walled; VoLva consisting of gelatinous layer, up

to 50 um thick, hyphae thin-walled, up to 33 um diam., volval elements 20-47

x 15-24 um, cylindric-clavate; StrPITIPELLIS hyphae made of cylindric, septate

hyphae, 5-30 um diam., terminal elements 30-45 x 12-13 um, with rounded

apex, caulocystidia absent; CLAMP-CONNECTIONS absent in all hyphae.

ADDITIONAL SPECIMENS EXAMINED: INDIA, MAHARASHTRA STATE, Pune, Agharkar

Research Institute Campus (18°52'N 73°83’E), on ground (soil), grass field, 21.06.2009,

coll. G. Senthilarasu (AMH 9443).

Justo et al. (2010, 2011) reassessed species traditionally assigned to Pluteaceae

Kotl. & Pouzar based on molecular data. Volvariella has been placed outside

the pluteoid clade with a new genus, Volvopluteus, erected to accommodate

species with an ixocutis pileipellis and spores > 11 um long. Consequently,

Volvariella gloiocephala, V. earlei, and V. acystidiata have been transferred to

Volvariella sathei sp. nov. (India) ... 471

PiaTE 2. Volvariella sathei: a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Pleurocystidia.

e. Volval elements. f. Terminal elements of stipitipellis. Scale bar = 10 um.

472 ... Senthilarasu, Sharma & Singh

Volvopluteus. Volvariella acystidiata is considered now a form, as Volvopluteus

earlei f. acystidiatus (N.C. Pathak) Vizzini & Contu (Justo et al. 2011).

Of the 20 species described from India, two belong in Volvopluteus

(Vp. gloiocephalus (DC.) Vizzini et al. and Vp. earlei (Murrill) Vizzini et al.),

and the remaining species belong in Volvariella. Volvariella sathei is compared

here with morphologically similar taxa belonging to both genera.

Volvopluteus earlei, which also produces white basidiomata, differs in smaller

(3-5 cm diam.) size and longer (>11 um) spores (Justo & Castro 2010; Justo et al.

2010). Volvopluteus earlei f. acystidiatus, originally described by Pathak (1975)

and later by Vizzini & Contu (2010), also closely resembles V. sathei but clearly

differs by its smaller (20-30 mm diam.) yellowish tinged pileus with pinkish

margin, shorter more slender stipe (30-40 x 2-3 mm), larger spores (10.5-16.5

x 7.5-10.5 um) and basidia (45-70 x 10-13.5), and absence of hymenial cystidia

(Justo et al. 2010). Phylogenetically Volvopluteus species are clearly distant from

Volvariella species, including V. sathei. There are certain regions within ITS

including 5.8S that distinguishes Volvariella from Volvopluteus.

Volvariella sathei is similar to Volvariella pusilla (Pers.) Singer (Pegler 1977,

1983, as V. parvula) in the white basidiomata with fibrillose pileus and remote

lamellae. However, V. pusilla differs in stature with smaller pilei (1.5-3 cm

diam.) and stipes (10-40 x 2-4 mm) as well as producing smaller basidia (17-29

x 5-7.5 um), pleurocystidia (36-53 x 11-17 um), and cheilocystidia (25-40 x

9-16 um).

Volvariella nivea T.H. Li & Xiang L. Chen described from China (Li et al.

2009) also produces large, white basidiomata but clearly differs in slightly larger

ovoid to broadly ellipsoid spores and considerably larger cheilo- (50-150 x

20-46 um) and pleurocystidia (60-132 x 19-44 um).

Volvariella bombycina (Schaeff.) Singer, V. bombycina var. microspora

Dennis, V. hypopithys (Fr.) Shaffer, V. liliputiana (Henn.) G.C. Rath, and

V. delicatula (Massee) Manjula are also similar to V. sathei. However,

V. bombycina is lignicolous, and the spores are larger (7.3-9.4 um in V.

bombycina; 6-7.5 um in V. bombycina var. microspora). Volvariella hypopithys

has smaller basidiomes (pileus < 5 cm diam.) with squamulose pilei and hairy

stipes, while Volvariella liliputiana and V. delicatula are much smaller (pileus

5-30 mm diam.) than V. sathei.

A BLASTn search comparing the AMH 9436 530bp nucleotide sequences

(comprising ITS1 = 210bp, 5.88 = 151bp, ITS2 = 157bp) with the Genbank

DNA ITS sequence database showed maximum (84%) similarity with V. terrea

(JF415141.1 query coverage 88%) and 83% similarity with V. dunensis

(JF415140.1 query coverage 88%). Twenty-two ITS sequences representing

13 Volvariella spp., 2 Volvopluteus sp., and 1 Schizophyllum sp. obtained from

Genbank database were aligned using CLUSTAL W with manual adjustments.

Volvariella sathei sp. nov. (India) ... 473

69 Volvariella lepiotospora voucher AJ155

Volvariella nullicystidiata SP393639

Volvariella dunensis voucher SCM3513

Volvariella caesiotincta voucher MA54717

5é | 6 Volvariella volvacea voucher TO AV143 la

Volvariella taylorii voucher AJ54

Volvariella sathei sp. nov. AMH 9436

36 Volvariella nivea GDGM25489

Volvariella terrea voucher LUG11010

69 Volvariella strangulata voucher TO AV141

g? Volvariella surrecta voucher AJ55

sa} 91 Volariella pusilla voucher AJ51(LOU) Ib

Volvariella pusilla voucher TO AV139

38] — Volvariella hypopithys voucher TO AV137

#3L Volvariella sp. LOU18924

Volvariella bombycina voucher AJ244

100 Volvariella volvacea strain ATCC MYA-4696

Schizophyllum commune strain: IFM 56967

Volvopluteus gloiocephalus(=Volvariella gloiocephala) voucher TO AV136

ti 100 Volvolputeus earlei (=Volvariella cookei) voucher TO AV133

gg | Volvopluteus earlei (=Volvariella earlei) voucher TO AV134

g7|| Volvopluteus earlei(=Volvariella media) voucher TO HG2001

65L_ Volvopluteus earlei (=Volvariella acystidiata) voucher TO HG1973

PLATE 3. Neighbour-joining tree showing phylogenetic position of Volvariella sathei, Volvariella,

and Volvopluteus species. Bootstrap values > 50% are indicated.

The 5.8S region was almost perfectly aligned, but numerous gaps (indels) and

misalignments among Volvariella species were observed in ITS1 and ITS2

regions. All analysed Volvariella species were divided into two main clusters

(Land II, PLare 3), with V. sathei, V. nivea, and V. taylorii together in subclade Ia.

Although V. sathei and V. nivea, both characterized by pure white basidiocarps,

are placed together with a strong (96%) bootstrap support, the branch lengths

show considerable sequence divergence between them. Our analysis places the

two sequences of Volvariella volvacea (HM246500.1, TO AV143; HQ999973.1,

ATCC MYA-4696) far apart (in completely two different clusters), suggesting

ambiguity in deposited sequences.

As Li et al. (2009) has indicated for V. nivea, we noted a high degree of

variability (including considerable misalignment and several indels) throughout

the ITS 1 and ITS 2 regions among Volvariella species, but the 5.8S region

remained conserved with relatively few changes.

474 ... Senthilarasu, Sharma & Singh

Key to species of Volvopluteus (Vp.) and Volvariella (V.) in India:

Two newly described species —Volvariella indica Pathak et al. (Manjula 1983) and

V. minuta Kumar et al. (Kumar et al. 2009-10)— and a new record of V. congolensis N.C.

Pathak (Kumar et al. 2009-10) from India are poorly known and have been excluded

from the key.

LeSpores > liljpuint lone: pileipellis-is ai txo Cutis 6 ot panes esas engage Package oka lage nae 2

L.spores <10:5; tim: long, pileipellis miotairOcutisn: a2 Sivertson i hens cesarean 3

2. Pileus 5-15 cm diam.; spores 11-18 x 6.5-9.5 um (ivory white to

light brownish grey, often darker on the disc) ................ Vp. gloiocephalus

2. Pileus 3-5 cm diam.; spores 11-15.5 um long ................. 000. e ee Vp. earlei

3. Lignicolous, on dead wood, pileus pale yellow then whitish ..................... 4

De SLOWING, ONE OLE TESUDSEGALA o: 2: «nates Mar art teteawa ia Bontecls dave a siawa tae titva Mena buaciNe Green Ne arey 7

4, Basidiome robust, > 5 cm diam; volva thick and fleshy..................0..0000. 5

4. Basidiome-siiall,-<Scmidiam:; volva-siiall hit t.. .12.29.28, ae 0t. ate. eee tet anak 6

S oporestrequentiv: O—1O0'S, tiny On ges Vest ne eles agen caer atau eh ahi V. diplasia

5: Spores:hot 9.5 um long -volva White. 2 eee s cee oie net eet wee oe V. bombycina

6. Volva internally rugoso-tomentose; lamellae pale flesh pink;

SPOres Oa ACS At, Ao eS Ph) Aes Pa en ee Oe ade Siew ee See V. glandiformis

6. Volva smooth; lamellae with brownish tints; spores 6.3-8 x 4-5 um .... V. thwaitesii

7. Basidiome growing on other mushrooms.................0 0 cee eee ee V. surrecta

7. Basidiome growing on soil or other organic substrates................000. 00 eee 8

5: Basidiome pure -wiaiteto; pinkish.white «1.23 Fant let Lah OE I ath be 9

SsBasidiome ash ereen-to: browns) Siac le ce Cees eee ee Token he 13

9. Pileus and stipe distinctly hairy (pileus < 5 cm diam., white;

SPOKES O- 7X ASS I) ey ass nee a dipn ¢ Fd) ber Edi poor gd Ainber Fai naer gainers V. hypopithys

9. Basidiome not hairy, glabrous, subvelutinate to silky fibrillose .................. 10

10. Pileus 8-9 cm diam. white, fibrillose; stipe 80-105 x 9-13 mm .......... V. sathei

10, Pileus.small'= 3:erm-diam-stipe short.and sletider ss:5. clone cadens odlein edn se ee 11

11. Pileus < 1 cm diam, subvelutinate; spores 4.5 x 4um............... V. liliputiana

11. Pileus < 3 cm diam., glabrous to silky fibrillose.......... 0.0... cece cece eee eae 12

12. Spores 5-6 x 3.2-4 um, pileus at first pinkish white ................. V. delicatula

12. Spores 4.5-7.5 x 4-6.5 um, pileus pure white to pale cream (silky smooth

to silky fibrillose, volva white, appressed) ........... 00... cece eee eee V. pusilla

13. Pileus ash-green to grayish brown, volva white .................. V. woodrowiana

PS" Biles -cifferent.Shaves.of DLO Wil. lech Son 1a soit rin eh lt dads hadnt ya tote i 14

14.Volva, fleshy, rigid, pale to dark brown to blackish brown above,

eracking into-areolatec SqUAaMUles. seks acy ches sts fovea sph febeg eb eben dh 5, Sgkdivrees V. terastia

14 Volya smooth onot squamoses... aif... 2a 5 8 Pape oh Rape Peps eB Pas enn hee o ens 15

15. Basidiome large and fleshy; pileus robust < 12 cm diam., tawny brown to

grayish brown; stipe 7-20 mm diam.; volva brown..................0. 00 eee 16

15. Basidioma small to medium < 6.5 cm diam.; stipe 2-6 mm diam.;

VOlVal STEY,: WHE OM DTOW IN. to Fics ferns Bags le Paes ogg Pape le Pies tte lee fo aeace 18

Volvariella sathei sp. nov. (India) ... 475

16. Spores frequently > 9 um long (6-10.4 x 4.5-7 um)... eee eee V. volvacea

1G; Spores sno ti, SHO Me. ew ues hew, Aer Oy Ns tp ile spa Made Pl neha Fede Woedek eabledbu, eal 17

17. Pileus light seal brown at the disc, paling to grayish vinaceous at the margin

with fine brown radial striae; spores 4.5-8.6 x 3-5.3 um............ V. cubensis

17. Pileus bright chestnut brown, spores 5-6.5 x 3.5-4 um... 0... eee V. castanea

L8eRileds oravish; Valve Oravichts!. (mst Oh SEN eee oye Fe ete Fawn de Sheds Oe iby

18, Pileus'browaishs volyaibrowir OF White si5% sein ksnp trace te eee area sane ev ate ge = 20

19. Pileus evenly pale to dark grey, slightly darker at the disc;

MaroMelOtStHiAtes eh. 327) Sy AE ORE Se TM Oe et Ree Pe Sees olan V. taylorii

19. Pileus evenly grayish with blackish disc; margin or the entire cap

ra Fog DIRE A (ALO 1 79 Bed Spt yt SGC, SPST Gh BIR Eagan Ang oN OLE AERA Tce V. nigrodisca

20. Pileus vinaceous brown at the disc, paler towards the margin; stipe very long,

4-10 cm x 3-6 mm; volva fuliginous brown ................ V. pseudovolvacea

20. Pileus ash gray to grayish brown with blackish radial fibrils; stipe short,

4-4.5 cm x 2-4 mm, volva white ........ 2... eee eee V. apalotricha

Acknowledgments

We thank Dr. Alfredo Justo (Biology Department, Clark University, Worcester,

MA, USA) and Dr. Zai-Wei Ge (Kunming Institute of Botany, Chinese Academy of

Sciences, Kunming, Yunnan, P.R. China) for critically reviewing the manuscript and

giving appropriate modifications in the presubmission of the manuscript. We express

our sincere thanks to Dr. Shaun Pennycook and Dr. Lorelei Norvell for their valuable

suggestions after manuscript submission. GS personally thanks Dr. Alfredo Justo and

Dr. Vadivelu Kumaresan for providing literature on Volvariella species. Sincere thanks

to the Director, Agharkar Research Institute, for providing all laboratory facilities. We

would like to thank the Department of Science and Technology (DST), Government

of India, New Delhi, for providing financial support under the IRPHA Programme for

setting up state-of-the-art National Facility for Culture Collection of Fungi (No. SP/SO/

PS-55/2005) at Agharkar Research Institute, Pune, India.

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Manjula B. 1983. A revised list of the agaricoid and boletoid basidiomycetes from India and Nepal.

Proceedings of Indian Academy of Sciences (Plant Science) 92: 81-213.

Natarajan K, Kumaresan V, Narayanan K. 2005. A checklist of Indian agarics and boletes

(1984-2002). Kavaka 33: 61-128.

Pathak NC. 1975. New species of Volvariella from Central Africa. Bull. Jard. Bot. Natl. Belg.

45: 195-196. http://dx.doi.org/10.2307/3667598

Pathak NC, Ghosh R, Singh MS. 1978. The genus Volvariella Speg. in India. 295-303, in: CK Atal

et al. (eds). Indian Mushroom Science-1. Indo-American Literature House.

Pegler DN. 1977. A preliminary agaric flora of East Africa. Kew Bull. Addit. Ser. 9: 615 p.

Pegler DN. 1983. Agaric flora of the Lesser Antilles. Kew Bull. Addit. Ser. 9. 668 p.

Pradeep CK, Vrinda KB, Mathew S, Abraham TK. 1998. The genus Volvariella in Kerala state, India.

Mushroom Research 7(2): 53-62.

Vizzini A, Contu M. 2010. Volvariella acystidiata (Agaricomycetes, Pluteaceae), an African

species new to Europe, with two new combinations in Volvariella. Mycotaxon 112: 25-29.

http://dx.doi.org/10.5248/112.24

White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal

RNA genes for phylogenetics. 315-322, in: MA Innis et al. (eds) PCR protocols: a guide to

methods and applications. Academic Press, San Diego California.

MY COTAXON

http://dx.doi.org/10.5248/119.477

Volume 119, pp. 477-482 January-March 2012

A new species of Pleurophragmium from India

Maria A. D'Souza” & D.J. BHAT ”

"Department of Botany, Dhempe College of Arts & Science, Miramar, Goa-403 002, India

*Department of Botany, Goa University, Goa-403 206, India

*CORRESPONDENCE TO: majorinal @rediffmail.com

ABSTRACT — A new anamorphic species, Pleurophragmium indicum (Ascomycota), is

described from India and illustrated. This litter-inhabiting fungus was collected from dead

Dendrocalamus strictus leaves and is characterized by ellipsoidal to obovoid, straight to curved,

transversely 3-septate, brown conidia. A key is provided to all accepted Pleurophragmium

species.

Key worps — biodiversity, microfungi, taxonomy

Introduction

During our studies on taxonomy and diversity of microfungi associated

with flowering plants of Western Ghat forests in southern India, a hitherto

unknown anamorphic ascomycete with affinities to Pleurophragmium

Costantin was recovered from moist-chamber incubated leaf litter of the

bamboo, Dendrocalamus strictus.

Materials & methods

Dried and freshly fallen leaves of Dendrocalamus strictus were thoroughly washed in

distilled water and incubated in a sterile moist chamber for one to several days at room

temp (22-25°C). The leaves were examined at 2-day intervals under a stereoscope fitted

with incidental light. Grey-brown colonies with erect to slightly flexuous conidiophores

with a crown of conidia appeared on the surface of the leaves after 5-7 days of incubation.

The fungus was mounted on a slide with lactophenol and examined under a bright field

light microscope.

Taxonomy

Pleurophragmium indicum M.A. D’Souza & Bhat, sp. nov. Fics. 1-4

MycoBank MB563115

Differs from P. bitunicatum in conidia that lack a distinct outer covering and from P

parvisporum in its larger conidia and varied colouration.

478 ... D'Souza & Bhat

20um

Fics. 1-3. Pleurophragmium indicum.

1. Habit. 2. Magnified view of conidiophores with conidia. 3. Conidia.

Type: India, Molem Wildlife Sanctuary, Goa, on fallen dead and decaying leaves of

Dendrocalamus strictus (Roxb.) Nees (Poaceae), 11 March 1999, leg. Maria A. D’Souza,

(Holotype, GUBH 367).

EryMo_oey: from the country of the type locality.

Co.tontgseffuse, dark brown to black, velvety on thenatural substrate. MyCELIUM

partly immersed and partly superficial. COoNIDIOPHORES mononematous,

scattered, erect, straight to slightly flexuous, swollen and rhizome-like at the

Pleurophragmium indicum sp. nov. (India) ... 479

Fic. 4. Pleurophragmium indicum. Conidiophores and conidia, 600x.

base, smooth, 4—6-septate, dark brown below, pale brown in the middle and

above, thick-walled, 100-160 um long, 7-22 um wide at base, 4.5-6.5 um wide

in the middle and near apex. CONIDIOGENOUS CELLS polyblastic, integrated,

terminal, cylindrical, sympodial, minutely denticulate, noncicatrized, smooth,

pale brown to subhyaline, 15-23 x 4.5-9.5 um. Conrp1A solitary, simple, dry,

ellipsoidal to obovoid, rounded at the apex, narrower at the base, straight to

slightly curved, smooth, transversely 3-septate, middle cells dark brown, end

cells pale brown, 20-30 x 4.5-11 um, 3-5 um wide at the ends.

480 ... D'Souza & Bhat

Discussion

The genus Pleurophragmium, with P. bicolor Costantin [= P. simplex (Berk. &

Broome) S. Hughes] as type, includes 37 currently accepted taxa, with 27 other

names placed in synonymy (Hughes 1958, Ellis 1971, 1976, Matsushima 1975,

Kirk et al., 2008). Although the conidia are similar in shape and pigmentation

to Curvularia Boedijn (Ellis 1971), P indicum is differentiated by its blastic

mode of conidial development.

Of those validly described species, P. bitunicatum Matsush., which resembles

P. indicum in pigmentation and conidial size, differs by having conidia with

distinct outer covering. Pleurophragmium parvisporum (Preuss) Hol.-Jech.

TABLE 1. Comparison of accepted species of Pleurophragmium

SUBSTRATE TYPE

SPECIES

LOCALITY

P. acutum (Grove) M.B. Ellis 1976 Dead stems of Urtica dioica England

P. angamosense Matsush. 1995 Decayed petiole of palm Peru

P. aquaticum R.F. Castafieda et al., 2007 Decaying wood submerged in a stream Mexico

P. arecae Matsush. 1987 Decaying leaves of Areca catechu Taiwan

P. bicolor Matsush. 1975, nom. illegit. Decayed leaves of Quercus, Podocarpus Japan

P. bitunicatum Matsush. 1975 Decaying leaves Japan

P. cylindrosporum Matsush. 1975 Decaying leaves Japan

P. harunganae Hansf.1946 Peele ones iam Kaas oF Uganda

Harungana madagascariensis

P. hippotrichoides Dead leaves, stems of Cortaderia selloana, Europe,

(Corda) M.B. Ellis 1976 Umbilicus rupestris, Pandanus sylvestris, Channel

and Phormium tenax Islands

P. indicum Dead and decaying leaves of ;

: India

Dendrocalamus strictus

P. malaysianum Matsush. 1996 Decaying leaves Malaysia

P. miniumbonatum (R.F. Castafieda et al.) Fallen decaying leaves Venezuela

R.E Castafieda 2007

P. naviculiforme Matsush. 1975 Decaying leaves Japan

P. obcampanuloides Matsush. 1995 Decaying stems Japan

P. parvisporum (Preuss) Hol.-Jech., 1972 Rotten leaves of Musa sapientum Taiwan

P. peruamazonicum Matsush.1993 Decaying leaves of palm Peru

P. peruamazonicum var. inflatum .

Matsush. 1993 Decaying and rotten leaves of palm Peru

P. simplex (Berk.& Broome) Partly decorticated dead stems of Barns

S. Hughes 1958 Urtica, Brassica, Polygonum P

P. subfusiforme Matsush. 1975 Decaying stems Japan

P. triseptatum Matsush. 1975 Decaying stems Japan

P. tritici M. B. Ellis 1976 Dead wheat culms Ireland

P. varieseptatum Matsush. 1975 Decaying leaves of Phyllostachys edulis Japan

P. verruculosum D.P. Tiwari 1970 Rhizosphere soil of Piper betle India

Pleurophragmium indicum sp. nov. (India) ... 481

produces similarly shaped conidia but differs in the uniformly pale ochraceous

conidia that also are markedly smaller (14.5-25 x 4.5-6.5 um).

Twenty-three accepted taxa of Pleurophragmium are compared (TABLE 1)

and keyed out below.

Key to accepted taxa of Pleurophragmium

Ie@onidia With 2tunicaees i... 24:nne dd noe a4 nad 4 nace ord 4 nee 84 nore P. bitunicatum

Has COM fa WAU a: CATIA ez ib sct-n Finck Shes chee ods cdea Fibs phon Fds sehen db bspctew aut scbow ayuipctoa 2

2. Conidia smooth or indistinctly granular, 12-18 x 5-7um........... P. harunganae

2a. Conidia verrucose, finely verruculose or minutely echinulate ................. 3

AD COUN CIA SIN GOW bit s.. ha pt lee da giees eagblere Peeler 3 egl eeagt oko wah ech ae fects foe arn fect pea 4

3. Conidia verrucose, subclavate to clavate, 10-16.5 x 3.3-5um ..... P. verruculosum

3a. Conidia finely verruculose, ellipsoidal, 9-14 x 6-8 um hyaline .. P. hippotrichoides

3b. Conidia minutely echinulate, ellipsoidal to fusiform, 6-11 x 3-4uum ...... P. tritici

An GOTTA MV ANTIO ricvarlstavacces teavncans teavRcgha Le eare Lede ne teen LedineseLeAgUA Toe aaaeN cot RRNA 5

4a, ‘Conidia-stb hyaline todemattaceOus, Bs, ic tie ton fsa tan cesatign saaetlgn cs atop st uerate ete 6

5- Conidianaviculate: 6=b2 6253 [inion sg atans ealettel oS ol BR ah P. naviculiforme

5a. Conidia fusiform to ellipsoidal, 18-32 x 6.5-10 um...............002- P. acutum

5b. Conidia cylindro-clavate, (20 -) 40-75 x 4-5 um ............... P. malaysianum

GxGonidia sublyalime ee oon oe een ae beeen tbe lee hate ee, BN Rene, OSE ree Alea, ds Mate epee 7

Ga sGonidiasdematiaceOuss So. tu fs sci s + tints ou wht sie is ie SNe a Sia A oe, atk os abd oe 8

7. Conidia fusiform, curved, 3-7-septate, 21-38 x 5.8-9.2um........ P. subfusiforme

7a. Conidia cylindrical, (1—)3(- 6)-septate, 14-25 x 4-5.7 uum......... P. triseptatum

7b. Conidia fusiform to obclavate, (2—)3(-4)-septate, 12-24 x 3-4.5um ..... P. arecae

HE GOS UPTGRE MRO ANT UCAN dF) Ae dnt im: Sel alae an OMe do Ras De de ders ke dir Reh ar des come Ae Ce ete 9

$a. Gonidia-oFothershapes,niot eylindtical’ ss. crust sta sietdotd Sisteote Sew td ete Sued Saale ds 12

OsGohtdia usually: 32septate: 22). 25536). 25 eke’ heat 8 mente Reng hong tS cee ante 10

9a-Conidia2= or 1-4-septate sa. «sash feos Vad Yao + tals took dent os Vig

10. Conidia pale to moderately brown, (15-) 20-28 x 2.8-4 um

PP Rahs Ee eed SRA ED aL ee PERSO EO A, P. bicolor Matsush., nom. illegit.

10a. Conidia central cells brown, terminal cells pale brown, 25-40 x 5-6 um

FEE he Pe Le eat Le Reet See Se eR Ae ee ee P. cylindrosporum

11. Conidia varicolored with a pale brown central cell and subhyaline terminal cells,

2ESe plate, IS—BOVS-AR G6 IIE esa \isers ett aes chats acea lye: scenutise cceeatl P. peruamazonicum

lla. Conidia pale olive, 1-4-septate, 8-22 x 3-5 um ............... P. varieseptatum

PD Meonidiall 2 sseplgte ecko. Peele: Dorie. Doe! ok tag! bak age bata ot Pet ote nda hohe od bate 13

(2a AS OMIdia DOL INOS. SCP tAle he. aN enticed Um add eM odd AM olde ad co UR ae elt Rly ae 14

13. Conidia obpyriform to turbinate, 1-2-septate, 11.5-15.5 x 5-6.5um.............

wana WP ok Se elas eh et ih ae ol ae YE Yaa a ee A P. obcampanuloides

13a. Conidia ovoid, (1-) 2-septate, 17-28 x 8-1.5 um

Ppt Eikee lle aur ch. au decnor Rar ote on Rot P. peruamazonicum var. inflatum

482 ... D'Souza & Bhat

14. Conidia 5-septate, fusiform, slightly curved or straight,

2A SAO D 7) (NTN EB atid tans elbeabery wMlbeakys tiratece tlinadory selec bord P. angamosense

14a. Conidia 3-septate, ellipsoidal to obovoid or fusiform to clavate .............. 15

14b. Conidia 3-4-septate, obovoid, oblong, ellipsoidal to subclavate .............. 16

15. Conidia ellipsoidal to obovoid, straight to curved, 20-30 x 3-11 um... P. indicum

15a. Conidia fusiform to clavate, sometimes navicular, 25-30 x 6.0-6.5 um

POOP LO Ae PTO Meh Ras gee ag er may etc Piper iy Pee cee P. aquaticum

16. Conidia ellipsoidal to subclavate, 10-21 x 3.5-6 um ..............00-. P. simplex

16a. Conidia obovoid, pyriform to broad clavate ........... 0.0 cee eee ee eee 17

17 -Conidia obovoid, 10=252% 46.5: pin: ho ance igs seacsaee ong sow anole ee anecae P. parvisporum

17a. Conidia obovoid, pyriform to broad clavate, umbonate at the apex,

MG = TOE AI Ba Seiad con dr pneieg dyhy wedoa hehe fokod ach beboa dh sckeadih abe P. miniumbonatum

Acknowledgments

This work was supported by a research grant to D.J. Bhat by the Department of

Science and Technology, Government of India, New Delhi. MD thanks the Department

of Science and Technology, New Delhi, for a research fellowship. The authors thank

Prof. Kevin Hyde (Thailand) and Prof. Bryce Kendrick (Canada) for their comments

and suggestions and Dr. Rafael F. Castafieda-Ruiz (Cuba) and Dr. Eric McKenzie (New

Zealand), for reviewing the manuscript.

Literature cited

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey,

England.

Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew,

Surrey, England.

Hansford CG. 1946. The foliicolous Ascomycetes, their parasites and associated fungi Mycological

Papers 15: 240 p.

Heredia Abarca G, Castafieda Ruiz RF, Arias RM, Saikawa M, Stadler M. 2007. Anamorphic fungi

from submerged plant material: Acumispora verruculosa sp. nov., Pleurophragmium aquaticum

sp. nov. and Pleurophragmium miniumbonatum comb. nov. Mycotaxon 101: 89-97.

Holubova-Jechova V. 1972. A note on Pleurophragmium simplex (hyphomycetes). Ceska Mykologie

26 223-25:

Hughes SJ. 1958. Revisiones Hyphomycetum aliquot cum appendice de nominibus rejiciendis.

Canadian Journal of Botany 36: 727-836. http://dx.doi.org/10.1139/b58-067

Kirk, PM, Cannon PF, Minter DW, Stalpers JA. 2008. Ainsworth & Bisby’s dictionary of the fungi.

10" edition. CAB International Wallingford, U.K.

Matsushima T. 1975. Icones microfungorum a Matsushima lectorum. Matsushima, Kobe, Japan.

Matsushima T. 1987. Matsushima Mycological Memoirs 5: 1-100.

Matsushima T. 1993. Matsushima Mycological Memoirs 7: 1-141.

Matsushima T. 1995. Matsushima Mycological Memoirs 8: 1-44.

Matsushima T. 1996. Matsushima Mycological Memoirs 9: 1-30.

Tiwari DP. 1970 [“1969”]. A new species of Pleurophragmium from rhizosphere of Piper betle.

Indian Phytopathology 22: 513-515.

MY COTAXON

http://dx.doi.org/10.5248/119.483

Volume 119, pp. 483-485 January-March 2012

New combinations in Lactifluus. 2. L. subg. Gerardii

D. STUBBE”’, X.-H. WANG? & A. VERBEKEN**

'Ghent University, Department of Biology, Research Group Mycology,

K.L. Ledeganckstraat 35, B-9000 Gent, Belgium

? Section Mycology and Aerobiology, Scientific Institute of Public Health,

Juliette Wytsmanstraat 14, 1050 Brussels, Belgium

> Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany,

Chinese Academy of Sciences, Kunming 650204, P. R. China

* CORRESPONDENCE TO: mieke.verbeken@ugent.be

ABsTRACT — In this second of a series of three papers, new combinations in the genus

Lactifluus are proposed. This paper treats Lactarius subg. Gerardii (proposed here as new

combination in Lactifluus). In this subgenus 17 combinations at species level are proposed.

Key worps — milkcaps, nomenclature, Lactarius, Russulaceae

Introduction

This is the second treatment of species now classified in Lactifluus. Lactifluus

subgenera Edules, Lactariopsis, and Russulopsis were discussed in the first article

(Verbeken et al. 2012).

Taxonomy

Lactifluus subg. Gerardii

This recently established subgenus in Lactarius represents a group of milk-

caps that is well defined both molecularly and morphologically. Species are

agaricoid with brown pileus and stipe, contrasting pale lamellae, and a white

spore print. Three deviant species —all three remarkably small, pleurotoid,

and white (Lactarius uyedae, L. conchatulus, L. genevievae)— also belong to

this subgenus. Their alliance with Lactifluus subg. Gerardii is confirmed by a

multi-gene phylogeny but is also substantiated by their microscopic features.

The ornamentation of the basidiospores in this subgenus is always reticulate

and composed of interconnecting ridges, usually between 0.5 and 1.5 um

high. The pileipellis is typically a palisade with a strongly developed cellular

subpellis. True pleurocystidia are rare. As of this writing, L. subg. Gerardii

484 ... Stubbe, Wang & Verbeken

contains 18 described species, all from North America, Asia, and the Australian

region (Stubbe et al. 2012), and the group is absent from Africa and Europe.

The type species, Lactarius gerardii, has already been recombined in Lactifluus.

Stubbe et al. (2010) has demonstrated the existence of several cryptic species

and species complexes in this subgenus, and the total number of species will

likely increase after more data are gathered on several North-American milk-

caps like Lactarius atro-olivaceus, L. pseudogerardii, L. louisii, L. subtomentosus,

L. subisabellinus var. murrillianus, and L. xanthydrorheus.

Lactifluus subg. Gerardii (A.H. Sm. & Hesler) Stubbe, comb. nov.

MycoBank MB 564016

= Lactarius ser. Gerardii A.H. Sm. & Hesler, Brittonia 14: 378. 1962

= Lactarius subg. Gerardii (A.H. Sm. & Hesler) Stubbe, Fungal Biology 114: 280. 2010.

Tye: Lactarius gerardii Peck

Lactifluus atrovelutinus (J.Z. Ying) X.H. Wang, comb. nov.

MycoBank MB 564017

= Lactarius atrovelutinus J.Z. Ying, Acta Mycol. Sin. 10: 191. 1991.

Lactifluus bicolor (Massee) Verbeken, comb. nov.

MycoBank MB 564018

= Lactarius bicolor Massee, Kew. Bull. Misc. Inf. 1914: 73. 1914.

Lactifluus conchatulus (Stubbe & H.T. Le) Stubbe, comb. nov.

MycoBank MB 564019

= Lactarius conchatulus Stubbe & H.T. Le, Fungal Diversity 52: 162. 2012.

Lactifluus coniculus (Stubbe & Verbeken) Verbeken, comb. nov.

MycoBank MB 564021

= Lactarius coniculus Stubbe & Verbeken, Fungal Diversity 52: 152. 2012.

Lactifluus genevievae (Stubbe & Verbeken) Stubbe, comb. nov.

MycoBank MB 564022

= Lactarius genevievae Stubbe & Verbeken, Fungal Diversity 52: 163. 2012.

Lactifluus gerardii (Peck) Kuntze, Revis. Gen. Pl. 2: 856. 1891.

= Lactarius gerardii Peck, Bull. Buffalo Soc. Nat. Sci. 1: 57. 1873, as “geradii”.

Lactifluus hora (Stubbe & Verbeken) Stubbe, comb. nov.

MycoBank MB 564023

= Lactarius hora Stubbe & Verbeken, Fungal Diversity 52: 160. 2012.

Lactifluus leae (Stubbe & Verbeken) Verbeken, comb. nov.

MycoBank MB 564024

= Lactarius leae Stubbe & Verbeken, Fungal Diversity 52: 154. 2012.

Lactifluus leonardii (Stubbe & Verbeken) Stubbe, comb. nov.

MycoBank MB 564025

= Lactarius leonardii Stubbe & Verbeken, Fungal Diversity 52: 156. 2012.

Lactifluus subg. Gerardii combs. nov. ...

Lactifluus limbatus (Stubbe & Verbeken) Stubbe, comb. nov.

MycoBank MB 564026

= Lactarius limbatus Stubbe & Verbeken, Fungal Diversity 52: 147. 2012.

Lactifluus ochrogalactus (Hashiya) X.H. Wang, comb. nov.

MycoBank MB 564027

= Lactarius ochrogalactus Hashiya, Mycoscience 47: 232. 2006.

Lactifluus petersenii (Hesler & A.H. Sm.) Stubbe, comb. nov.

MycoBank MB 564029

= Lactarius petersenii Hesler & A.H. Sm., N. Am. Species Lactarius: 106. 1979.

Lactifluus reticulatovenosus (Verbeken & E. Horak) Verbeken, comb. nov.

MycoBank MB 564030

= Lactarius reticulatovenosus Verbeken & E. Horak, Sydowia 53: 285. 2001.

Lactifluus sepiaceus (McNabb) Stubbe, comb. nov.

MycoBank MB 564031

= Lactarius sepiaceus McNabb, New Zealand J. Bot. 9: 50. 1971.

Lactifluus subgerardii (Hesler & A.H. Sm.) Stubbe, comb. nov.

MycoBank MB 564032

= Lactarius subgerardii Hesler & A.H. Sm., N. Am. Species Lactarius: 107. 1979.

Lactifluus uyedae (Singer) Verbeken, comb. nov.

MycoBank MB 564033

= Lactarius uyedae Singer, Nov. Hedw. 40: 436. 1985.

= Pleurogala uyedae (Singer) Redhead & Norvell, Mycotaxon 48: 377. 1993.

Lactifluus venosus (Verbeken & E. Horak) Verbeken, comb. nov.

MycoBAnk MB 564034

= Lactarius venosus Verbeken & E. Horak, Austr. Syst. Bot. 13: 692. 2000.

Lactifluus wirrabara (Grgur.) Stubbe, comb. nov.

MycoBank MB 564035

= Lactarius wirrabara Grgur., Larger Fungi S. Australia: 58. 1997.

Acknowledgments

485

The authors acknowledge Scott Redhead and Shaun Pennycook for valuable

comments and for reviewing the manuscript.

Literature cited

Stubbe D, Nuytinck J, Verbeken A. 2010. Critical assessment of the Lactarius gerardii species complex

(Russulales). Fungal Biology 114: 271-283. http://dx.doi.org/10.1016/j.funbio.2010.01.008

Stubbe D., Le H.T., Wang X.-H., Nuytinck J., Van de Putte K., Verbeken A. 2012. The Australasian

species of Lactarius subgenus Gerardii (Russulales). Fungal Diversity 52: 141-167.

http://dx.doi.org/10.1007/s13225-011-0111-3

Verbeken A, Nuytinck J, Buyck B. 2012 [“2011”]. New combinations in Lactifluus. 1. L. subgenera

Edules, Lactariopsis, and Russulopsis. Mycotaxon 118: 455-458.

http://dx.doi.org/10.5248/118.447

MYCOTAXON

http://dx.doi.org/10.5248/119.487

Volume 119, pp. 487-492 January-March 2012

BOOK REVIEWS AND NOTICES

ELSE C. VELLINGA, Book Review Editor*

861 Keeler Avenue, Berkeley CA 94708 U.S.A.

CORRESPONDENCE TO: bookreviews@mycotaxon.com

INTRODUCTION

The books reviewed here are as diverse as the fungi themselves, with

molecular systematics of Penicillium and a field guide to the species of one

genus in one country for conservation purposes as extremes of one continuum.

Several publications stand out because of the high quality photos.

Book announcements include a volume on smut fungi in the series FLORA

FUNGORUM SINICORUM, a Spanish guide, and a Festschrift for the lichenologist

Thomas H. Nash III.

ASCOMYCETES

Phylogenetic and taxonomic studies on the genera Penicillium and

Talaromyces. Edited by R.A. Samson and J. Houbraken. 2011. SrupiEs In

Myco.Loey no. 70. CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85176,

3508 AD Utrecht, The Netherlands. <info@cbs.knaw.nl>. Pp. 183, illustr.

ISBN 978-90-70351-87-8. Price: 60 € (paper copy), download free.

This work is significant in two respects. First, with a publication date of 15

November 2011, it is the first major work on the taxonomy of Trichocomaceae

to have appeared since the separate nomenclatural status of anamorph-typified

and teleomorph-typified names of pleomorphic fungi ended on 30 July 2011.

Second, it is a significant contribution to the phylogenetic systematics of

Penicillium and Talaromyces in its own right, presenting the results of four fresh

studies.

Of particular importance is Houbraken & Samson's overview and revised

phylogenetic system for not only penicillioid but also aspergilloid fungi. This

‘Books for consideration for coverage in this column should be mailed to the Book Review Editor

at the address above. All unsigned entries are by the Book Review Editor.

488 ... Vellinga, BOOK REVIEW EDITOR

may come as a shock to some — with even two new families being recognized.

Molecular phylogenetic studies using several gene show that three families

are warranted: (1) Aspergillaceae for Aspergillus s. str. (syn. Eurotium, Fenellia,

Neocarpentales, Neosartorya, other teleomorphs, and _ Stilbothamnium),

Hamigera (incl. Merimbla), Leiothecium, Monascus (incl. Basipetospora),

Penicilliopsis, Penicillium s. str. (syn. Chromocleista, Citromyces, Eladia,

Eupenicillium, Hemicarpentales, Thysanophora, and Toluromyces), Phialomyces,

Phialosimplex, Polypaecilum, Sclerocleista, Warcupiella (incl. Raperia), and

Xeromyces; (2) Thermoascaceae for Paecilomyces (incl. Byssochlamys), and

Thermoascus (incl. Coonemeria, and Dactylomyces); and (3) Trichocomaceae

for Rasamsonia, Sagenomella, Talaromyces (incl. Erythrogymnotheca, and

Sagenoma), Thermomyces, Trichocoma, and tentatively for Dendrosphaera.

Four genera remained unplaced: Ascorhiza, Dichlaena, Pseudocordyceps, and

Sarophorum. While some might think the authors “jumped-the-gun,” as the

names Aspergillus and Penicillium are such widely used names compared

with those of the teleomorph-typified generic synonyms, the wording of the

Melbourne Cope can be taken as indicating they were not obliged to have

this approved by the General Committee on Nomenclature established by the

Congress. In any event, there are no nomenclatural penalties for their choices,

which follow the rules applying to all non-pleomorphic fungi. Following this

revision of the familial and generic concepts and names, more detailed accounts

of the subgenera and sections of Penicillium, with detailed lists of the species to

be included in them, are presented.

This overview is followed by a critical revision of Penicillium sect. Citrina by

Houbraken, Frisvad & Samson, based on molecular phylogeny and physiology

(especially extrolites produced). Thirty-nine species are accepted, of which 17

are described as new. Extrolite profiles and calmodulin and B-tubulin sequences

can be used for species separations, but only about half of the species could

be unambiguously identified by ITS sequences alone. Rivera & Seifert address

the P. sclerotiorum complex with an equally critical and polyphasic approach,

recognizing seven species, of which three are newly described; in this case ITS

barcodes satisfactorily separated the taxa. The final contribution by Samson

and seven colleagues addresses Talaromyces (incl. P. subgen. Biverticillium) and

considers all taxa described in the group; 69 species are accepted, one newly

described, many new combinations are made from Penicillium into the genus,

and a list of excluded names and synonyms is provided. As expected with Jens

Frisvad as one of the co-authors, detailed information is given on the extrolites,

with the “exometabolites” tabulated in detail.

All four contributions are superbly illustrated with coloured photo-

micrographs of colonies as well as fine shots of conidiophores and conidiogenous

cells. This is a carefully and well-produced work, which is a “must” for all workers

MycotTaxon 119 Book Reviews ... 489

on penicillioid fungi and their allies, and is sure to attract many citations. My

one regret is that no culture or morphology based keys are included. ‘This

means that it is becoming increasingly impossible to identify fungi in this

group precisely without molecular sequence data for several genes.

Davip L. HAWKSWORTH

Departamento de Biologia vegetal II, Facultad de Farmacia,

Universidad Complutense de Madrid, Plaza Ramon y Cajal, Madrid 28040, Spain

davidh@farm.ucm.es; d.hawksworth@nhm.ac.uk

BASIDIOMYCETES

Riisikad. The genus Lactarius in Estonia. By K. Kalamees, 2010. Natural History

Museum, University of Tartu. Available from Katrin Kolnes (katrin.kolnes@ut.ee).

ISBN 978-9985-4-0619-9. Pp. 187, many col. pl. Price 21 €

This modern treatment of the genus Lactarius in Estonia includes a short

introduction to the taxonomic placement of the genus, list of species recorded

for Estonia, an ecology chapter, illustrated overview of the main characters for

identification, and a key to the species. The key, which is illustrated, is in both

Estonian and English. The main part of the book is taken up by the 60 species

descriptions, with short descriptions of another 22 that might exist in the

country. A glossary, list of references, and species index conclude the book.

Each species is presented by a good colour photo (often several) to illustrate

colour variation and development. There are no microscopic details depicted.

The text accompanying the photos covers sequence data, herbarium data, and

ample ecological details. Original data are used throughout. A short English

characterization of each species is very helpful for non-Estonian speakers.

The number of species is smaller than that treated in the work by Heilmann-

Clausen et al. (1998) for northern Europe. But the extra information, the high

quality photos, and the low price make this a good choice. In short, a well

executed, well researched and very valuable contribution to our knowledge of

milk caps.

Heilmann-Clausen J, Verbeken A, Vesterholt J. 1998. The genus Lactarius. Fungi

of Northern Europe Vol. 2.

Slaktet Hygrophorus, Skogsvaxskivlingar i Sverige. En faltguide till SMF:s

svampvakteri “Vaxvakt”. By E. Larsson, S. Jacobsson & A. Stridvall, 2011.

MYKOLOGISKA PUBLIKATIONER 3. Sveriges Mykologiska Férening, Institutionen

for vaxt- och miljévetenskpaer, Géteborgs Universitetet, Box 461, 40530 Goteborg,

Sweden. <www.svampar.se> . 56 pp.

In 2011 the Swedish Mycological Society started a project to record and monitor

Hygrophorus species throughout Sweden. With only 15 species common and

490 ... Vellinga, BOOK REVIEW EDITOR

well known, many species are currently considered vulnerable or under threat

and included on the red data list.

To help identify the focus species, a small booklet has been produced

offering species keys with descriptions and colour photos of all 37 taxa that

enable field identification. The emphasis is on macroscopic characters, with

spore dimensions as the only microscopic characters presented. Notes on

ecology and distribution are given for each species, showing that many species

are host tree specific, and some notes on possible confusion with other species

complete the species information. There is also a recording sheet for the 15

rare species for which material should be sampled and sent to Ellen Larsson,

the first author. Though written in Swedish, the booklet is easy to understand

with some help from modern translation web sites and knowledge of another

Germanic or Scandinavian language.

GENERAL

Systematics and evolution of fungi. By J.K. Misra, J.P. Tewari & S.K. Deshmukh

(eds), 2012 (‘2011’). PRoGREss IN MYCOLOGICAL RESEARCH Vol. 2. CRC Press,

Taylor & Francis Group, 6000 Broken Sound Parkway, NW, Suite 300, Boca Raton,

FL 33487, U.S.A. <www.crcpress.com> ISBN 978-1-57808-723-5. Pp. 412, Pl. 2.

Price US$119.95.

The second book in a series of four on ‘Progress in Mycological Research

focuses on systematics and phylogenetics, and 13 individual chapters treat

various fungal groups.

An introduction by Hawksworth in the first chapter presents the merits and

problems surrounding sequence-based systematics. Chapter 2 is devoted to

fossil fungi, Chytridiomycota are treated by Powell and Letcher (Chapter 3),

and Benny gives an overview of the Zygomycota, their systematics, and possible

segregation into four groups (Chapter 4). Lichtwardt (Chapter 5) presents the

evolution of the Trichomycetes, followed by Misra (Chapter 6), who covers the

systematics of the two largest genera of the Harpellales (Stachylina and Smittium)

along with keys and species descriptions. Chapter 7 focuses on two genera of

edible mushrooms, Morchella and Macrolepiota, in Israel, and Barseghyan et

al. try to delineate species based on ITS and EF-lalpha sequence analyses but

describe no new species. Chapter 8 highlights morphological characters that

distinguish groups of mushroom forming fungi, and authors Zmitrovich &

Wasser warn that groupings based solely on RNA genes should be considered

with skepticism. Tura et al. (chapter 9) gives an overview of Phellinus s.l. and

Inonotus s.l. in Israel based on ITS sequences and morphology. In Chapter

10 Uli-Mattila, who discusses toxigenic Fusarium species from small cereal

MycotTaxon 119 Book Reviews ... 491

grains, advocates a combined approach using all available characters for

species recognition, while Gannibal (Chapter 11) presents the consequences

of phylogenetic studies of alternarioid hyphomycetes. Tewari et al. (Chapter

12) shift gears to note methods for rapidly diagnosing Candida-related human

pathogens. In the last chapter, Nagy et al. provide an overview of various

phylogenetic methods and models to unravel the evolution in fungi.

Many articles are reviews and literature compilations, but some new

research is presented. In all it gives an idea of the state of knowledge of the

groups covered here.

REGIONAL FLORAS

Fungi of tropical China. By X.-L. Wu, Y.-C. Dai, T.-H. Li, Z-L. Yang & B. Song,

2011. Science Press, Beijing, <www.sciencep.com>. ISBN-13: 9787030294708.

Pp. 548, col. plates 495. Price circa US$ 220 [480.00 Yuan]

It is a pleasure to leaf through this book full of good mushroom photos from

tropical China. In total 495 species are depicted, out of the 2065 listed for the

area in the back. The book, authored by five mycologists and written in Chinese

with Latin nomenclature and English chapter headings, gives an overview of

the habitats (with their fungi), a checklist of all tropical fungi in China, and an

extensive reference list; the main part of the book is devoted to species photos

and descriptions. On the right hand page are two or three photos opposite the

description on the even numbered pages. The majority of the taxa belong to

the basidiomycetes. The species are ordered according to their phylogenetic

position, for instance Auriscalpium can be found near the genus Russula. It

is really nice to see so many ectomycorrhizal species depicted; Amanita, for

instance, is well represented by 37 species and at least 35 species of boletes are

included. Among the wood inhabiting fungi, Ganoderma stands out with 30

species, while stinkhorns in all their weird beautiful shapes and colours are

represented by 21 species.

It is obvious that in some groups, names from species described from other

regions have been applied to southeastern Asian specimens and are not always

correct. Two such misapplied names are ‘Lactarius deliciosus' and ‘Anthurus

archeri, and the photo of Suillus bovinus does not look typical for that species.

Most species have been photographed in the field, but lab photos have been

used for a very small number.

The book shows the diversity and richness of fungi in tropical China, and

can certainly be used outside China. I encountered many taxa that are familiar

from northern Thailand. I am looking forward to seeing an English translation

of this book to make it more accessible to the rest of the world.

492 ... Vellinga, BOOK REVIEW EDITOR

Mycota of Rhode Island: A checklist of the fungi recorded in Rhode Island

(including lichens and myxomycetes). By R.D. Goos, 2010. THE Biota

OF RHODE IsLAND vol. 4. Rhode Island Natural History Survey, P.O. Box 1858,

Kingston, RI 02881, <programadmin@rinhs.org>. ISBN 1-887771-09-3. Pp. 228.

Price US$60.00

Strangely enough, there do not seem to be many checklists of fungi for the

US.A., either for separate states or as a whole. I tried to find such lists on line

but was only able to find the database for plant pathogenic fungi at the U.S.

National Fungus Collections. For European countries, checklists of macrofungi

are seen as a prerequisite for recording and mapping programs, on which

conservation efforts and red data lists can be based.

The present list for the state of Rhode Island, the smallest of the 50 states

(comparable in size to the Cape Verde Islands), encompasses all fungi, from

Chytridiomycota and Glomeromycota to Basidiomycota; slime molds and

Oomycota are also covered. ‘The list is restricted almost exclusively to verified

records and based on a number of different sources (herbaria, literature). The

presence of voucher specimens is a must.

A short history of the mycology practiced in the state forms the prelude to

the main part of the book with the species lists. For each species the following

information is given: full name including authority, host and substrate, an

indication of distribution and rarity, and references and or herbarium records.

The book is illustrated with nice line drawings by Roberta Calore.

It is great to have this information available, but of course this is the kind of

data that should be available for all, and searchable in various ways on line.

Book ANNOUNCEMENTS

Biomonitoring, Ecology and Systematics of Lichens. Recognizing the Lichenological

Legacy of Thomas H. Nash III on his 65th Birthday. By S.T. Bates, F. Bungartz, R. Licking,

M.A. Herrera-Campos & A. Zambrano (eds), 2011. BIBLIOTHECA LICHENOLOGICA

Band 106. ISBN 978-3-443-58085-8. Pp. 442, col. pl. 16, figs 102. Price 109 €

Flora Fungorum Sinicorum Volume 39. Tilletiales Urocystidales Entorrhizales

Doassansiales Entylomatales Georgefischeriales. By G. Lin, 2011. ISBN 9787030314536

Pp. 152 p., figs. Price around US$ 49.00.

Guia de hongos de la peninsula ibérica. By G. Moreno & J.L. Manjon, 2011. Ediciones

Omega, S.A. Calle Platé, 2608006 Barcelona, Spain. <www.ediciones-omega.es>. ISBN

978-84-282-1349-3. Pp. 1440, 590 col. pl. Price 100 €

MYCOTAXON

http://dx.doi.org/10.5248/119.493

Volume 119, pp. 493-494 January-March 2012

Regional annotated mycobiotas new to www.mycotaxon.com

Mycotaxon is pleased to announce the addition of five new species distribution lists to

our “web-list” page at www.mycotaxon.com//resources/weblists.html, where a total of 89

checklist PDFs are now available for free download. The content and design of each list

is the sole responsibility of its authors and their three (or more) expert reviewers. Those

meeting MycoTaxon’s scientific and nomenclatural criteria are accepted for posting. The

authors, titles, and abstracts of our most recently accepted web-lists are provided below.

SOUTH AMERICA

Argentina

N. Niveiro & E. Alberté. Checklist of the Argentine Agaricales I. Amanitaceae, Pluteaceae

and Hygrophoraceae. 24 p.

Asstract — A checklist of species belonging to families Amanitaceae, Pluteaceae,

and Hygrophoraceae was made for Argentina. The list includes all species published

till year 2011. Eleven genera and 150 species were recorded. The family Pluteaceae is

the most cited for the country with 93 species.

Brazil

Isabella Pereira de Melo Wanderley Costa, Marcia Maria Costa Assuncao, Thais

Emanuelle Feijé de Lima, Rafael José Vilela de Oliveira & Maria Auxiliadora de

Queiroz Cavalcanti. Checklist of endophytic fungi from tropical regions. 26 p.

ABSTRACT — Endophytes have great ecological and biotechnological importance and

occur in a wide variety of plant families. This paper lists the genera and species of

endophytic fungi from tropical regions with their hosts and references.

EUROPE

Czech & Slovak Republics

Alena Novakova, Alexandra Simonovicova & Alena Kubdatova. List of cultivable

microfungi recorded from soils, soil related substrates and underground

environment of the Czech and Slovak Republics. 186 p.

ABSTRACT — A current list of the published microfungal records isolated from

various soils and soil related substrates are presented from the territory of the Czech

and Slovak Republics (formerly Czechoslovakia). Rhizosphere, entomopathogenic,

494 ... New regional mycobiotas online

ovicidal, nematophagous, coprophilous, keratinophilic, dermatophytic, and

thermoresistant fungi are also reported. The review contains microfungal species

from peat and peat substrates, litter, vermicompost, enchytraeid and earthworm

intestine and casts, and cattle environs, as well as records from soils affected by

windthrow, fire, heavy metals and human activity. In addition, microfungi which are

isolated from various substrates from underground environments, with the exception

of airborne fungi, are included. A total of 1,332 microfungal records are reported in

this list (54 of them belonging to yeasts or yeast-like microfungi and 1,278 belonging

to filamentous fungi), representing 385 genera and 1,283 species of microscopic

fungi. One microfungal taxon belongs to Microsporidiomycota, seven taxa belong to

Chromista, 164 to Mucoromycotina, 1,133 to Ascomycota, and 28 to Basidiomycota.

Forty-nine records were reported as undetermined fungi or sterile mycelia.

MID-EAST

Iran

Masoomeh Ghobad-Nejhad & Nils Hallenberg. Checklist of Iranian non-gilled/

non-gasteroid hymenomycetes (Agaricomycotina). 41 p.

ABSTRACT — This list includes 395 species of Iranian non-gilled/non-gasteroid

hymenomycetes. Corticioid, poroid (polypores), hydnoid, clavarioid, cantharelloid,

thelephoroid, and cyphelloid fungi are covered, together with heterobasidiomycetes.

Almost all the species are supported by at least one herbarium specimen. ‘The list

has been compiled mainly from material collected by the authors, verified reports

and literature. Nineteen species are reported for the first time from the country,

and 118 species are added to the most recent account of aphyllophoroid fungi of

North Iran by Hallenberg (1981, Mycotaxon 12: 473-502). The occurrence in Iran

of six species is regarded as doubtful, while 12 species are excluded. The number

of species known for each province in Iran is listed, with Golestan shown to be the

most species-rich province. The total number of species recognized for major Iranian

phytogeographical territories is also noted and Funalia trogii, Inonotus hispidus, and

Schizophyllum commune are shown to be the most widespread species. A permanent

repository for continuous updates has been made available via the Myco-Lich website,

and future new records for the country will be published online. Users can access

the latest changes to the checklist via subscription to a feed (RSS). The link to the

checklist is http://www.myco-lich.com/mycology-of-iran/basidiomycota/checklists/

National-checklist.

Turkey

Bahcecioglu, Zeliha & Sanli Kabaktepe. Checklist of rust fungi in Turkey. 81 p.

ABSTRACT — This study presents a checklist of rust fungi (Uredinales) of Turkey

together with their known host plants. The checklist enumerates 351 species of rust

fungi belonging to 26 genera and 9 families. Including 778 host plants belonging to

325 genera and 63 families. A host index is also provided.

MY COTAXON

http://dx.doi.org/10.5248/119.495

Volume 119, pp. 495-507 January-March 2012

Minimizing the chaos following the loss of Article 59:

Suggestions for a discussion

WALTER GAMS'’, RICHARD A. HUMBER’, WALTER JAKLITSCH?,

ROLAND KIRSCHNER‘ & MARC STADLER>™

" Molenweg 15, 3743CK Baarn, Netherlands

? USDA-ARS BioIPM Research, RW Holley Center for Agriculture & Health,

538 Tower Road, Ithaca, N. Y. 14853-2901, USA

> Department of Systematic and Evolutionary Botany, University of Vienna,

Rennweg 14, A-1030 Vienna, Austria

* Department of Life Sciences, National Central University,

No. 300, Jhongda Rd., Jhongli City, Taoyuan County 32001, Taiwan (R.O.C.)

° Helmholtz Institute of Infection Research (HZI), Dept Microbial Drugs, Braunschweig, Germany

* CORRESPONDENCE TO: walter.gams@online.nl

‘ (authors in alphabetic sequence)

ABSTRACT — We assert that the suppression at the 2011 Melbourne Botanical Congress of

the dual nomenclature for pleomorphic fungi was premature since most fungal genera still

remain inadequately typified and need molecularly based revisions. The new system turns a

rule-determined nomenclature into a system of names to be decided by committees. The new

system cannot be implemented until experts publish well-documented rationales about the

generic names to be accepted or suppressed among alternative morphs and the responsible

committees act officially. Prioritization vs. suppression is not the same as conservation vs.

rejection. If a teleomorphic generic name is suppressed, it still remains valid and legitimate

and can still serve as a basis for names of higher-rank taxa. In most cases the teleomorph genus

name should be prioritized unless convincing arguments favor preference of the anamorph

name. Paraphyletic genera must be recognized. Unless the phylogenetic positions of all type

species for possible morphs are confirmed, no prioritization can be fixed. When a binomial

in a prioritized genus has a younger epithet than the corresponding name in the suppressed

genus, only priority of extant names in the prioritized genus should count.

KEY woRDs — anamorph-teleomorph connection, competing morph names

Unification of fungal nomenclature (one name, whether teleomorphic or

anamorphic, is allowed for each taxon) had to come eventually (Hawksworth

et al. 2011, Norvell et al. 2011), but the move is being enforced much too early

while only a minority of fungal taxa has been examined phylogenetically

496 ... Gams & al.

(Hibbett et al. 2007). As currently circumscribed, too many fungal genera

are unconfirmed as homogeneous and monophyletic (such proofs may not be

possible for some genera), and knowledge of correlations between anamorph

and teleomorph genera is far from comprehensive (Gams et al. 2011); however,

Hawksworth et al. (2011) cited the ability to link anamorphs and their

teleomorphs by molecular means as a key reason for the rapid adoption of the

new rules. Names of lichenized fungi have always been excluded from dual

nomenclature and are little affected by the issues discussed here (Lendemer

2011). While the previous Article 59 might not be reinstated (and cannot

even be considered until the 2017 International Botanical Congress), we must

find solutions to minimize the disruptive effects of this decision for fungal

nomenclature and systematics. That the new procedures will depend on

committee decisions listed in databases rather than on adherence to clear-cut

rules of a CODE will obviously be a source of conflicts.

A key sentence in Paragraph 1 of the PREAMBLE in the ICBN (McNeill et al.

2006) is crucial to our argument: “This Code aims at the provision of a stable

method of naming taxonomic groups, avoiding and rejecting the use of names

which may cause error or ambiguity or throw science into confusion.” This

fundamental principle seems to be ignored by what has happened: Previous

editions of the Cope established democratically accepted rules subject to

adjustment every six years by general voting in the Nomenclature Sessions.

The newly prescribed rules and processes for determining the correct names

for pleomorphic fungi reject these long understood, fair rules in favor of non-

appealable decisions rendered by a series of Committees. It is still unknown

how these newly adopted processes will be put into practice. Language adopted

in Melbourne requests “consultation with” but does not actually mandate the

involvement of non-IAPT committees (that have no legal standing vis-a-vis

the CoDE). NOMENCLATURAL STABILITY, a principle long stated in the CobE,

seems to have been abandoned in favor of committee-based decisions governing

the choices and applications of names for exactly those fungi whose complex

biologies and multiple reproductive forms need governance by clear rules, not

by consensus in committees.

Gene-based data can never be the sole basis for meaningful, informative

classifications of any organism without also incorporating the essential

foundations of morphology and other traditional characters. Imperfectly

known taxa (new or old) need to be recollected for more study of their genes

and morphologies before being reclassified. Fieldwork and ecological and

morphological studies will be indispensible to the additions into GenBank of

genetic data for innumerable neglected taxa. Deposits of accurate gene sequence

data from incorrectly identified source organisms is a widespread problem that

can confuse and seriously damage later analyses based on false assumptions.

Minimizing nomenclatural chaos after Art. 59... 497

While environmental DNA sampling might indicate that fungi are present, it

reveals nothing about the appearance and biology of the fungi from which that

DNA came, how to recognize them, or whether and how these fungi interact

with any other organisms.

Some supporters of the one-fungus one-name (1F=1N) concept seem

to assume that all fungi will readily yield good DNA and clean sequences.

However, many entomophthoraleans resist repeated attempts by skilled

technicians to obtain clean DNA and useful amplifications of genes now

widely accepted (and expected) to provide phylogenetically informative data.

Some taxa in the Xylariaceae yield either no cultures from too-scanty available

materials or no useful PCR products from even freshly collected stromata.

Further, some xylarioid stromata have at times provided contaminant DNA

that was later shown to represent different taxa of the same or closely related

xylariaceous genus (Stadler et al. 2010, Stadler 2011). Although Xylariaceae

harbors few known endophytic species, some do grow hyperparasitically on

xylarioid taxa; consequently, DNA extraction from stromata should be strongly

discouraged, given that which organism provides the DNA sequence within a

xylarioid stroma may remain uncertain. Endolichenic fungi may cause even

more confusion (Arnold et al. 2009). As fungi generally grow in complex

aggregations in nature, often only a skilled morphologist can ascertain that a

culture actually represents what was thought to be the original inoculum. Some

fungi may never be culturable unless new techniques are developed for isolation

and growth, while special methods may also be needed to purify and to amplify

their DNA. Anamorph-teleomorph relationships can hardly be proven in such

cases, and phenotype-based taxonomic concepts MusT be maintained for the

time being.

Implementation of the new rules

New paragraph approved in Melbourne:

Art. 57.2 (new) In pleomorphic fungi (including lichenicolous fungi, but

excluding lichen-forming fungi and those fungi traditionally associated with

them taxonomically, e.g. Mycocaliciaceae), in cases where, prior to 1 January

2013, both teleomorph-typified and anamorph-typified names were widely used

for a taxon, an anamorph-typified name that has priority is not to be taken

up until retention of the teleomorph-typified name has been considered by the

General Committee and rejected [OUR EMPHASIS].

This rule implies that the duality of names should not be suppressed until the

responsible committees have rendered and publicized their decision about

the preferential use of a particular name. Thus, mycological editors should

be patient and tolerant with authors who may still prefer to use teleomorph-

typified generic names, even if the corresponding anamorph name is older.

A498 ... Gams & al.

Prioritization—choosing between competing morph names for an individual

pleomorphic fungus—vs. conservation.

New text passed in Melbourne to appear in the ICN:

Art. 14.13 (new) In the interest of nomenclatural stability, for organisms

treated as fungi (including lichenicolous fungi, but excluding lichen-forming

fungi and those fungi traditionally associated with them taxonomically, e.g.

Mycocaliciaceae), lists of names may be submitted to the General Committee,

which will refer them to the Nomenclature Committee for Fungi (see Div. II) for

examination by subcommittees established by that Committee in consultation

with the General Committee and appropriate international bodies. Accepted

names on these lists, which become Appendices of this CopE once reviewed

and approved by the Nomenclature Committee for Fungi and the General

Committee, are to be listed with their types together with those competing

synonyms (including sanctioned names) against which they are treated as

conserved (see also Art. 56.3).

PRIORITIZATION. We strongly suggest referring to the choice among competing

morph names as prioritization, an action that is not equivalent to conservation.

While Art. 14.13 requires such choices to be “treated as conserved,’ such status

would confer permanent validity to frequently misapplied names.

To clarify this, Art. 14.13 should be amended to deal with suppressed

names. Among competing anamorphic (occasionally synanamorphic) and

teleomorphic names for the same taxon, one of the “name pair” will be given

preference while the use of the other(s) is suppressed. Although suppressed

names are not available for use, however, they remain nomenclaturally valid

and legitimate. This condition differs fundamentally from the status of a NOMEN

REJICIENDUM. In contrast to conservation, an act of prioritization need not be

irreversibly fixed if convincing arguments support a change.

Consequently, names of teleomorphic genera that may be suppressed in

favor of associated anamorphic generic names remain available as bases for

HIGHER-RANK NAMES and would not disrupt the current general systematics of

Ascomycota that would be severely affected by the issues discussed here.

Until July 31, 2011, teleomorph-typified names had precedence over

anamorph-typified names irrespective of temporal priorities. Under the new

rules, priority of publication should generally determine the choice. It is,

however, true that the great majority of names for fungal families (and higher

ranks) are based on teleomorph-typified genera. Because all teleotypified

generic names remain legitimate and available as a basis for higher-rank taxon

names, there is no objection to this (Art. 18.3 ICBN), and no changes at higher-

rank names will become necessary for purely nomenclatural reasons. Basing

any higher-rank names on anamorph-typified generic names might be desirable

in some cases, but it must be discussed. If, for example, the name Trichoderma

Minimizing nomenclatural chaos after Art. 59 ... 499

were prioritized over Hypocrea, should this require introducing the names

‘Trichodermataceae, ‘Trichodermatales, etc.¢ Because family names adhere to

the rule of priority, would the name “Trichodermataceae’ have precedence over

Hypocreaceae simply because of the priority of the genus name?

The new rule abandoning the preference of teleomorph-names for higher

ranks of fungi may encourage the description of many higher taxa (families,

orders, etc.) for clades of exclusively anamorphic Ascomycota. This would be a

new — and undesirable — situation. Certain anamorph names have later been

correlated with symplesiomorphic characters that have little or no use in formal

taxonomy but do serve as descriptive terms.

For the prioritization of single or listed fungal names well-documented

PROPOSALS must be submitted and published, e.g. in Taxon, as well as posted

on an appropriate website. It might be helpful to first propose a preliminary

prioritization that would become definitive only after a certain period of

testing (e.g., after 2 years).

Decisions about prioritization will be made by authorities comprising a)

a committee of experts (the largest, most diverse group reasonably possible),

b) the International Committee for the Taxonomy of Fungi (ICTF), and c) the

IBC Permanent Nomenclature Committee for Fungi (NCF). The NCF then will

report to the General Committee. A majority of at least 60% of the votes (the

NCF present practice—including abstentions, but suggested for (a) excluding

abstentions) should be required for a decision. Art. 14.13 alludes to committees

a) and b) as “appropriate international bodies.” A primary concern is, who will

bear the responsibility of establishing individual committees of experts?

The size and scope of any expert committee must vary according to the

fungi under consideration to assure diverse points of view and geography; the

constitution of these committees should and must be debated — and consensus

reached — before they are established. The greatest problem will be finding

mycologists competent in all fungal groups to be tackled. If enough experts are

not found for a particular group or if no molecular analyses are available, then

no prioritization of names should be decided. It will be a task of each expert

committee's members to draft proposals of preferred names and vote on them.

For example, Hypocreavs. Trichoderma is being debated within the International

Subcommission on Trichoderma and Hypocrea Taxonomy (ISTH); its outcome

will be reported to the ICTF and NCE, who may (or may not) confirm it and

publish their decision. In this case, the cellulase producer Trichoderma reesei and

some other biocontrol strains of Trichoderma spp. are commercially important.

Although the name of the T. reesei correlated teleomorph, Hypocrea jecorina, is

also becoming widely used, preference would be given to Trichoderma, which

is the older name anyhow. Does a preference for Trichoderma then force species

500 ... Gams & al.

like H. pulvinata (in the anamorphically little-differentiated Trichoderma sect.

Hypocreanum) and H. danica, H. peltata, H. rhododendri, H. spinulosa, etc. (no

anamorphs) (Jaklitsch 2011) into Trichoderma?

The problem is that there are a) several groups of fungal taxa on which

nobody is really working and b) other groups that are being well studied but

by different authors with entirely incompatible concepts. How can a committee

scenario work in these cases?

The implication that “expert groups” shall join in committees to treat their

respective fields of specialty may pose serious problems. Taxonomic concepts

in the Eumycota have always been highly variable (including even the name

for all fungi). The introduction of molecular phylogenies raised the hope that

such controversies might be resolved, but we now know this will not be the

case. In fact, progress with molecular phylogenetic studies for diverse higher

fungal taxa has been highly variable and depends on the variably patchy

funding available to taxonomic specialists in different fungal groups. Sound

molecular phylogenies are very difficult to achieve for innumerable fungal

taxa for which most constituent species are neither epitypified nor available

in cultures. Many who supported implementation of the 1F=1N concept seem

not to have realized that morphological and other phenotypic data still provide

more information on which to base classifications, especially where a limited

number of sequenced genes fails to provide convincing proof of relatedness.

Itis gravely worrying that TOO MANY new layers of authorities whose opinions

may differ (and, indeed, be mutable) are injected into what was previously a

process with universally recognized rules based on priority and an established

line of authority to achieve stable, enduring nomenclatural decisions. The

new system of committees adjudicating names EN MASSE means that research

findings not favored by a committee can simply (and dangerously) be “voted

out” of official recognition.

Suggested general working rules for committees

Names of genera

Anamorph and teleomorph genera were hitherto based on anamorph and

teleomorph traits, respectively. Adoption of unitary nomenclature necessitates

emending all pleomorphic generic diagnoses to recognize both anamorphic

and teleomorphic characters, whether present or not. All mycologists should

welcome such improvements because separating information about alternate

morphs into multiple descriptions fails to communicate vital and complete

information about the whole organism. Although we have often recognized

some degree of polyphyly in many important anamorphic genera, we have until

now preferred that teleomorphic genera be (as far as possible) monophyletic.

The inverse situation may arise, however, when a more narrowly defined

Minimizing nomenclatural chaos after Art. 59 ... 501

anamorphic genus takes precedence over a teleomorphic one. In that case, the

teleomorph genus may continue to be usedas an artificial “dustbin” ofteleomorph

species with or without known anamorphs. For example, many Mycosphaerella

anamorphs may be submerged into some morphologically and phylogenetically

well-defined anamorphic genera such as Cercospora and Ramularia (Crous

et al. 2009, Crous 2010), thus forcing Mycosphaerella to become a “dustbin”

for species without known anamorphs. An equally undesirable alternative

would be combining many hundreds of species from several anamorph genera

into Mycosphaerella, which already contains ca. 2000 species. Morphological

comparisons among species would become impossible in such a huge genus

unless anamorph characters were correlated with phylogenetic ones. To avoid

such problems, will we be forced to accept gigantic genera characterized only

by statistics based on (minimal) DNA data?

If Puccinia were prioritized over Uredo s.str., the synonym ‘Uredo’ would

not be available for species with different teleomorphs, thus requiring the

proposal of new generic names to accommodate them. Art. 14.13 may need to

be amended in order to repair this undesirable situation.

Serious problems will arise for teleomorph genera such as Cordyceps and its

recent segregates, each of which is associated with multiple distinct anamorph

genera — many of which may form distinct clades and have names with

nomenclatural priority over their linked teleomorph genus. Further splitting of

(teleomorphic) genera may become advisable in such cases rather than lumping

multiple disparate anamorphs within one teleomorphic genus.

This situation is exemplified by the relatively rare Asian teleomorphic genus

Metacordyceps G.H. Sung et al. (Hypocreales: Clavicipitaceae) (Sung et al. 2007;

also see Kepler et al. 2012) whose anamorphs include the globally distributed,

commercially important entomopathogenic genera Metarhizium Sorokin

1883 and Nomuraea Maubl. 1903, as well as Pochonia Bat. & O.M. Fonseca

1965 (including species used for biocontrol of nematodes) and other species

not placed in any anamorphic genus after the phylogenetic reclassification of

Paecilomyces sect. Isarioidea Samson 1974 (Luangsa-ard et al. 2005, Sung et

al. 2007). That these anamorph genera are phylogenetically and ecologically

distinct may justify the preferential use of the anamorph names. Nevertheless,

the younger teleomorph name Metacordyceps also carries relevant information.

Although we generally prefer to retain precedence for the teleomorph-

typified generic name over its correlated anamorphic name, we must accept

exceptions.

Exceptions to the precedence of teleomorph-generic names

Exceptions from this rule are allowed, although objective criteria are not

yet clearly worded in the new Art. 57.2. To the Trichoderma-Hypocrea and

Metarhizium-Metacordyceps examples noted above, we add the so-called

502 ... Gams & al.

“macromycete” groups (those forming conspicuous stromata) in Ascomycota

for possible exception.

Some Hypocreales (as in other ascomycete groups with economically

important species, such as plant parasites or industrially cultivated fungi) have

already been studied thoroughly by molecular methods. Taxonomic authorities

familiar with these fungi will tend to follow the molecular results to adjudicate

the status of the core taxa. However, their anamorph names for some of these

fungi may prevail in future nomenclature, because the anamorphic stages

that cause the conspicuous disease symptoms were described long before

their corresponding teleomorphs. Nonetheless, nearly all xylarialean experts

will admit that the number of taxa whose molecular characters have already

been thoroughly evaluated is wholly insufficient to make final nomenclatural

determinations, since the phylogenies based on housekeeping genes vs. rDNA

present wholly divergent results about relationships among these fungi. The

numerous experts (all also signatories of Gams et al. 2011) will surely agree

that there is no need to change the current taxonomic system. Anamorphs

of Xylariales are widely known but treated as synapomorphies, so that dual

nomenclature was abandoned for these genera decades ago, whereas such

well-known and conserved generic names as Xylaria, Hypoxylon, and Daldinia

continue to take preference. Their anamorphic names are actually used as

features that characterize subfamilies (or, in the future, perhaps, families or

orders). The so far prevailing preference for teleomorph names could or should

be adapted for all Ascomycota unless very serious reasons (including priority)

make it more practical to adopt the anamorph name (as for Penicillium and

anamorphs of Metacordyceps).

The latest revision of Penicillium (Samson et al. 2011) provides more

examples: the teleomorphic name Eupenicillium has had little application

or relevance, with Penicillium already treated as holomorphic by Raper &

Thom (1948); however, for species of the more distantly related teleomorphic

Talaromyces (whose anamorphs in Penicillium sect. Biverticillium still needed

a new generic name), the teleomorph name was chosen to replace the better

known anamorphic names in Penicillium. Must all these cases be decided again

in the newly prescribed High Courts of Mycology by jurors yet to be selected?

In the future, teachers of mycology may be hard pressed to explain to their

students why some genera of the Xylariaceae have nodulisporium-like or

geniculosporium-like anamorphs but those of the “Trichodermataceae” (or

“Trichodermatales”) have Hypocrea-like teleomorphs. Recruitment of young

fungal taxonomists is already very difficult; those who led the 1F=1N movement

and championed the decisions adopted in Melbourne even used this difficulty

as a major argument in their favor. Teaching of mycology MIGHT become more

complex if suprageneric taxonomic ranks were often based on anamorphic

Minimizing nomenclatural chaos after Art. 59 ... 503

names, but such an argument seems to give little credit to the ability of students

to understand these complications.

A further consequence of complex connections between anamorph and

teleomorph genera is the necessary recognition of paraphyletic genera, such

as Lecanicillium, which Gams & Zare (2001) described, at the same time

recognizing its paraphyly. The current PCR-driven, monophyletic world view

also seems to reject the perfectly valid, widely held position that paraphyletic

taxa are not only acceptable, but that recognizing them as such is natural and

appropriate. Zoologists, who have long accepted the reptiles as a transitional

group from which the birds have arisen, do not deny Reptilia its own taxonomic

status, even where both molecular and comparative anatomic studies prove

that birds + reptiles are a paraphyletic class. It is hard to grasp why some

mycologists fervently want to impose strict monophyly on all fungal taxa even

where gene-based data supporting such a conclusion are unavailable or may be

unobtainable) (Hérandl & Stuessy 2010).

When a new teleomorph is discovered for a known anamorph, it will no

longer be possible to describe it in a new genus unless that species is proven

phylogenetically not to be congeneric with the type of the anamorph genus.

If the type species of a genus has not been studied by gene-based methods,

the genetic homogeneity of the genus cannot be proven. Similarly, no unification

of anamorphs and teleomorphs can be forced if the type species of both the

anamorph and teleomorph genera have not been analysed.

Names of species

If the rule of priority were strictly applied to epithets in both teleomorphic

and anamorphic genera, many name changes would ensue unless a particular,

well-established name is ‘rescued’ by formal conservation. To avoid this, we

suggest a procedural rule for the committees (probably eventually for a new

article in the Cope) that binomials established before July 31, 2011 for either

morph type must not be replaced by combining older epithets available in the

‘opposite’ morph (e.g., Trichoderma reesei should not become ‘T. jecorinum’ and

T. citrinoviride should not become “T. schweinitzii’ despite the older available

name Hypocrea schweinitzii). If Pleospora were retained over Stemphylium,

would the new combination ‘Pleospora botryosa’ then have to replace the

established but younger P. tarda? In parallel with the ICBN Art. 11.2, such a

rule would stipulate that priority applies only to names of the same rank or the

same prioritized genus.

Another problem concerns species for which no sequence data are available.

Can they be recombined into the alternative morph’s genus or not? Would

they remain in limbo? After proving the linkage of the type species of both

anamorph and teleomorph genera, it would be better to combine all other

names from the suppressed genus into the prioritized genus to keep them

504 ... Gams & al.

visible and usable, even if those types were never restudied by any means. If not,

they will be forgotten, any further revisions of them would halt, and homonyms

or synonyms might be created in the prioritized genus.

Such issues are further complicated because innumerable older (and many

highly important) species were not typified when described or their type

specimens are lost; these taxa would require lecto-, neo-, and/or epitypifications

before drawing any gene-based taxonomic conclusions about their status. The

problems raised by so many fungi that were described from single collections

and that may be extremely rare or difficult to re-collect again for DNA-based

studies are nothing less than a daunting impediment for the establishment of a

workable 1F=1N taxonomy.

Conclusions

A major aim of the one-fungus one-name policy was to simplify fungal

nomenclature. The question, however, remains: simplification for whom? For

the convenience of present-day mycologists, it would be desirable not to change

the names of higher taxa. When in extreme cases an anamorph-generic name

replaces the type genus of teleomorph-based higher taxa (e.g., Hypocreales,

Hypocreaceae), we still remember the meaning of the higher taxon because

we are used to it. The situation becomes complicated for future generations,

however, who must deal with many previously teleomorph-based higher taxa

when the teleomorph genus is no longer in use; mycologists must then learn

that these names were based on a now-rejected nomenclatural practice and try

to understand the superseded taxonomies BEFORE they can apply the systematic

concepts that led to the current use of a name. The intended “simplification”

imposed by the 1F=1N standard will thus not be achieved in a long-term

vision, and future mycologists will still need to understand the suppressed dual

nomenclature to deal with the historical literature. Moreover, many recently

introduced teleomorph-generic names do reflect improved phylogenetic

knowledge, although they will not successfully compete with the correlated

anamorph names; these names undoubtedly will retain their informative value

in naming clades and should not, therefore, be completely eliminated.

In a carefully balanced system of prioritization, future mycologists may

appreciate having the choice between either continuing to use the present

higher-rank taxon names or creating new anamorph-based names.

An essential, permanent remnant of the dual nomenclature will be the

need for listing all teleomorphic and anamorphic synonyms of a fungus in any

monographic treatment together with the typifications of all these names. It

will often be the case that more species were described under the suppressed

morph-generic name than in the officially recognized morph genus. In any case,

all these suppressed species will have to be re-studied before any new species can

Minimizing nomenclatural chaos after Art. 59... 505

be described. This does not simplify future taxonomic work. Re-examination of

types must be possible any time, and this will involve morphological (including

ultrastructural), biochemical, and molecular approaches equally, all still subject

to improvement. Curiously, it is often more valuable to preserve teleomorphic

material of most ascomycetes in a herbarium than their anamorph which, if the

type is a dried agar culture, can display its properties only when reactivated in

culture.

In a unified nomenclature, fungi will continue to have complicated life

cycles displaying a teleomorph and one or more anamorphs, each with its

distinctive gene expression and biological and ecological role. Future students

of mycology will have to understand the concept of anamorph and teleomorph

to understand the biology of the fungus, even if all stages are encompassed

under a single name.

The first paragraph of the Preamble of the ICBN contains an odd and

ominously predictive stipulation immediately before the sentence quoted

above: “The purpose of giving a name to a taxonomic group is not to indicate its

characters or history [our emphasis added], but to supply a means of referring

to it and to indicate its taxonomic rank.” It seems probable that few mycologists

who deal with pleomorphic fungi would appreciate the strict application

of such a stipulation. The nomenclaturally required consolidation of many

morphologically disparate teleomorphic and anamorphic forms and even genera

into single genera now required by the newly adopted standard of 1F=1N may

satisfy both the new Cope and advocates of strict monophyleticism but this

process will create many significantly large and diverse — but nomenclaturally

supported! — genera that will also be self-accreting mycological “black holes.

The names of such ‘megataxa will bear no useful biological information

about their species. While this result will conform with the newly redefined

(if artificial) sense of nomenclatural stability, it cannot be hailed as progress

toward biological or even taxonomic sensibility.

We should also note that the availability of cultures of hitherto unknown

fungal organisms is an absolute prerequisite for their future exploitation in

biotechnological applications. Fungi have historically been used for many

beneficial purposes; curiously, the roots for the new concepts and changes

adopted in Melbourne seem to have been elaborated mainly for organisms that

were studied intensively for their diverse beneficial or detrimental properties

— e.g., pathogenicity, production of mycotoxins or other primary or secondary

gene products, uses as biological control agents, or many other potential

concerns. We should realize that in view of future applications, we have so far

only dealt with the low-hanging fruits. Large-scale bioprospecting programs

are now under way, and potential uses for fungi are by no means restricted to

the traditional search for novel, badly needed antibiotics and other compounds

506 ... Gams & al.

with potential utility in pharmaceutical or agrochemical indications. The search

for industrial enzymes and commodity chemicals for “White Biotechnology’,

as well as renewable resources (biofuel production), bioremediation, etc., will

almost certainly become more important in the future. Innovative methods of

genomics research will allow for the exploitation of slow-growing organisms,

e.g., by transfer of their genes into heterologous hosts where biotechnological

applications appear feasible.

Acknowledgments

We are grateful to John McNeill for communicating and explaining the new texts

of Articles 14.13 and 57.2. Sincere thanks also to Keith A. Seifert, James Lendemer and

Scott Redhead for helpful pre-submission reviews. Lorelei L. Norvell and Shaun R.

Pennycook also contributed significantly to improving the text.

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MY COTAXON

http://dx.doi.org/10.5248/119.509

Volume 119, pp. 509-511 January-March 2012

NOMENCLATURAL NOVELTIES AND TYPIFICATIONS

PROPOSED IN MYCOTAXON 119

Abrachium Baseia & T.S. Cabral, p. 424

Abrachium floriforme (Baseia & Calonge) Baseia & T.S. Cabral, p. 424

Acarospora gallica H. Magn. 1929, neotypified, p. 374

Alveariospora Meir. Silva, R.F. Castafieda, O.L. Pereira & R.W. Barreto, p. 110

Alveariospora distoseptata Meir. Silva, R.E. Castafieda, O.L. Pereira &

R.W. Barreto, p. 110

Anaseptoidium R.E. Castaneda, Heredia & R.M. Arias, p. 141

Anaseptoidium mycophilum R.F. Castafieda, Heredia & R.M. Arias, p. 142

Antrodia madronae Vlasak & Ryvarden, p. 220

Ceriporiopsis pseudoplacenta Vlasak & Ryvarden, p. 222

Coltricia indica Baltazar, p. 386

= Coltricia pusilla J.R. Sharma & J.E. Wright 1992 [‘1989°], nom. illeg.

= Coltricia parvula J.R. Sharma & J.E. Wright 1997, nom. illeg.

Corynesporella obclavata L.G. Ma & X.G. Zhang, p. 83

Cylindrosympodium sosae R.F. Castafieda, Heredia & R.M. Arias, p. 145

Deconica deconicoides (E. Horak, Guzman & Desjardin) Guzman, p. 67

Diorygma agumbense B.O. Sharma & Khadilkar, p. 2

Diorygma dandeliense B.O. Sharma & Khadilkar, p. 3

Diorygma karnatakense B.O. Sharma & Khadilkar, p. 4

Diorygma rupicola B.O. Sharma & Makhija, p. 5

Endophragmiella gardeniae Jian Ma & X.G. Zhang, p. 104

Endophragmiella machili Jian Ma & X.G. Zhang, p. 105

Fulvifomes cambodiensis L.W. Zhou & W.M. Zhang, p. 176

Fuscoporia palomari Vlasak & Ryvarden, p. 218

Gymnopus inusitatus var. cystidiatus Antonin, Finy & TomSovsky, p. 294

Helvella fusca Gillet (1879), lectotypified & epitypified, p. 434

Herpothallon weii Y.L. Cheng & H.Y. Wang, p. 440

510... MYCOTAXON 119

Heteroconium annesleae S.C. Ren & X.G. Zhang, p. 363

Heteroconium neolitseae S.C. Ren & X.G. Zhang, p. 363

Hyphoderma bicystidiatum Priyanka & Dhingra, p. 255

Hyphoderma subglobosum Priyanka & Dhingra, p. 257

Hysterostomella connari Inacio, Pereira-Carvalho, E.S.C. Souza, H.B. Sales &

Dianese, p. 308

Tjuhya oenanthicola Lechat & Hairaud, p. 249

Intraornatosporaceae B.T. Goto & Oehl, p. 121

Intraornatospora B.T. Goto, Oehl & G.A. Silva, p. 122

Intraornatospora intraornata (B.T. Goto & Oehl) B.T. Goto, Oehl &

G.A. Silva, p. 122

Lactifluus subg. Gerardii (A.H. Sm. & Hesler) Stubbe, p. 484

Lactifluus atrovelutinus (J.Z. Ying) X.H. Wang, p. 484

Lactifluus bicolor (Massee) Verbeken, p. 484

Lactifluus conchatulus (Stubbe & H.T. Le) Stubbe, p. 484

Lactifluus coniculus (Stubbe & Verbeken) Verbeken, p. 484

Lactifluus genevievae (Stubbe & Verbeken) Stubbe, p. 484

Lactifluus hora (Stubbe & Verbeken) Stubbe, p. 484

Lactifluus leae (Stubbe & Verbeken) Verbeken, p. 484

Lactifluus leonardii (Stubbe & Verbeken) Stubbe, p. 484

Lactifluus limbatus (Stubbe & Verbeken) Stubbe, p. 485

Lactifluus ochrogalactus (Hashiya) X.H. Wang, p. 485

Lactifluus petersenii (Hesler & A.H. Sm.) Stubbe, p. 485

Lactifluus reticulatovenosus (Verbeken & E. Horak) Verbeken, p. 485

Lactifluus sepiaceus (McNabb) Stubbe, p. 485

Lactifluus subgerardii (Hesler & A.H. Sm.) Stubbe, p. 485

Lactifluus uyedae (Singer) Verbeken, p. 485

Lactifluus venosus (Verbeken & E. Horak) Verbeken, p. 485

Lactifluus wirrabara (Grgur.) Stubbe, p. 485

Lecanora expallens Ach. (1810), lectotypified, p. 198

Lecanora expallens var. conizaea Ach. (1810), lectotypified, p. 198

Leptogium weii H.J. Liu & S. Guan, p. 413

Marasmiellus milicae Me&i¢, Tkaléec & Antonin, p. 234

Mycena moconensis Niveiro, Alberté & Desjardin, p. 168

Paradentiscutata B.T. Goto, Oehl & G.A. Silva, p. 122

Paradentiscutata bahiana Oehl, Magna, B.T. Goto & G.A. Silva, p. 123

Paradentiscutata maritima B.T. Goto, D.K. Silva, Oehl & G.A. Silva, p. 126

Passalora macfadyenae Meir. Silva, O.L. Pereira & R.W. Barreto, p. 187

NOMENCLATURAL NOVELTIES & TYPIFICATIONS ...

Penicillium guanacastense K.G. Rivera, Urb & Seifert, p. 324

Penicillium mallochii K.G. Rivera, Urb & Seifert, p. 322

Phlebopus spongiosus Pham & Har. Takah., p. 28

Pleurophragmium indicum M.A. D’Souza & Bhat, p. 477

Prospodium macfadyenae Meir. Silva, O.L. Pereira & R.W. Barreto, p. 191

Pseudobaeospora deckeri C.F. Schwarz, p. 466

Pseudocercosporella stellariicola M.J. Park, J.H. Park & H.D. Shin, p. 270

Psilocybe thaiaerugineomaculans Guzman, Karunarathna & Ram.-Guill., p. 69

Psilocybe thaicordispora Guzman, Ram.-Guill. & Karunarathna, p. 72

Psilocybe thaiduplicatocystidiata Guzman, Karunarathna & Ram.-Guill., p. 74

Psilocybe thaizapoteca Guzman, Karunarathna & Ram.-Guill., p. 77

Radulomycetopsis Dhingra, Priyanka & J. Kaur, p. 133

Radulomycetopsis cystidiata Dhingra, Priyanka & J. Kaur, p. 135

Remototrachyna aguirrei (Sipman, Elix & T.H. Nash) Flakus, Kukwa &

Sipman, p. 161

Remototrachyna consimilis (Vain.) Flakus, Kukwa & Sipman, p. 161

Remototrachyna singularis (Hale) Flakus, Kukwa & Sipman, p. 162

Remototrachyna sipmaniana Kukwa & Flakus, p. 162

Skeletocutis bambusicola L.W. Zhou & W.M. Qin, p. 346

Skeletocutis subodora Vlasak & Ryvarden, p. 228

Solicorynespora cryptocaryae Jian Ma & X.G. Zhang, p. 98

Solicorynespora ligustri Jian Ma & X.G. Zhang, p. 96

Solicorynespora machili Jian Ma & X.G. Zhang, p. 97

Sporidesmium antidesmatis Jian Ma & X.G. Zhang, p. 19

Sporidesmium liquidambaris Jian Ma & X.G. Zhang, p. 18

Sporidesmium machili Jian Ma & X.G. Zhang, p. 21

Terriera petrakii C.L. Hou, p. 332

Terriera rotundata C.L. Hou, p. 330

Tuber microspermum L. Fan & J.Z. Cao, p. 392

Tuber microspiculatum L. Fan & Yu Li, p. 392

Volvariella sathei Senthil., Rahul Sharma & S.K. Singh, p. 470

Xylaria atroglobosa H.X. Ma, Lar.N. Vassiljeva & Yu Li, p. 382

Zelodactylaria A.C. Cruz, Gusmao & R.F. Castafieda, p. 242

Zelodactylaria veticillata A.C. Cruz, Gusmao & R.F. Castafieda, p. 242

Sat