IV ... MYCOTAXON 126

MYCOTAXON

VOLUME ONE HUNDRED TWENTY-SIX — TABLE OF CONTENTS

COVER SECTION

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RREVIC WERE: brs 25h at Sean Fe coe BSE S Wis bole Noite Ee, eel a teria tne vii

SUBMISSION PYOCCUUTCs: 2 84.5 25 hokey taht FA Doe ATS RA Ee gs aoe ae on Viii

PROUT TOE so is a a aes ore ey Me oa gh Men oat We Ratt aie ee alg ix

RESEARCH ARTICLES

Biogeography and taxonomy of pyrenomycetous fungi 3.

Three new species from the area around the Sea of Japan

LARISSA N. VASILYEVA, HAI-x1A MA & STEVEN L. STEPHENSON

Hyphomycetes: Linodochium sinense sp. nov. and new records

from fallen Sycopsis sinensis leaves in China

DeE-WE! LI, JINGYUAN CHEN & YIXUN WANG

Colletotrichum fructicola, first record of bitter rot of apple in China

DAN-DAN Fu, WEI WANG, RUI-FENG QIN,

RONG ZHANG, GUANG-YU SUN & MARK L. GLEASON

Phyllobaeis crustacea sp. nov. from China

SHUNAN CAo, XINLI WEI, QIMING ZHOU & JIANGCHUN WEI

Nipponoparmelia perplicata sp. nov. (Parmeliaceae, Ascomycota) from

eastern Asia SERGIJ YA. KONDRATYUK, SVETLANA I. TSCHABANENKO,

JoHN A. ELIx, SOON OK OH, ARNE THELL & JAE-SEOUN HuR

Buelliella, Codonmyces, and Polycoccum species new to Turkey

KENAN YAZICI & JAVIER ETAYO

Passalora lepistemonis sp. nov. from China

LEI X1A, YING-LAN GUO & Yu LI

Two new species of Spadicoides from southern China

JI-WEN X1A, L1-Guo Ma, JIAN MA & X1tU-GUO ZHANG

Entocybe haastii from Watagans National Park,

New South Wales, Australia SARAH E. BERGEMANN,

Davip L. LARGENT, & SANDRA E. ABELL-DAVIS

Craterium corniculatum sp. nov. from northwestern China

Bo ZHANG & YU LI

Embryonispora, a new genus of hyphomycetes from China

KAI-NING ZHAO, GUO-HUA YIN, GUO-ZHU ZHAO & AI-XIN CAO

is)

igs

OCTOBER-DECEMBER 2013 ... V

The genus Neopaxillus (Crepidotaceae)

Roy WATLING & M. CATHERINE AIME 83

New records of Aspicilia from China

SHU-XIA LI, XING-RAN KOU & QIANG REN 91

Ambomucor gen. & spp. nov. from China

RU-YONG ZHENG & XIAO-YONG LIU 97

Two new species of Rhytismataceae on fagaceous trees

from Anhui, China Li CHEN, D.W. MINTER,

SHI-JUAN WANG & YING-REN LIN 109

Three newly recorded species of Parasympodiella and Chalara

from China X1a0-X1A Li, L1-Guo Ma,

JI-WEN XIA & XIU-GUO ZHANG 121

Tuber bomiense, a new truffle species from Tibet, China

Kat-MEI Su, WEI-PING XIONG, YUN WANG,

SHU-HonG LI, RONG XIE & DANZENG BAIMA 127

Lepiota brunneoincarnata and L. subincarnata:

distribution and phylogeny A. RAZAQ, E.C. VELLINGA,

S. ILyAs & A.N. KHALID 133

New combinations in Trichoderma (Hypocreaceae, Hypocreales)

WALTER M. JAKLITSCH & HERMANN VOGLMAYR 143

Lactarius taxa (Basidiomycota, Russulales) by Zdenék Schaefer -

types and other collections in PRM

JAN HOLEC, KAMILA PESICOVA & LENKA EDROVA 157

Phytophthora virginiana sp. nov., a high-temperature tolerant species

from irrigation water in Virginia XIAO YANG & CHUANXUE HONG 167

New records of Puccinia on Poaceae from Khyber Pakhtunkhwa,

Pakistan A. IsHAQ, N.S. AFSHAN & A.N. KHALID 177

The olive goblet: Peziza oliviae, a new cup fungus

growing underwater in Oregon JONATHAN L. FRANK 183

One hundred fourteen years of Pluteus in Brazil:

collections studied by Hennings and Rick

NELSON MENOLLI JR. & MARINA CAPELARI 191

A new species of Bahusutrabeeja from Guangxi, China

XIAO-XIA LI, JI-WEN X1A, L1-Guo Ma,

RAFAEL F. CASTANEDA-RUIZ & XIU-GUO ZHANG 227

Flavophlebia sphaerospora, a new corticoid species from India

MANINDER Kaur, AVNEET P. SINGH & G.S. DHINGRA 231

Peniophora hallenbergii sp. nov. from India SAMITA & G.S. DHINGRA 235

BOOK REVIEWS AND NOTICES ELsE C. VELLINGA (Editor) 239

VI ... MYCOTAXON 126

REGIONAL MYCOBIOTAS NEW TO THE MYCOTAXON WEBSITE

The Cortinariaceae of Argentinas Nothofagus forests

G.M. ROMANO & N.E. LECHNER 247

Lichenized and lichenicolous fungi from the Pititises Archipelago (Eivissa

and Formentera Islands and Islets), Balearic Islands, Spain

VIOLETA ATIENZA, ELENA ARAUJO,

MarIA DEL ROSARIO ARROYO, ANA ROSA BURGAZ, GEMMA FIGUERAS,

ANTONIO GOMEZ-BOLEA, NESTOR L. HLADUN, XAVIER LLIMONA,

ESTEVE Liop, Maria EUGENIA LOPEZ DE SILANES, BERNARDA MARCOS,

ISRAEL PEREZ- VARGAS, RAQUEL PINO-BODAS, ESTELA SERINA

& VicTOR J. Rico 247

Lichen flora of the Ilam Province, South West Iran TAHEREH VALADBEIGI 248

NOMENCLATURAL NOVELTIES AND TYPIFICATIONS

PROPOSED IN VOLUME 126 249

ERRATUM FROM MYCOTAXON 125

p.196, line 27 FOR: Wuiding — READ: Wuding —

OcTOBER-DECEMBER 2013 ... VII

REVIEWERS — VOLUME ONE HUNDRED TWENTY-SIX

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.

VLADIMIR ANTONIN

A. APTROOT

TrmoTHy J. BARONI

GERALD L. BENNY

WOLFGANG VON BRACKEL

VICENT CALATAYUD

RAFAEL FE, CASTANEDA RUIZ

GREGORIO DELGADO

PRADEEP K. DIVAKAR

KEITH EGGER

Li Fan

MANNON GALLEGLY

WALTER GAMS

ZAI-WEI GE

IAN R. HALL

NILS HALLENBERG

TERRY W. HENKEL

Hon H. Ho

PETER JOHNSTON

INGVAR KARNEFELT

BRYCE KENDRICK

PAUL M. KirK

KERRY KNUDSEN

FRANCISCO KUHAR

T.K. ARUN KUMAR

DeE-WE!I LI

JUN-FENG LIANG

D. JEAN LODGE

Guo-ZHONG LU

ROBERT LUCKING

HELMUT MAYRHOFER

Eric H.C. McKENZIE

ANDREW M. MINNIS

D.W. MINTER

A.R. NIAZI

LORELEI L. NORVELL

DavID ORLOVICH

OMAR Paino PERDOMO

SHAUN R. PENNYCOOK

SERGIO PEREZ-ORTEGA

DONALD H. PFISTER

Gary J. SAMUELS

B.M. SHARMA

CAROL A. SHEARER

H.J.M. SIPMAN

AVE SUIJA

ELsE C. VELLINGA

JOHN C. VILLELLA

ANNEMIEKE VERBEKEN

FELIPE WARTCHOW

A.J.S. WHALLEY

MING YE

M. ZHANG

ZHONGYI ZHANG

WEN- YING ZHUANG

PUBLICATION DATE FOR VOLUME ONE HUNDRED TWENTY-FIVE

MYCOTAXON for JuLy-SEPTEMBER, VOLUME 125 (I-x1I + 1-306)

was issued on November 20, 2013

vu ... MYCOTAXON 126

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OcCTOBER-DECEMBER 2013 ... IX

FROM THE EDITOR-IN-CHIEF

MycoTaxoNn 126, which owes its slimness to a temporary presubmission review

bottleneck, presents 27 excellent papers by 87 authors from 21 countries and revised

by 50 expert reviewers. Within its pages are two new genera (Ambomucor and

Embryonispora from China) plus 22 other taxa new to science representing Ambomucor,

Bahusutrabeeja, Craterium, Embryonispora, Hypoxylon, Linodochium, Lophodermium,

Passolora, Phyllobaeis, Spadicoides, Terriera, and Tuber from China; Annulohypoxylon

and Nectria from Russia; Flavophlebia and Peniophora from India; Nipponoparmelia

from Russia and South Korea, and from the United States a new Phytophthora from

irrigation water in Virginia and a new underwater (!) Peziza from Oregon.

In addition to range extensions and new hosts for previously named taxa, we also

offer one new Entocybe combination from Australia, 46 new (and eagerly awaited)

combinations in Trichoderma, a reevaluation of Neopaxillus, and two excellent

taxonomic revisions —one on Zdenék Schaefer's Lactarius taxa in Prague, the other

a comprehensive treatment of Pluteus collections previously studied by Hennings and

Rick in Brazil. We close with some excellent book reviews!

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Warm regards,

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9 March 2014

MYCOTAXON

http://dx.doi.org/10.5248/126.1

Volume 126, pp. 1-14 October-December 2013

Biogeography and taxonomy of pyrenomycetous fungi 3.

The area around the Sea of Japan

LARISSA N. VASILYEVA’*, HAI-x1IA MA*? & STEVEN L. STEPHENSON‘

‘Institute of Biology & Soil Science, Far East Branch of the Russian Academy of Sciences,

Vladivostok 690022, Russia

?Institute of Tropical Bioscience and Biotechnology, Chinese Academy of

Tropical Agricultural Sciences, Haikou 571101, China

*Institute of Mycology, Jilin Agricultural University, Changchun 130118, China

‘Department of Biological Sciences, University of Arkansas, Fayetteville, AR 72701, USA

*CORRESPONDENCE TO: vasilyeva@biosoil.ru

AsstRACT — The peculiar species composition of the assemblage of pyrenomycetous fungi

known from the area around the Sea of Japan is discussed. Three new species—Annulohypoxylon

orientale, Hypoxylon cyanescens, and Nectria araliae—are described and illustrated.

KEY worps — Ascomycota, northeastern Asia, ecological distribution

Introduction

One of the most interesting regions of northeastern Asia is the area around

the Sea of Japan. Many pyrenomycetous fungi are restricted to this region

of the world, and a number of economically important pathogenic fungi are

known only from northeastern China, Japan, Korea and southeastern Russia.

Unfortunately, often many of these fungi have been confused with other species,

so phytopathologists cannot recognize them and thus develop the appropriate

control measures.

At the 2011 Asian Mycological Congress held in Incheon (Korea), Dr.

Tsuyoshi Hosoya—during his talk on the reassessment of Hyaloscyphaceae

(Helotiales, Leotiomycetes) based on molecular phylogenetic analysis—repeated

Dr. Richard Korf’s statement that “for the discosystematist, Japan is a relatively

unexplored paradise” and that the region is “rich in apparently endemic forms,

an understanding of which is essential for phylogenetic and phytogeographic

studies” (Korf 1958: 7). However, it seems that not only Japan but rather the

entire area around the Sea of Japan is the real ‘paradise’ for many groups of

fungi.

2 ... Vasilyeva, Ma & Stephenson

Materials & methods

The specimens reported herein are deposited in the Herbarium of Institute of Biology

and Soil Science, Far East Branch of the Russian Academy of Sciences (VLA) and the

Herbarium of Cryptogams of the Kunming Institute of Botany, Chinese Academy of

Sciences (HKAS). Photographs of ascomata were obtained using a Nikon D40x digital

camera and $70 Canon camera. The photographs of ascospores were taken by using

a VHX-600E microscope of the Keyence Corporation. External stromatal colors were

recorded and coded according to Rayner (1970).

Some pathogenic and nonpathogenic pyrenomycetous fungi

restricted to the area around the Sea of Japan

One pathogen restricted to the Sea of Japan area is Venturia nashicola

S. Tanaka & S. Yamam., the scab fungus of Japanese and Chinese pears. Soon

after Tanaka & Yamamoto (1964) described this species, its name was reduced

to a synonym of V. pirina Aderh., which causes a similar disease in European

pears (Sivanesan 1977). Although Khokhryakova (1978) had already discussed

that V. nashicola has shorter ascospores and conidia than V. pirina, her studies

and special references to their biogeographical differences (Khokhryakova

1978, 1980) were initially ignored, and only recently have her findings been

supported by the data obtained by other researchers (Ishii & Yanase 2000). The

differences in ascospore size between V. nashicola and V. pirina (on Pyrus) are

exactly the same as between V. chlorospora (Ces.) P. Karst. and V. subcutanea

Dearn. (on Salix) or V. canadensis M.E. Barr and V. rumicis (Desm.) G. Winter

(on Rumex), and if the latter species pairs are accepted as independent (Barr

1968), there seems little reason to unite V. nashicola and V. pirina.

Another pathogen restricted to the Sea of Japan area is Polystigma ussuriense

(Jacz. & Natalyina) Prots., first described from the Russian Far East (Natalyina

1931) but not acknowledged until after almost a half century by Khokhryakova

(1978, 1980). This species was re-examined even later (Cannon 1996) and

accepted as only a subspecies of P. rubrum (Pers.) DC. However, even at the

level of subspecies the Sea of Japan area has its own taxon. The ascospores and

conidia of P ussuriense are longer than those of P rubrum — the same characters

separating Venturia nashicola and V. pirina. Although the same characters

have unequal value in different (especially remote) genera, size differences in

ascospores and conidia almost universally discriminate species in numerous

pyrenomycetous groups (cf. Rappaz 1987; Rossman et al. 1999; Gryzenhout

et al. 2009; Jaklitsch 2011). Consequently, P ussuriense might be considered as

an independent species occurring in northeastern Asia and probably replaces

P. rubrum in this area, although the P. rubrum has been reported from China,

Japan, and Korea (Cannon 1996; Katumoto 2010).

Another example, Valsa mali Miyabe & G. Yamada, is a very aggressive

pathogen that causes canker of apple trees in various countries surrounding

the Sea of Japan. Described more than a century ago (Ideta 1911), V. mali

Pyrenomycetes of the Sea of Japan area ... 3

was confused with other species during the past century, most often with

V. ceratosperma (Tode) Maire (e.g., Kobayashi 1970; Otani 1995), although the

two species can be easily distinguished superficially (Fic. 1). The arrangement

of ostioles in V. mali as a ring around the margin of an ectostromatic disc is

characteristic of Valsa sect. Circinatae (Nitschke 1867; Urban 1957) or sect.

Valsa (Spielman 1985). The character of an ectostromatic disc filled with closely

packed ostioles places V. ceratosperma in section Monostichae.

The independent status of V. mali began to be discussed only relatively

recently (Vasilyeva & Kim 2000), although its anamorph, Cytospora orientis-

extremi Gvrit., had been described (with reference to V. mali as the teleomorph)

as a separate species even earlier (Gvritishvili 1973). However, the first

molecular study to consider V. mali (Adams et al. 2002) suggested that it be

combined with Leucostoma persoonii (Nitschke) Héhn., since the two species

clustered together on a phylogenetic tree. Although they look completely

different superficially (Fic. 1), the clustering is actually not surprising, given

that these two species share a number of similarities (e.g., the kind and size

of ascospores, stromatal shape, Cytospora anamorphs, restriction to rosaceous

hosts). The problem is that those same similarities comprise a repetitive internal

polymorphism within Valsa and Leucostoma and outweigh the few differences

that exist between them. When molecular trees are constructed based on

phenetic principles, thus relying upon the quantity of shared characters without

discriminating their level in the taxonomic hierarchy, such mistakes in the

unification of species from different genera are potentially common (Vasilyeva

& Stephenson 2010a). A more recent sequence analysis by Wang et al. (2011)

Fic. 1. Stromata. A. Valsa mali (VLA P-1981); B. Valsa ceratosperma (VLA P-1845); C. Leucostoma

persoonii (VLA P-230). Scale bars: A, C = 0.5 mm; B = 0.3 mm.

4 ... Vasilyeva, Ma & Stephenson

Fic. 2. Known localities of: A. Biscogniauxia maritima; B. Biscogniauxia mandshurica;

C. Cryptosphaeria venusta; D. Cryptosphaeria exornata. (The map, taken from http://www.fas.org,

has been modified to show our data).

supports V. mali as independent, and its possible distribution in northeastern

Asia might be shared with a number of non-pathogenic pyrenomycetous fungi.

Some species described originally from the Russian Far East (Vasilyeva 1998)

— Biscogniauxia mandshurica Lar.N. Vassiljeva, B. maritima Lar.N. Vassiljeva,

Cryptosphaeria exornata Lar.N. Vassiljeva, C. venusta Lar.N. Vassiljeva — were

later found in China, Korea or Japan; since their English descriptions and

illustrations of stromata have been provided elsewhere (Vasilyeva et al. 2009),

only their known localities are presented here (Fic. 2). Diaporthella corylina

Lar.N. Vassiljeva is known from only two localities in China and Russia, and

Leucodiaporthe acerina M.E. Barr & Lar.N. Vassiljeva from only two localities

in Korea and Russia (Vasilyeva et al. 2007). Podostroma giganteum S. Imai,

described from Japan (Imai 1932), was reported much later for the Russian Far

East.

Pyrenomycetes of the Sea of Japan area... 5

Among the species reported previously from northeastern Asia (Tsuneda

1982; Tsuneda & Arita 1984; Vasilyeva 1998) in need of taxonomic

reconsideration is Hypoxylon truncatum (Schwein.) J.H. Mill. As can be seen

from Fic. 3, the stromata of ‘H. truncatum’ from the Russian Far East surely

differ from those observed for H. truncatum specimens from the eastern United

States; hence we assign the Russian specimens to Annulohypoxylon orientale, a

new species described herein. In addition, Vasilyeva (1998) reported Nectria

coryli Fuckel from the southern Russian Far East but emphasized that it had

never been found on either of the two Corylus species (C. heterophylla Fisch.

ex Trautv., C. mandshurica Maxim.) that are widely distributed in the region.

Instead, the fungus was restricted to the Araliaceae—mostly to Aralia elata

(Miq.) Seem. and Eleutherococcus senticosus (Rupr. & Maxim.) Maxim.—and

was characterized by the presence of black ascomata similar to those of Nectria

austroamericana (Speg.) Cooke (Samuels et al. 2006), a marked contrast to the

red ascomata in N. coryli. Ascospores of Nectria species from the Sea of Japan

area appear even smaller than those characteristic of N. coryli. Therefore, on

biogeographical, ecological, and morphological grounds, the former warrants

being described as the new species N. araliae.

The specimens of the third new species described in this paper were repeatedly

found in the southern Russian Far East, mostly on Ligustrina amurensis (Rupt.)

Rupr. (Oleaceae), which is distributed throughout northeastern China, Korea,

and the Russian Far East. However, the new species was not described until

the same fungus was collected in Jilin Province (northeastern China), as

discussed in the Ph.D. dissertation by Haixia Ma (Ma 2011), where it was listed

as “Hypoxylon aeneo-viridis’; herein it is formally named as H. cyanescens.

Taxonomy

Annulohypoxylon orientale Lar.N. Vassiljeva & S.L. Stephenson, sp.nov. Fic. 3B

MycoBank MB 802.460

A Annulohypoxylone truncato in disco ostiolato plano ejusdem typi ac A. bovei differt.

Type: Russia: Primorsky Territory, Khasan District, Trinity Bay, on dead branches of

Quercus mongolica Fisch. ex Lebed. (Fagaceae), 3 Nov. 1995, L. Vasilyeva (Holotype

VLA P-382).

Erymo.ocy: The epithet refers to the eastern part of Asia where this species was

collected.

STROMATA Solitary to densely aggregated, semi-globose, 2-4 mm diam., 1.3-1.5

mm high, with perithecial mounds %-% exposed, surface initially brown with

shining-black ostiolar caps, then becoming blackened, with KOH-extractable

pigments olivaceous (48); the tissue below the perithecial layer blackish.

Perithecia spherical, 0.25-0.35 mm diam., ostioles finely papillate, encircled

with a flat, bovei-type disc 0.2-0.25 mm diam. Asci in the spore-bearing

6 ... Vasilyeva, Ma & Stephenson

Fic. 3. Stromata. A. Annulohypoxylon truncatum from Texas, USA (VLA P-2446);

B. Annulohypoxylon orientale (VLA P-381). Scale bars: A = 1.2 mm; B = 2.7 mm. (Photo E.M.

Bulakh).

portion 70-80 x 4.5-5.5 um, stipitate, stipes 30-50 um long, with an apical

ring bluing in Melzer’s iodine reagent, discoid, about 1 um broad. Ascospores

brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, 8-10

x 4-4.5(-5) um, with a straight germ slit shorter than the length of the spore;

perispore dehiscent in 10% KOH, smooth.

ADDITIONAL SPECIMENS EXAMINED: RUSSIA: PRiMoRSKY TERRITORY, Khasan

District, Ryazanovka vicinity, on Quercus mongolica, 30 Jun 1982, M. Nazarova (VLA

P-381); Trinity Bay, on Quercus mongolica, 4 Aug 2006, L. Vasilyeva (VLA P-1803).

KOREA: vicinity of Seoul, Suwon, on Quercus sp., 30 May 1999, L. Vasilyeva (VLA

P-1649)

ComMMENTS—In the Russian Far East, this species has been found only in the

Khasan District, at the very southern extreme of the Primorsky Territory. It

differs from A. truncatum (Schwein.) Y.M. Ju et al. in having smaller ostiolar

discs that are of the bovei-type instead of the truncatum-type. The general

appearance of the stromata is characterized by the presence of the shining-

black and convex ostiolar caps, which persist for a long time but finally fall

off to expose the flat ostiolar discs. The image of stromata of a specimen from

Japan (Tsuneda & Arita 1984: Fig. 1) corresponds to that noted for specimens

from southeastern Russia.

Pyrenomycetes of the Sea of Japan area ... 7

Fig. 4. Hypoxylon cyanescens: A-B. Stromata; C. Stromatal surface; D. Ascospores. Scale bars:

A = 3.6 mm, B = 2.7 mm, C = 1.4 mm, D = 9.4 um.

Hypoxylon cyanescens H.X. Ma, Lar.N. Vassiljeva & Yu Li, sp. nov. Fic. 4

MycoBank MB 802459

A Hypoxylone aeruginoso in ascosporis majoribus et a H. aeruginoso var. macrosporo in

granulis in KOH dissolutis differt.

Type: China: Jilin Province, Changbai Mountain, alt. 1300 m, on rotten wood, 7 Sept

1998, Pei-gui Liu (Holotype HKAS32650).

Erymo.oey: The epithet refers to the color of the stroma.

STROMATA pulvinate or effused-pulvinate, flat or with conspicuous perithecial

mounds 2-8 mm diam. and 0.5-1 mm thick; surface brown to dark vinaceous

at the sterile margins, dark cyan blue, isabelline or gray olivaceous over the

ostiolar portion, later becoming darkened, with KOH-extractable pigments bay

(6) to livid purple (81). Perithecia 0.1-0.3 mm, ostioles finely papillate. Asci

not observed. Ascospores brown, unicellular, ellipsoid-inequilateral or almost

equilateral, with narrowly or broadly rounded ends, 11.5-13.5 x 5-6 um, with

a straight germ slit extending the length of the spore; perispore indehiscent in

10% KOH, smooth.

ADDITIONAL SPECIMENS EXAMINED: RUSSIA: PRimoRSKY TERRITORY, Lazovsky State

Nature Reserve, on Ligustrina amurensis (Oleaceae), 2 Aug 1986, L. Vasilyeva (VLA

8 ... Vasilyeva, Ma & Stephenson

P-2638); AMUR REGION, in the vicinity of Kundur, 24 Aug 1992, L. Vasilyeva (VLA

P-2639). KOREA: Gangwon province, Pyeongchang county, Mt. Odaesan, 20 Sep 2006,

L. Vasilyeva (VLA P-2746).

CoMMENTS—Hypoxylon aeruginosum J.H. Mill. and its large-spored variety

H. aeruginosum var. macrosporum J.D. Rogers are the only previously known

taxa with a dark cyan blue stromatal surface. Both varieties of H. aeruginosum

are known from the western hemisphere (Guyana, Mexico, and Louisiana in

the United States). Hypoxylon cyanescens is similar to the large-spored variety

of H. aeruginosum in ascospore size but differs in the presence of bay to livid

purple KOH-extractable pigments, whereas the latter variety has stromata

apparently without KOH-extractable pigments.

Nectria araliae Lar.N. Vassiljeva & S.L. Stephenson, sp. nov.

MycoBaAnk MB 802461

A Nectria coryli in ascosporis minoribus et ascomatis nigrescentibus differt.

Type: Russia: Primorsky Territory, Khasan District, Kedrovaya Pad Nature Reserve, on

Aralia elata (Araliaceae), 25 Oct 1987, L. Vasilyeva (Holotype VLA P-1383).

Erymo.oey: The epithet refers to the host genus of the holotype specimen.

ASCOMATA crowded in clusters, superficial or slightly embedded on a stromatic

basis, dark-grey or black, globose to turbinate, becoming collapsed when dry,

smooth, 200-250 um diam., with small papillate ostioles. Asci cylindrical,

70-80 x 4-4.5 um. Ascospores biseriate, two-celled, about 8-10 x 3-4 um, with

small setose appendages, usually budding in the asci.

ADDITIONAL SPECIMENS EXAMINED: RUSSIA, PRimoRSKY TERRITORY, Ussuriysk

District, Mountain-Taiga Station, on Aralia elata, 23 Sep 1984, M. Gvritischvili

(VLA P-1993); Sikhote-Alin Nature Biosphere Reserve, on A. elata, 3 Aug 1985,

L. Vasilyeva (VLA P-1381); KurILE IsLANDs, Kunashir Island, on A. elata, 31 Jul

1987, L. Vasilyeva (VLA P-1386); KHABAROVSK TERRITORY, Komsomolsk Nature

Reserve, on Eleutherococcus senticosus, 2 Jul 1986, L. Vasilyeva (VLA P-2139); JEwIsH

AUTONOMOUS REGION, Bastak Nature Reserve, on E. senticosus, 19 Aug 2004,

L. Vasilyeva (VLA P-2124).

ComMENTS— The similarity of Nectria araliae with N. coryli is impressive, and

the ascospores have setose appendages and are usually budding in both species.

Only recently, the ecological preferences of the north-Asian specimens and

their distribution peculiarities were taken into consideration. Superficially, the

black ascomata of N. araliae are similar to those of Nectria austroamericana

(Samuels et al. 2006: photo on p. 120), but the latter seems to be restricted

to members of the Fabaceae (Acacia and Gleditsia), has ascospores that are

irregularly septate, with 3-6-transverse and one longitudinal septa, and has

been reported for North and South America.

Pyrenomycetes of the Sea of Japan area ... 9

Discussion

The area considered in this paper is known as “the Eastern Asiatic Region

(Oriasiaticum, Sino-Japanese Region, East Asian Region, Temperate Eastern

Region)” and has been described as “the richest floristic region within the

Holarctic Kingdom situated in temperate East Asia.” Moreover, “it has been

recognized as a natural floristic area since the publication in 1872 of August

Grisebach’ss volume ‘Die Vegetation der Erde’ and was later delineated by

such geobotanists as Ludwig Diels, Adolf Engler (as the Temperate Eastern

region), Ronald Good (as the Sino-Japanese Region), and Armen Takhtajan”

(http://en.wikipedia.org).

This special center of biodiversity is said to comprise “the southern part

of the Russian Far East, southern part of Sakhalin, Manchuria, Korea, Japan,

Taiwan, the relatively humid eastern part of mainland China extending from

Manchuria and the coastline to the Eastern Himalaya and Kali Gandaki Valley

in Nepal, including Sikkim, northern Burma (Myanmar) and northernmost

Vietnam (parts of Tonkin)” (http://en.wikipedia.org). Although knowledge

of the mycobiota in many of the countries in southern Asia (such as Burma

or Vietnam) is rather poor, the scattered data reported in the rather limited

literature and our preliminary observations indicate that the fungal biodiversity

in countries (e.g., China, Japan, Korea, and southeastern Russia) located

around the Sea of Japan differs from that of more southern portions of this

world region. Moreover, it also should be noted that some current studies of

vegetation also exclude the south of China, Burma, and Vietnam from the Sino-

Japanese Floristic Region (Yih 2012: map at p. 16). It seems that the apparently

unique area in question is primarily limited by the distribution in northeastern

Asia of what are referred to as ‘cool-temperate rainforests’ (Krestov et al. 2011;

Nakamura et al. 2011), whereas the excluded countries are characterized by

different types of climate and vegetation.

Many pyrenomycetes have been collected exclusively in the Sea of Japan area

and assigned to new species or genera (Vasilyeva 1990, 1993, 2001, 2007, 2010;

Ju et al. 1999, 2009; Stadler et al. 2005; Vasilyeva & Melnik 2006; Vasilyeva &

Stadler 2008; Vasilyeva et al. 2007, 2010; Hirooka et al. 2011). The main purpose

of the Vasilyeva & Stephenson (2010b, 2011) series (for which this is the third

contribution) is to emphasize that by neglecting biogeographical patterns,

mycologists often have reached the wrong conclusions regarding application

of species names to fungi in northeastern Asia. For example, Cryptosphaeria

eunomioides (G.H. Otth) Hohn., Diatrype disciformis (Hoftm.) Fr., Diatrypella

quercina (Pers.) Cooke, Hypoxylon fragiforme (Scop.) J. Kickx f., and Rosellinia

aquila (Fr.) Ces. & De Not. have all been reported from the Russian Far East

10 ... Vasilyeva, Ma & Stephenson

(e.g., Koval 1972) but actually do not occur there, and any of these names

applied to Chinese fungi (Teng 1996) should be checked.

It is possible that species restricted to the Sea of Japan area had a wider

distribution prior to the Glacial Age but were pushed by glaciers to the edge

of the Pacific Ocean where the Sea of Japan formed after separation of the

island arc of Japan from the Asian mainland about 20 million years ago. It is

possible that this area became a kind of ‘trap’ for fungi that depend upon air

currents for spore dispersal, whereas the air currents themselves often depend

upon sea currents. If one looks at the map of sea currents around the Japanese

Archipelago (http://valeriecourreges.blogspot.com), one can see that the main

water circulation pattern in the Sea of Japan is created by two main currents,

with the Tsushima Current being warm and the Liman Current being cold.

In fact, these two currents create a closed system of water as well as the air

currents responsible for transferring fungal spores. The partial exit from

this closed system toward Sakhalin Island and the southern Kurile Islands is

provided by the Soya Current. This might explain why some fungal species

(e.g., Loranitschkia viticola Lar.N. Vassiljeva, Nectria araliae) are known not

only from the Asian mainland near the Sea of Japan but also from Kunashir

Island.

Only a few species have distribution patterns that extend southward from

the Sea of Japan area, and the currents around the Japanese Archipelago are

probably responsible for this distribution, too. The prevailing winds associated

with the warm waters of the Kuroshio Current, which begin along the eastern

shores of Taiwan, might carry fungal spores to Japan, and there are interesting

examples of distribution patterns that extend along the eastern coast of Asia

from the Kurile Islands to an area of the mainland at approximately the same

latitude as Taiwan. For example, Spirodecospora melnikii (Lar.N. Vassiljeva)

K.D. Hyde & Melnik, described from Kunashir Island (Vasilyeva 1990: as

Anthostomella melnikii), also has been found in Korea (Melnik et al. 2005)

and near Hong Kong (Lu et al. 1998). The Hong Kong fungus was described

originally as Spirodecospora bambusicola B.S. Lu et al., now regarded as a

synonym of S. melnikii (Melnik & Hyde 2003). The mainland coastline of China

is mentioned above as one characteristic area of the Sino-Japanese Floristic

Region, and further investigations may reveal a whole complex of species with

the same distribution.

Similar distributional regularities also occur in a very distantly related

group of fungi. Fifty-three of 123 rust species (43%) found in Russia

belonging to Chaconiaceae, Coleosporiaceae, Cronartiaceae, Melampsoraceae,

Mikronegeriaceae, Phakopsoraceae, and Pucciniastraceae are restricted to the

Sea of Japan area (Azbukina & Karatygin 2010). Within this group, a number

of species seem to have had their spores carried by the air current associated

Pyrenomycetes of the Sea of Japan area... 11

with the Soya Current from Japan to Sakhalin Island and southern Kurile

Islands. Consequently, Coleosporium yamabense (Saho & I. Takah.) Hirats.

f., Naohidemyces fujisanensis S. Sato et al., and Uredinopsis woodsiae Kamei

are shared by Japan and Sakhalin Island (Azbukina 2005); Pucciniastrum

hydrangeae-petiolaris Hirats. f. is known from Japan, Sakhalin Island, and

Kunashir Island; Thekopsora tripetaleiae Hirats. f. has been reported from Japan

and Iturup Island; and T: menziesiae Hirats. f. is known from Japan, Kunashir

Island, and Iturup Island. The Taiwanese connection is also found in the rust

fungi: Blastospora itoana Togashi & Onuma and Milesina miyabei Kamei are

found both in the Sea of Japan area (China, Japan, the southern portion of the

Russian Far East) and Taiwan (Ono et al. 1987; Azbukina 2005).

Zoogeographical studies clearly also distinguish the so-called ‘Sino-Japanese

realm (Holt et al. 2013). This realm does not encompass such southern countries

as Burma, Nepal, or Vietnam but extends further to the west of northern China

than suggested for the center of fungal biodiversity around the Sea of Japan. In

this context, a rather interesting fungus described recently by Zhuang & Bau

(2008), namely Sinofavus allantosporus W.Y. Zhuang & T. Bau, merits attention.

One specimen was collected in 2003 in the Magadan region (north-eastern

Russia) by Nina Sazanova and sent to Larissa Vasilyeva but not described.

A second collection found again in 2007 by Tolgor Bau in the westernmost

Chinese province of Xinjiang (Tuomer Peak) was designated as the holotype of

the species. Sinofavus allantosporus seems to be restricted to Populus species, the

substrate of the Nina Sazanova specimen; the illustration provided in Zhuang &

Bau (2008: Fig. 1) shows the branch (apparently from Populus) upon which the

Chinese specimen fruited. As such, a double restriction—first to substrate and

then to the northern mountainous region of northeastern Russia and northern

China—is characteristic of S. allantosporus, and it is extremely surprising that

such a peculiar discomycete with compound fruitbodies remained completely

unnoticed until the 21* century. All these preliminary observations indicate

the need for further investigation of several species complexes in north-eastern

Asia that appear to have their own biogeographical patterns.

Acknowledgments

We thank Dr. David Orlovich (University of Otago) and Prof. Carol Shearer

(University of Illinois) for serving as presubmission reviewers and for providing helpful

comments and suggestions.

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MY COTAXON

http://dx.doi.org/10.5248/126.15

Volume 126, pp. 15-22 October-December 2013

Hyphomycetes: Linodochium sinense sp. nov. and

new records from fallen Sycopsis sinensis leaves in China

DeE-WEI L1*", JINGYUAN CHEN? & YIXUN WANG?

'The Connecticut Agricultural Experiment Station, Valley Laboratory,

153 Cook Hill Road, Windsor, CT 06095, USA

?Institute of Forest Disease and Insect Control, Hubei Academy of Forestry,

I Lion Peak, Jiufeng, Wuhan, Hubei, 430075, China

* CORRESPONDENCE TO: dewei.li@ct.gov

ABSTRACT — Among hyphomycetes collected from dead leaves of Sycopsis sinensis in

Houhe National Nature Reserve, Wufeng, Hubei, China, a sporodochial fungus new to

science, Linodochium sinense, is described and illustrated. It develops yellow conidiomata

and 2-celled conidia that are filiform, colorless, smooth, 34-40 x 1.1-1.5 um. Candelabrum

microsporum, Chalara affinis, Cylindrocladium madagascariense, and Subulispora britannica

are new records from China, while Menisporopsis novae-zelandiae is newly reported from

mainland China.

KEY worpDs — anamorphic fungi, saprobe

Introduction

Houhe National Nature Reserve in Wufeng Tujia Autonomous County,

Hubei, China is exceptional for its rich biodiversity and rare species. With over

2292 species of vascular plants including 29 endangered species (Song & Liu

1999, Li et al. 2005), the Reserve also possesses some species native only to

China (Dang et al. 2004, Li et al. 2005). Its fungal diversity is almost entirely

unexplored, and only two studies (on ascomycetes and polypores) have been

conducted there (Li et al. 2007, Zhuang et al. 2007).

Sycopsis sinensis Oliv. (Hamamelidaceae) is a native evergreen broadleaf

woody plant that grows sporadically in mixed evergreen broadleaf forests or

bushes at elevations of 1000-2000 m in southern China (Li et al. 2003). The

largest population of S. sinensis as dominant species has been reported in

Houhe National Nature Reserve (Liu et al. 1999).

Two field trips were made to the Reserve to collect hyphomycetes in August

2008 and May 2009. Hyphomycetes on dead leaves of S. sinensis were studied

16 ... Li, Chen & Wang

and categorized. Three new hyphomycetes from the two surveys have already

been published, one of them from S. sinensis (Li et al. 2010, Li et al. 2011).

Five additional hyphomycetes from S. sinensis are reported here, one as a

new species, three as new records from China, and one as a new record from

mainland China.

Materials & methods

Sporodochia, conidiophores and conidia of the fungus were mounted in lacto-

fuchsin (0.1 g acid fuchsin, 100 ml 85% lactic acid; Carmichael 1955) or 85% lactic

acid. Microscopic observations were made using bright field and Nomarski differential

interference contrast optics. Photomicrographs were taken with an Olympus MicroFire

digital camera (Goleta, CA). Fungal structures were measured with PictureFrame 2.3

software (Goleta, CA). Measurements of the fungal structures were statistically analyzed

with Microsoft Office Excel 2010 with 95% confidence interval of means. The results

are presented as ranges and mean + standard deviation. Q = length/width ratio and

n = the number of fungal structures measured. Herbarium acronyms follow Index

Herbariorum (Thiers 2013).

Taxonomy

Linodochium sinense D.W. Li, Jing Y. Chen & Yi X. Wang, sp. nov. Fics 1-6

MycoBank MB 804385

Differs from other Linodochium spp. by conidiogenous cells that bear 1-3 (mostly 2)

1-septate conidia with dichotomous denticulate apices and conidial anastomoses that

form x-shaped derivative conidia.

Type: China, Hubei, Wufeng, Houhe National Nature Reserve, on dead leaves of Sycopsis

sinensis, 20 May 2009, Yixun Wang (Holotype: BPI 892531).

EryMmo oey: referring to China for its origin.

CoNIDIOMATA sporodochial, circular to irregular, bright yellow when young,

orange at maturity, scattered, hypophyllous, occasionally amphigenous,

200-300 um diam. and 90-110 um high. Conrp1opHorss differentiated, erect,

branched, flexuous or straight, colorless to pale yellow, smooth, septate, packed

together. CONIDIOGENOUS CELLS integrated, terminal, holoblastic, straight or

flexuous, bacilliform or ampulliform, sympodially elongated, smooth, thin-

walled, colorless, (11-)12-16(-19) x (1-)1.2-1.4(-1.7) um (mean = 14.4

+ 1.8 x 1.3 + 0.1, n = 30), bearing mostly (1-)2(-3) conidia; apices of some

conidiogenous cells ending with two flat-topped denticles of different lengths

Figures 1-8. Linodochium sinense (Holotype: BPI 892531). 1. Sporodochia on lower epidermis of a

leaf of Sycopsis sinensis, 2. Cross-sections of sporodochia. 3. Conidiogenous cells and conidiophores.

4. Conidia on conidiogenous cells and four pairs of conidia on the right anastomosed into x-shaped

derivative conidia. 5. An x-shaped derivative conidium. 6-7. Conidia on a conidiogenous cell with

a dichotomous denticulate apex. 8. Three conidia on a conidiogenous cell with a trichotomous

denticulate apex. Scale bars: 1 = 1 mm; 2 = 10 um; 3-8 = 5 um.

Linodochium sinense sp. nov. (China) ... 17

18 ... Li, Chen & Wang

forming a dichotomy, some with one denticle and one flat conidiogenous

locus on the other side of the conidiogenous cell apex, rarely, conidiogenous

cells ending in a trichotomous denticulate apex. Conrp1A 1-septate, filiform,

colorless, smooth, guttulate, (32-)34-40(-45) x (1-)1.1-1.5(-1.6) um (mean =

37.543 x 1.3 + 0.2, n = 30), Q = (22.5-)24.5-32.1(-39.3) (mean = 28.3 + 3.8, n

= 30); some conidial pairs anastomosing to form x-shaped derivative conidia.

TELEOMORPH: unknown.

GEOGRAPHICAL DISTRIBUTION: Hubei, China.

Hasirat: dead leaves of Sycopsis sinensis.

Notes: Linodochium was erected by Hohnel (1909) and typified by Linodochium

hyalinum (Lib.) Hohn. It remained monotypic for seven decades. Dyko &

Sutton (1979) redescribed and illustrated the genus and L. hyalinum, its only

species. Two years later, Minter & Holubova-Jechova (1981) added a second

species, L. formosum Minter & Hol.-Jech. During the 1990’s two more species

were described: Linodochium album R.E. Castafieda & W.B. Kendr. (Castaneda

& Kendrick 1990) and L. hyalinum, L. sideroxyli Dulym. et al. (Dulymamode et

al. 1999). Linodochium sinense is the fifth species proposed in the genus. Much

longer conidia differentiate L. formosum (80-140 um with 2-4 septa; Dyko &

Sutton 1979) and L. hyalinum (36-89 um with <12 septa; Minter & Holubova-

Jechova 1981) from L. sinense. White sporodochia and falcate conidia of

L. album (Castafieda & Kendrick 1990) differ from the yellow sporodochia and

filiform conidia of L. sinense. Linodochium sideroxyli is morphologically similar

to L. sinense but differs in having wider conidia (1.5-2 um, to 2.5 um at the

apex, 0-1 septum; Dulymamode et al. 1999) without dichotomous denticulate

apices.

The significance of conidial anastomosis to form x-shaped derivative conidia

in L. sinense is not clear. It might function as parasexual behavior or as part

of sexual reproduction. However, further study is necessary to determine its

function.

The teleomorph of L. hyalinum was found to be Pseudohelotium pineti

(Batsch) Fuckel (Minter & Holubova-Jechova 1981). However, we found no

teleomorph for L. sinense.

There are no DNA sequence data for any Linodochium species deposited

in GenBank, and we were unable to isolate DNA from the L. sinense type

specimen.

Key to species of Linodochium

LiSperodochia yellowsambercon orange r.t.05 kit.o8 Aut ott ed hd ee he et 2

1. Sporodochia white to grayish white ........ 0... eee eee eee ee eee 4

2, Gonidia-2—4 septate; 36-89 x AR 1, Soin aces Vera abs evade Mavacly Hevacde Bera L. hyalinum

2 Gontdia 0= Tasep tate: Fy, sos aton tig tar pte aates enue Beane Bena Pat nat eae 3

Linodochium sinense sp. nov. (China) ... 19

3. Conidiogenous cells bearing 1-4 conidia, no denticles at apical end;

conidia 0-1-septate, 32-48 x 1.5-2 um, wider toward apex ........ L. sideroxyli

3. Conidiogenous cells bearing mostly 2 conidia, with dichotomous denticles at apex;

conidia 1-septate, 34-40 x 1.1-1.5 um, conidial anastomosis forming x-shaped

Gerivalivestomicids &.c4 eid hee eee ee ge ee ee ety L. sinense

4, Conidia straight, <12-septate, 80-140 x 2-3 um...............0008. L. formosum

4. Conidia falcate, 1-2-septate, 30-45 x 1-2 um .... 6... eee eee ee L. album

New records for China

Candelabrum microsporum R.F. Castafieda & W.B. Kendr., Univ. Waterloo Biol. Ser.

35: 16 (1991)

TELEOMORPH: unknown.

CONIDIOPHORES reduced to conidiogenous cells. CONIDIOGENOUS CELLS

monoblastic. Conip1A subhyaline, solitary; conidial body multi-branched

dicho- or sometimes trichotomously, with minute apical tubercles, forming

globose, subglobose or elongated conidia, 35-55 um diam.

SPECIMEN EXAMINED: CHINA, HuBeI, Wufeng, Houhe National Nature Reserve, dead

leaf of Sycopsis sinensis, 28 August 2008, De-Wei Li (BPI 892532.

ECOLOGY & DISTRIBUTION: On dead leaves of Sycopsis sinensis; Palmae.

Previously known from Cuba; new record from China.

Note: Some conidia from the Chinese specimen are elongated and deviate

slightly from the conidial shape of the holotype. Zhao et al. (2007) reported two

other Candelabrum species in China, C. brocchiatum Tubaki and C. spinulosum

Beverw. Both species were collected on decaying branches of an undetermined

tree submerged in water in Dashuwang, Mount Tianmu, Zhejiang.

Chalara affinis Sacc. & Berl., Atti Ins. Veneto Sci. Lett.. Arti, Ser. 6 3: 741 (1885)

TELEOMORPH: unknown.

CONIDIOPHORES simple, erect, straight or slightly flexuous, cylindrical with

an inflated base, <4 septate, sometimes constricted at the septa, pale brown,

smooth, 40-70 um long, 4-6 um wide at the base. PH1aLipEs lageniform,

pale brown, smooth, 35-50 x 4-6.5 um. Conip1A 1-celled, colorless, smooth,

cylindrical, ends flat or slightly rounded, 8-11 x 1.5-2 um, catenate.

SPECIMEN EXAMINED: CHINA, HuBe!, Wufeng, Houhe National Nature Reserve, dead

leaf of Sycopsis sinensis, 28 August 2008, De-Wei Li (BPI 892533).

ECOLOGY & DISTRIBUTION: On dead leaves of Sycopsis sinensis; Pinus sylvestris

(Minter & Holubova-Jechova 1981). Previously known from Czech Republic

and the United Kingdom; new record from China.

NoTE: the conidia from the Chinese material are slightly smaller than those

of the type.

20 ... Li, Chen & Wang

Cylindrocladium madagascariense Crous, Tax. Path. Cylindrocladium: 112 (2002)

TELEOMORPH: Calonectria madagascariensis Crous.

CONIDIOPHORES colorless, smooth, penicillate, 60-85 x 5-6.5 um, with a

septate extension, 130-350 um long, terminating in a clavate vesicle. PHIALIDES,

colorless, smooth, allantoid, cylindrical, or doliiform, 9-11 x 3-4 um. CONIDIA

cylindrical, rounded at both ends, straight, colorless, smooth, thin-walled,

3-spetate, 50-55 x 4-5 um.

SPECIMEN EXAMINED: CHINA, HuBeI, Wufeng, Houhe National Nature Reserve, dead

leaf of Sycopsis sinensis, 20 May 2009, Yixun Wang (BPI 892534).

ECOLOGY & DISTRIBUTION: On fallen leaves of Sycopsis sinensis; Eucalyptus sp.,

soil. Previously known from Congo and Madagascar (Crous 2002); new record

from China.

Notes: The Chinese collection had shorter conidiophore extensions and

slightly shorter conidia than those of the holotype. Crous (2002) reported that

Cylindrocladium madagascariense is pathogenic to Eucalyptus sp., causing a leaf

spot. It remains unknown whether this species does the same to S. sinensis.

Further observation of S. sinensis foliage prior to leaf abscission is necessary to

clarify this.

Subulispora britannica B. Sutton, Trans. Br. Mycol. Soc. 61(3): 422 (1973)

TELEOMORPH: unknown.

CONIDIOPHORES 20-40 x 3.5-4 um, differentiate, solitary, straight or

flexuous, 1-4 septate, smooth, thick-walled, pale brown to brown, sometimes

with an enlarged base. CONIDIOGENOUS CELLS integrated, terminal polyblastic,

sympodial, subhyaline to pale brown, thin-walled with 1-3 conspicuous flat

unthickened scars. ConIpIA solitary, 3-10-septate, colorless, filiform, straight,

thin-walled, smooth, 50-90 um long and 2.5-3 um wide at the base, attenuated

toward the apex.

SPECIMEN EXAMINED: CHINA, HuBe!, Wufeng, Houhe National Nature Reserve, dead

leaf of Sycopsis sinensis, 28 August 2008, De-Wei Li (BPI 892535).

ECOLOGY & DISTRIBUTION: on dead leaves of Sycopsis sinensis; Hedera helix,

Ilex aquifolium, I. pernyi, and Quercus suber (Kirk 1982). Previously known

from Russia and UK; new record from China.

Notes: The Chinese conidiophores and conidia are slightly longer than

those of the type.

New record for Mainland China

Menisporopsis novae-zelandiae S. Hughes & W.B. Kendr., N.Z. J. Bot. 6: 369 (1968)

TELEOMORPH: unknown.

SETAE solitary, dark brown, pale brown toward the apex, <17 septate,

250-400 um long, 8-14 um at the base. ConipIOpHORES 40-75 x 1.5-2 um.

Linodochium sinense sp. nov. (China) ... 21

CONIDIOGENOUS CELLS monophialidic, smooth, colorless or subhyaline,

straight or slightly curved, 15-20 x 3-4 um. COLLARETTE conspicuous,

infundibuliform or cupulate, 2-3 um wide, 1.5-3.5 um deep. CONIDIA

1-septate, allantoid, colorless, smooth, 16-20 x 2.5-3 um with a simple setula

at each end, 4-6 um long.

SPECIMEN EXAMINED: CHINA, HuBEI, Wufeng, Houhe National Nature Reserve, dead

leaf of Sycopsis sinensis, 20 August 2009, Yixun Wang (BPI 892536).

ECOLOGY & DISTRIBUTION: on dead leaves of Sycopsis sinensis; Beilschmiedia

tarairi, Cinnamomum osmophloeum, C. pedunculatum, C. zeylanicum,

Cryptocarya mackinnoniana, Knightia excelsa, Litsea japonica, palm, Pasania

edulis, P. kawakamii (Hughes & Kendrick 1968, Matsushima 1980, 1989, 1993).

Previously known from Australia, Ecuador, Japan, New Zealand (Matsushima

1975, 1980, 1989, 1993). This is the first report from Mainland China.

Matsushima (1980) reported this species from Taiwan.

Notes: The Chinese collection had shorter setae than those of the type and

its conidiogenous cells are not polyphialidic.

Acknowledgments

The authors express their sincere gratitude to Dr. Bryce Kendrick and Dr. Rafael F.

Castafieda Ruiz for their critical review of the manuscript and to Drs. Louis Magnarelli

and James A. LaMondia for their pre-submission review. The authors are also grateful to

Dengkui Tang, Zhizhang Zheng, Yeqing Wang, Sanshan Cai, and Jihong Yang for their

assistance during the field trips. Dr. Lorelei L. Norvell’s editorial review and Dr. Shaun

Pennycook’s nomenclature review are greatly appreciated. This paper is dedicated to Dr.

Louis Magnarelli, director of The Connecticut Agricultural Experiment Station, who

passed away on July 11, 2013. DWL expresses his heartfelt gratitude to Dr. Magnarelli

for his support in the past nine years.

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MY COTAXON

http://dx.doi.org/10.5248/126.23

Volume 126, pp. 23-30 October-December 2013

Colletotrichum fructicola, first record of

bitter rot of apple in China

DaN-DAN Fu*?, WEI WANG’, RUI-FENG QIN’, RONG ZHANG",

GUANG-YU SUN™ & MARK L. GLEASON?

'State Key Laboratory of Crop Stress Biology in Arid Areas and College of Plant Protection,

Northwest A&F University, Yangling, Shaanxi, 712100, P. R. China

College of Food & Bioengineering, Henan University of Science and Technology,

Luoyang, Henan, 471023, P. R. China

*Department of Plant Pathology and Microbiology, Iowa State University, Ames, Iowa 50011, USA

“CORRESPONDENCE TO: “rongzh@nwsuaf.edu.cn OR ®* sgy@nwsuaf.edu.cn

ABSTRACT — Colletotrichum fructicola, isolated from diseased apple fruit with bitter

rot symptoms collected in orchards of Henan Province, is a new record for China. It also

represents a new pathogen species for apple bitter rot. Phylogeny inferred from combined data

sets of the complete rDNA ITS1-5.8S-ITS2 region, partial actin (ACT), B-tubulin-2 (TUB2)

and glyceraldehyde-3-phosphate dehydrogenase (GPDH) confirmed that the fungus should

be included within C. fructicola, and this grouping was also congruent with morphology and

culture characteristics.

KEY worpDs — anamorph, molecular, taxonomy, pathogenicity

Introduction

A worldwide fungal disease, bitter rot of apple, can cause serious economic

loss. Since 1990, the major pathogens causing bitter rot were reported as

Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. and its teleomorph,

Glomerella cingulata (Stoneman) Spauld. & H. Schrenk; C. acutatum J.H.

Simmonds plays a relatively minor role (Gonzalez et al. 2006).

Colletotrichum is one of the most economically important fungal genera

(Hyde et al. 2009a). Colletotrichum species are cosmopolitan; multiple species

can occur on a single host, and a single species is often found on multiple hosts

(Cai et al. 2009). Sutton (1992) noted that morphology alone is not sufficient fora

precise identification because the lack of reliable morphological features makes

species boundaries ambiguous and confusing, especially in the C. acutatum,

C. gloeosporioides, and C. dematium complexes. Reliable species identification is

important for biosecurity, plant breeding, and integrated disease management.

24 ... Fu & al.

Since 2009, MycoBank has recorded 28 new Colletotrichum species,

including six previously considered to belong to the C. gloeosporioides complex.

One species, C. fructicola isolated from coffee berries in Thailand, was separated

from C. gloeosporioides complex by Prihastuti et al. (2009) based on multi-gene

phylogeny and morphological characters.

In China, C. gloeosporioides was assumed to be the sole pathogen causing

apple bitter rot (Wang & Li 1987) until Zhang et al. (2008) reported that

C. acutatum could also cause the disease. In order to identify the Colletotrichum

species causing bitter rot of apple in China, we collected and isolated the

pathogen from diseased apple fruit in orchards in Henan Province in 2011 and

2012,

Materials & methods

Isolates & morphology

Apple orchards in Henan Province, China were surveyed for bitter rot symptoms in

the autumn of 2011 and 2012. A small piece of decayed tissue was cut from arbitrarily

selected lesions on fruits and disinfected with 70% ethanol before placed on potato

dextrose agar (PDA). Four strains were isolated and incubated at 28°C in darkness for

7 days on PDA. A mycelial disc was then removed from the actively growing edge of

a mono-conidial colony and transferred to new PDA. Three replicate cultures of each

isolate were investigated. After 7 days at 25°C in darkness, size and shape of 50 conidia

were measured (Than et al. 2008), and the colony diameter, conidial mass colors, and

colony zonations were also recorded. Appressoria were produced using a slide culture

technique, in which a 1 cm? segment of PDA containing the isolate was placed in sterile

water in a sterile Petri dish, covered with a sterile cover slip, and incubated under high

humidity at 25°C in darkness. After 2 days, the shape and size of 50 appressoria on

the cover slip were recorded. The F11PGZHO03 living culture was deposited in China

General Microbiological Culture Collection Center (CGMCC), Beijing, China.

DNA extraction & PCR sequencing

The protocol of Barnes et al. (2001) was used to extract genomic DNA from

mycelium growing on PDA. The primers ITS1-F and ITS4 were used to amplify the

ITS region of rDNA. The ACT gene was amplified with primer pair ACT512F and

ACT783R; TUB2 with Bt2a and Bt2b; and GPDH with GDF1 and GDR1 (Prihastuti

et al. 2009). The amplification conditions of PCR reactions were: initial denaturation

at 95°C for 3 min followed by 34 cycles of denaturation at 95°C for 60 s, annealing at

52°C for 30 s, extension at 72°C for 1 min and a final cycle of 10 min at 72°C. The PCR

products were sequenced by Sangon Biotech Co., Shanghai China, and the results were

manually checked for errors.

Phylogenetic analysis

Sequence data from ITS, ACT, TUB2, and GPDH genes of our isolate were deposited

in GenBank and compared with other Colletotrichum sequences from GenBank

Colletotrichum fructicola on apple (China) ... 25

TABLE 1. Colletotrichum sequences used in the phylogenetic analysis

i GENBANK ACCESSION NUMBERS*

SPECIES / STRAIN

ITS ACT TUB2 GPDH

C. asianum BML 13 ' FJ972605 FJ903188 FJ907434 FJ972571

BPD I4! FJ972612 FJ907424 FJ907439 FJ972576

C. falcatum FAL ' FJ972606 FJ907431 GQ289454 FJ972585

C. fructicola BPD 112! FJ972611 FJ907425 FJ907440 FJ972577

BPD 116? FJ972603 FJ907426 FJ907441 FJ972578

F11PGZHO03 ? KC012513 KC012505 KC012517 KC012509

C. gloeosporioides CBS 953.97 ! FJ972609 FJ907430 FJ907445 FJ972582

C. hymenocallidis CSSN2 ° GQ485600 GQ856775 GQ849438 GQ856757

CSSN3 ? GQ485601 GQ856776 GQ849439 GQ856759

C. kahawae IMI 319418 ! FJ972608 FJ907432 FJ907446 FJ972583

IMI 363578 ! FJ972607 FJ907433 FJ907447 FJ972584

C. siamense BPD 12! FJ972613 FJ907423 FJ907438 FJ972575

BMLI15! FJ972614 FJ907422 FJ907437 FJ972574

C. simmondsii BRIP 28519 * FJ972601 FJ907428 FJ907443 FJ972580

*Type strains and sequences in bold font

'Prihastuti et al. (2009); * this paper; * Yang et al. (2009); * Hyde et al. (2009b).

(TABLE 1), including those of several type specimens (Hyde et al. 2009b, Prihastuti et

al. 2009, Yang et al. 2009). Preliminary alignments of the four gene sequences were

conducted using Clustal X (Thompson et al. 1997) with manual adjustment, and using

BioEdit for visual improvement wherever necessary. Phylogenetic analysis of the

combined ITS/ACT/TUB2/GPDH sequences was performed using PAUP (Phylogenetic

Analysis Using Parsimony) v. 4.0b10 (Swofford 2003). All characters were unordered

and of equal weight and gaps were treated as missing data. Clade stability of the tree

resulting from maximum parsimony analysis (MP) and neighbor-joining analysis (NJ)

was evaluated by 1000 bootstrap replications on the same aligned dataset (Hillis & Bull

1993). Measures calculated for parsimony included tree length (TL), consistency index

(CI), retention index (RI), and rescaled consistency index (RC). The outgroup was

Colletotrichum falcatum.

Pathogenicity testing

The spores of isolate F11PGZH03 were harvested from PDA cultures, and the

concentration of spore suspension in sterile water was adjusted to 10° conidia/ml using

a haemocytometer. Mature and non-wounded Fuji apple fruits were surface-sterilized

with 70% ethanol prior to inoculation. Apples were inoculated by using wound/drop and

non-wound/drop inoculation methods (Prihastuti et al. 2009). The isolate inoculation

was repeated three times. Control fruits were treated similarly to the inoculated fruit, but

with sterile water instead of spore suspension. After inoculation, the fruits were placed

in a closed plastic bag to maintain high humidity and held at 25°C in the darkness. The

length and width of typical symptoms on the skin of fruit were recorded after 5 and 10

days.

26 ... Fu & al.

Fic. 1. Colletotrichum fructicola (F11PGZH03). A-B, colony on PDA after 7 days, (A = upper;

B = reverse). C, bitter rot symptom on apple skin. D-E, conidia. F-H, appressoria. Bars: D-H = 10 um.

Colletotrichum fructicola on apple (China) ... 27

Results

Taxonomy

Based on morphological, cultural, and multigene phylogenetic data, the

isolates were identified as:

Colletotrichum fructicola Prihastuti, L. Cai & K.D. Hyde, Fungal Diversity 39: 96.

2009. Fie. 1

SEXUAL MORPH not observed. ASEXUAL MORPH on PDA. Hypuae hyaline,

smooth-walled, septate, branched. Conrp1opHorEs formed directly on hyphae.

SETAE not observed. Conip1A hyaline, smooth-walled, aseptate, straight,

cylindrical with both ends round or one end slightly acute, (12.3-)1-3.5-

15.5(-17.1) x (4.3-)4.7-5.5(-6.3) um, mean + SD = 14.59 + 1.16 x 5.11 + 0.38,

L/W ratio = 2.9. AppREssoRIA single or in loose groups, pale to dark brown,

ovoid and slightly irregular to irregular in shape and often becoming complex

with age, (6.4—)7-9(-10.5) x (4.7—)5.2-6.2(-6.9) um, mean + SD = 8.08 + 1.03

x 5.78 + 0.54, L/W ratio = 1.4.

CULTURAL CHARACTERISTICS — Colonies grown from single conidia on

Difco PDA 78-80 mm diam after 7 d. Aerial mycelium white to pale grey,

dense, cottony, surface of agar pale grey to dark grey towards centre, more of

less white at the edge; acervuli not observed. In reverse grayish green towards

centre with white halo. In 7 days at 28°C, growth rate 8.5-9.25 mm/day (mean

+ SDi=; 9-064 O37 sen 3/9):

SPECIMENS EXAMINED: CHINA, HENAN PROVINCE: ZHENGZHOU CIry, on fruit surface

of apple (Malus domestica Baumg.), 28 Sept. 2011, Dandan Fu F11PGZHO03 (living

culture, CGMCC 3.15207; GenBank KC012513, KC012505, KC012517, KC012509);

SHANGQIU CITY, 6 Sept. 2012, F12PGSQ5; F12PGSQ6; X1ay1 CouNTY, 7 Sept. 2012,

F12PGXY1.

Phylogenetic analysis

Of 1863 characters aligned with Clustal X and optimized manually to allow

maximum alignment and maximum sequence similarity by BioEdit, 246 were

parsimony informative. Ambiguously aligned regions were excluded from all

analyses. The Kishino-Hasegawa (KH) test showed that three trees generated

from parsimony analysis were retained. One of the most parsimonious trees is

shown in Fia. 2 (TL = 867, CI = 0.9331, HI = 0.0669, RI = 0.8578, RC = 0.8005).

MP and NJ bootstrap values are provided above branches. The phylogram in

Fic. 2 illustrates that our isolate F11PGZH03 grouped with two C. fructicola

isolates in a single clade with 100%/100% bootstrap support, indicating that

they may represent the same species.

In addition to isolating F11PGZH03, we isolated three strains of C. fructicola

in Henan Province in 2012. The grouping of these three strains was well

supported by phylogenetic analysis of ITS sequences and multilocus sequence

28 ... Fu & al.

99/100 C. fructicola BPDI12

100/100 |! ©. fructicola BPDI16 C. fructicola

F11PGZHO03

C. asianum BPDI4

100/100

82/69 C. asianum BMLI3

90/1 J C. hymenocallidis CSSN3

C. hymenocallidis CSSN2

96/100

C. siamense BMLI15

C. siamense BPDI2

90/96

100/100 100/100

C. gloeosponoides CBS953.97

C. kahawae IMI319418

tOi1G0 C. kahawae |IMI363578

C. simmondsii BRIP 28519

10 Colletotrichum falcatum FAL

Fic. 2. Phylogram generated from maximum parsimony and neighbor-joining analyses based on

combined ITS, ACT, TUB2, and GPDH sequence data. Clade stabilities shown above branches

were calculated from maximum parsimony (>50%) and distance (>50%). The scale bar shows 10

changes. The tree is rooted with Colletotrichum falcatum.

Colletotrichum fructicola on apple (China) ... 29

data (results not shown). The colonies, conidia, and appressoria of our isolates

were morphologically distinguishable from those of C. gloeosporioides complex.

Pathogenicity tests

All four isolates produced typical symptoms of bitter rot in Fuji apples by

wound inoculation, except for isolate F11PGZH03, which did not produce

rot symptoms after non-wound inoculation. Symptoms produced by each

isolate were similar, and there were no apparent differences in lesion size and

characters among isolates.

Discussion

Based on phylogenetic analysis of combined ITS/ACT/TUB2/GPDH

datasets and morphological characters of the anamorph, we identified our

isolates as Colletotrichum fructicola, which represent a new species record for

the genus in China.

Colletotrichum fructicola was previously described only from coffee berries

and leaf spots of peanut (Prihastuti et al. 2009). In this study we isolated the

fungus from apple, which is a newly reported host for C. fructicola. Based

on the pathogenicity test, C. fructicola was also determined to be a bitter rot

pathogen of apple fruit.

Acknowledgments

This work was supported by National Natural Science Foundation of China

(31171797), Program for Changjiang Scholars and Innovative Research Team in

University (IRT0748), the 111 Project from Education Ministry of China (B07049) and

Top Talent Project of Northwest A&F University. The authors wish to thank Dr Eric

H.C. McKenzie (Landcare Research, Private Bag 92170, Auckland, New Zealand) and

Professor Zhongyi Zhang (College of Plant Protection, Yunnan Agricultural University,

Kunming, Yunnan, China) for reviewing the manuscript.

Literature cited

Barnes I, Roux J, Wingfield MJ, Coetzee MPA, Wingfield BD. 2001. Characterization of Seiridium

spp. associated with cypress canker based on $-tubulin and histone sequences. Plant Disease

85: 317-321. http:// dx.doi-org/10.1094/PDIS.2001.85.3.317

Cai L, Hyde KD, Taylor PWJ, Weir BS, Waller JM, Abang MM, Zhang JZ, Yang YL, Phoulivong S,

Liu ZY, Prihastuti H, Shivas RG, McKenzie EHC, Johnston PR. 2009. A polyphasic approach

for studying Colletotrichum. Fungal Diversity 39: 183-204.

Gonzalez E, Sutton TB, Correll JC. 2006. Clarification of the etiology of Glomerella leaf spot and

bitter rot of apple caused by Colletotrichum spp. based on morphology and genetic, molecular,

and pathogenicity tests. Phytopathology 96: 982-992.

http://dx.doi.org/10.1094/PHY TO-96-0982

Hillis DM, Bull JJ. 1993. An empirical-test of bootstrapping as a method for assessing confidence

in phylogenetic analysis. Systematic Biology 42: 182-192. http://dx.doi.org/10.2307/2992540

Hyde KD, Cai L, McKenzie EHC, Yang YL, Zhang JZ, Prihastuti H. 2009a. Colletotrichum: a

catalogue of confusion. Fungal Diversity 39: 1-17.

30 ... Fu & al.

Hyde KD, Cai L, Cannon PF, Crouch JA, Crous PW, Damm U, Goodwin PH, Chen H, Johnston

PR, Jones EBG, Liu ZY, McKenzie EHC, Moriwaki J, Noireung P, Pennycook SR, Pfenning

LH, Prihastuti H, Sato T, Shivas RG, Tan YP, Taylor PWJ, Weir BS, Yang YL, Zhang JZ. 2009b.

Colletotrichum - names in current use. Fungal Diversity 39: 147-182.

Prihastuti H, Cai L, Chen H, McKenzie EHC, Hyde KD. 2009. Characterization of Colletotrichum

species associated with coffee berries in northern Thailand. Fungal Diversity 39: 89-109.

Sutton BC. 1992. The genus Glomerella and its anamorph Colletotrichum. 1-26, in: JA Bailey, MJ

Jeger (eds). Colletotrichum: biology, pathology and control. CAB International: Wallingford.

Swofford DL. 2003. PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version

4.0. Sinauer Associates, Sunderland, Massachusetts, USA.

Than PP, Shivas RG, Jeewon R, Pongsupasamit S$, Marney TS, Taylor PWJ, Hyde KD. 2008.

Epitypification and phylogeny of Colletotrichum acutatum J.H. Simmonds. Fungal Diversity

28: 97-108.

Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. 1997. The Clustal_X windows

interface: flexible strategies for multiple sequence alignment aided by quality analysis tools.

Nucleic Acids Research 25: 4876-4882. http://dx.doi.org/10.1093/nar/25.24.4876

Wang XM, Li JY. 1987. Studies on species of Colletotrichum in Province Shaanxi, China. Acta

Mycologica Sinica 6: 211-218.

Yang YL, Liu ZY, Cai L, Hyde KD, Yu ZN, McKenzie EHC. 2009. Colletotrichum anthracnose of

Amaryllidaceae. Fungal Diversity 39: 123-146.

Zhang R, Wang SF, Cui JQ, Sun GY. 2008. First report of bitter rot caused by Colletotrichum

acutatum on apple in China. Plant Disease 92: 1474.

http://dx.doi.org/10.1094/PDIS-92-10-1474B

MY COTAXON

http://dx.doi.org/10.5248/126.31

Volume 126, pp. 31-36 October-December 2013

Phyllobaeis crustacea sp. nov. from China

SHUNAN Cao’’, XINLI WEI’, QIMING ZHOU’ & JIANGCHUN WEI’*

'State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences,

No. 1 Beichen West Road, Chaoyang District, Beijing 100101, China

*University of Chinese Academy of Sciences,

No. 19A Yuquan Road, Beijing 100049, China

*CORRESPONDENCE TO: weijc2004@126.com

ABSTRACT — ‘The lichen-forming fungus genus, Phyllobaeis, is reported for the first time

from China. A new crustose species, P. crustacea, is described and illustrated.

Key worps — Ascomycota, Baeomycetaceae, chemistry, molecular systematics, morphology

Introduction

The lichen-forming fungus genus Phyllobaeis is known from five squamulose

species (Gierl & Kalb 1993, Index Fungorum 2013) occurring in the Neotropics.

During our studies on the lichen flora of China we encountered a species that

showed affinities with this genus but deviated conspicuously by its crustose

(rather than squamulose) thallus. We present our analysis of the morphology,

anatomy, chemistry, and phylogeny of this species to clarify its taxonomy.

Materials & methods

The material used in this study was collected from Hainan, Yunnan, and Qinghai

provinces and Xizang (Tibet) autonomous region of China in 2010 and 2012, and from

the Antarctica in 2011. The collections examined are preserved in the lichen section of

the Herbarium Mycologicum Academiae Sinicae (HMAS-L). A specimen of Phyllobaeis

imbricata was borrowed from the former Herbarium Universitatis Amstedamensis, now

in Museum Naturalis in Leiden (L). A compound microscope (ZEIss Axioskop 2 plus)

and a dissecting microscope (Motic SMZ-168) were used for the study of morphology

and anatomy. A 10% solution of potassium hydroxide (KOH), a 5% bleaching solution

(sodium hypochlorite, NaOCl), concentrated alcoholic p-phenylenediamine (PD),

Lugol’s solution of Iodine, and thin-layer chromatography (TLC) (Culberson &

Kristinsson 1970; Culberson 1972; White & James 1985) were used for the detection of

lichen substances.

32 ... Cao & al.

TABLE 1. Twenty-two nrDNA ITS sequences used in phylogenetic analysis.

SPECIES LOCALITY; VOUCHER SPECIMEN * GENBANK NO. 4

Baeomyces placophyllus Xizang, China; HMAS-L 124223 KC414621

Xizang, China; HMAS-L 124222 KC414620

China; — DQ001274

Baeomyces rufus Qinghai, China; HMAS-L 124225 KC414623

Yunnan, China; HMAS-L 124226 KC414622

France; — AF448457

France; — AF448458

Dibaeis absoluta Hainan, China; HMAS-L 118071 KC414625

Hainan, China; HMAS-L 118073 KC414626

Dibaeis baeomyces — DQ782844

Dibaeis sorediata Hainan, China; HMAS-L 118090 KC414627

Hainan, China; HMAS-L 118097 KC414628

Icmadophila japonica Japan; — AB623070

Phyllobaeis crustacea Hainan, China; HMAS-L 118086 KC414614

Hainan, China; HMAS-L 118087 KC414615

Hainan, China; HMAS-L 118089 KC414616

Hainan, China; HMAS-L 118095 (holotype) KC414617

Hainan, China; HMAS-L 118096 KC414618

Phyllobaeis imbricata Carchi, Ecuador; L 0790053 KC414619

— HQ650635

Placopsis contortuplicata Antarctica; HMAS-L 124227 KC414624

Antarctica; — DQ534479

* Missing data indicated with “—” “ New sequences are shown in bold font.

DNA extraction, PCR amplification and sequencing

The extraction procedure followed the modified CTAB method (Wang et al. 2011).

PCR amplifications were performed using a Biometra T-Gradient thermal cycler.

The primer pair ITS5 and ITS4 (White et al. 1990) was used to amplify the nrDNA

ITS region. Reactions were carried out in 50 ul reaction volume and the components

used were 1 ul total DNA, 2 ul each primer (10 uM), 1 pl Taq polymerase (rTaq DNA

Polymerase, 5 U/l), 4 ul dNTP (2.5 mM each), 5 ul amplification buffer (10x, 25 mM

MgCl, contained), 35 ul ddH,O. Cycling parameters were set to an initial denaturation at

95°C for 5 min, followed by 30 cycles of denaturation at 94°C for 40 s, annealing at 52°C

for 40 s, extension at 72°C for 2 min, and a final extension at 72°C for 10 min. Negative

control, without DNA template, was prepared in every series of amplification in order

to minimize the possibility of contamination. Finally, PCR products were purified by gel

purification kit (Biocolor BioScience & Technology Co. Ltd.). Then, PCR products were

sequenced using ABI 3730 XL DNA Sequencer.

Altogether 22 nrDNA ITS sequences belonging to nine species were used for the

phylogenetic analysis (TABLE 1). Fifteen samples representing seven species were

Phyllobaeis crustacea sp. nov. (China) ... 33

sequenced by the authors, and another seven samples (belonging to six species) were

downloaded from GenBank.

Phylogenetic analysis

All sequences were aligned using ClustalW 1.6 (Higgins et al. 1994). The phylogenetic

analysis was executed with software Mega5.10 (Tamura et al. 2011). The Kimura-2-

parameter was selected as the nucleotide substitution model, and gaps or missing data

were set as pairwise deletion. The maximum likelihood (ML) method was used in

constructing the phylogenetic tree and the reliability of the inferred tree was tested by

1000 bootstrap replications.

Results

The ML-tree (Fic. 1) of the ITS rDNA sequences shows that the five

Phyllobaeis crustacea individuals cluster with 100% bootstrap support onto a

separate branch, which is most closely related to P. imbricata. Together these

two species form a common Phyllobaeis branch with 79% bootstrap support.

Baeomyces placophyllus and B. rufus cluster together with 98% bootstrap

support. Phyllobaeis and Baeomyces represent Baeomycetaceae with 59%

ePhyllobaeis crustacea KC414614

eP. crustacea KC414615

eP. crustacea KC414616

eP. crustacea KC414617 (T)

781 eP. crustacea KC414618

eP. imbricata KC414619 g

“4 100L. P. imbricata HQ650635 3

30) eBaeomyces placophylius KC414620 8 ow

100}! eB. placophyllus KC414621 s &

B. placophyllus DQ001274 © =

go) eB. rufus KC414623 S

©B. rufus KC414622 2

100}) B. rufus AF448457 f

261 B. rufus AF448458 8

ePlacopsis contortuplicata KC414624 S

PI. contortuplicata DQ534479 8

Icmadophila japonica AB623070 2

Dibaeis baeomyces DQ782844 g A

400) @D. absoluta KC414625 8 a

‘a ©D. absoluta KC414626 8 8

eD. sorediata KC414627 3 7

961 eD. sorediata KC414628 2

0.05

Fic. 1. ML tree based on nrDNA ITS region sequences. The species in the tree marked with “e”

were sequenced by the authors. Nucleotide: K2 model, gaps or missing data were partial deletion,

bootstrap = 1000. Genetic distance scale = 0.05. Numbers at nodes present the bootstrap support

value (numbers <50 not shown).

34 ... Cao & al.

bootstrap. Meanwhile, the Baeomycetales, containing the Baeomycetaceae and

Trapeliaceae (represented by Placopsis), is supported by a bootstrap value of

100%, and the Dibaeis species (formerly included in Baeomyces) are clearly

shown to belong to the outgroup, Icmadophilaceae.

Summarizing, the ITS rDNA sequences of P. crustacea differ significantly

from the other species of Baeomycetaceae that are most closely related to the

Phyllobaeis group.

Taxonomy

Phyllobaeis Kalb & Gierl, Herzogia 9:610.1993; emend. S.N. Cao & J.C. Wei

ORIGINAL DIAGNOSIS (Gierl & Kalb 1993: 610): Genus novum a genere Baeomyces differt

thallo squamuloso, superne et infra corticato acido norstictico continente; regionibus

tropicis distributum.

EMENDED DIAGNOsIs: Differs from Baeomyces by its production of norstictic acid and

its tropical distribution.

Phyllobaeis crustacea S.N. Cao & J.C. Wei, sp. nov. FIG. 2

FUNGAL NAME FN570052

Differs from the other Phyllobaeis species by its crustose thallus.

TYPE: CHINA, HAINAN: Changjiang County, Mt. Bawangling, 19°16’ N 109°03’ E, alt. 300

m, on rock, 25 Nov. 2010, S. N. Cao CSN047 (Holotype, HMAS-L 118095, GenBank

KC414617; Isotype: HMAS-L 127984).

Erymo_oey: Latin crustaceus, referring to the crustose thallus.

THALLUS crustose, grayish green, matt, varnish-like, tightly attached to the

substrate, forming a patch of 2.5-5 cm in diameter, lacking cortical layers,

irregularly delimited; algae layer continuous, algal cells green, ovoid or ellipsoid,

single, 5-7.5 x 3.75-5 um.

APOTHECIA pale reddish brown to brownish, round and _ plump,

0.3-0.5mm in diameter, short-stiped, without clear margin, scattered over the

thallus; podetia whitish, 0.1-0.5 mm tall, 0.5 mm in diameter, lacking algae;

hymenium 112.5-125 um thick, I-; paraphyses simple, non-septated; asci long-

clavate, 8-spored, with apex I-, 80-87.5(-92.5) x 7.5 um; ascospores oblong or

fusiform, hyaline, one-septate, 10-12.5 x 5 um.

CHEMISTRY: Spot tests: Thallus K+ yellow turning red, C-, KC-, P+ yellow.

All specimens contain norstictic acid (TLC).

ADDITIONAL MATERIAL EXAMINED: CHINA, HAINAN: CHANGJIANG CouNTY, Mt.

Bawangling, 19°16’ N 109°03’ E, alt. 300 m, on rock, 25 November 2010, S.N. Cao

CSN048 (HMAS-L 118096, GenBank KC414618); CSN049 (HMAS-L 118089, GenBank

KC414616); CSN050 (HMAS-L 118087, GenBank KC414615); CSN051 (HMAS-L

118086, GenBank KC414614).

Phyllobaeis crustacea sp. nov. (China) ... 35

—.—

Fic. 2. Phyllobaeis crustacea (holotype): A, habit; B, apothecia; C, ascus with eight ascospores,

apex I- (arrow). Scale bars: A, B = 0.5 mm; C = 10 um.

Acknowledgments

We are very grateful to Dr. Harrie Sipman and Dr. Robert Lticking for reviewing the

manuscript. Our thanks are also given to Drs. Shaun Pennycook, Wen-Ying Zhuang,

and Klaus Kalb for valuable and constructive comments on previous versions of this

paper and providing an important literature, and to the curator of L who kindly sent

36 ... Cao & al.

a specimen on loan. Special thanks are due to Ms. H. Deng for giving considerable

assistance during the studies in HMAS-L. This research was supported by the Ministry

of Science and Technology of PRC (2006FY120100) and the Chinese Arctic and

Antarctic Administration (2011GW12016).

Literature cited

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

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

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/10.1016/0021-9673(72)80013-X

Gierl C, Kalb K. 1993. Die Flechtengattung Dibaeis. Eine Ubersicht iiber die rosafriichtigen Arten

von Baeomyces sens. lat. nebst Anmerkungen zu Phyllobaeis gen. nov. Herzogia 9: 593-645.

Index Fungorum. 2013. http://www.indexfungorum.org/Names/Names.asp

Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. 2011. MEGA5: Molecular

evolutionary genetics analysis using maximum likelihood, evolutionary distance, and

maximum parsimony methods. Molecular Biology and Evolution 28: 2731-2739.

http://dx.doi.org/10.1093/molbev/msr121

Thompson JD, Higgins DG, Gibson TJ. 1994. CLUSTAL W: improving the sensitivity

of progressive multiple sequence alignment through sequence weighting, position-

specific gap penalties and weight matrix choice. Nucleic Acids Research. 22: 4673-4680.

http://dx.doi.org/10.1093/nar/22.22.4673

Wang YY, Zhang T, Zhou QM, Wei JC. 2011. Construction and characterization of a

full-length cDNA library from mycobiont of Endocarpon pusillum (lichen-forming

Ascomycota). World Journal of Microbiology and Biotechnology. 27: 2873-2884.

http://dx.doi.org/10.1007/s11274-011-0768-5

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

lichen substances. British Lichen Society Bulletin 57(Suppl.): 1-41.

White TJ, Bruns TD, Lee SB, Taylor JW. 1990. Amplification and direct sequencing of fungal

ribosomal RNA genes for phylogenetics. 315-222, in: MA Innis et al. (eds). PCR protocols: a

guide to methods and applications. Academic Press, New York.

MY COTAXON

http://dx.doi.org/10.5248/126.37

Volume 126, pp. 37-44 October-December 2013

Nipponoparmelia perplicata sp. nov. (Parmeliaceae, Ascomycota)

from eastern Asia

SERGI YA. KONDRATYUK”™, SVETLANA I. TSCHABANENKO?’,

JoHN A. EL1x3, SOON-OK OH*4, ARNE THELL® & JAE-SEOUN HuR?’

'M.H. Kholodny Institute of Botany of National Academy of Sciences of Ukraine, Kiev, Ukraine

Sakhalin Botanical Garden, P.O. Box 34, Gorkogo Str. 25, Yuzhno-Sakhalinsk, 693023, Russia

’Research School of Chemistry, Building 137, Australian National University,

Canberra, ACT 0200, Australia

‘Korean Lichen Research Institute, Sunchon National University,

315 Mae-gok dong, Sunchon, 540-742, Korea

SBotanical Museum, Lund University, Box 117, SE-22100 Lund, Sweden

* CORRESPONDENCE TO: ksya_net@ukr.net

ABSTRACT — A new species, Nipponoparmelia perplicata described from Korea and Russia,

is segregated from N. laevior, from which it differs in having shorter and wider lobes that

characteristically turn down towards the margins. The pseudocyphellae are inconspicuous,

positioned along the lobe margins or on the underside. An improved description of

N. pseudolaevior is presented, including new data on isidia with illustrations. A key to Korean

Nipponoparmelia species is provided.

KEY worps — taxonomy, thallus morphology, distribution, ecology, secondary lobules

Introduction

The genus Nipponoparmelia (Kurok.) K.H. Moon et al. forms a separate

main clade in the Parmeliaceae (Crespo et al. 2010), which includes a group

of East Asian species previously placed in Parmelia Ach. s. str. According to

Kurokawa (1994), who treated the group as a subgenus of three species within

Parmelia, they differ morphologically from other Parmelia species by forming

lateral punctiform pseudocyphellae. The pseudocyphellae resemble those of

Punctelia Krog and Flavopunctelia (Krog) Hale in lacking a persistent epicortex

roof and not forming angular or linear pore aggregates (Krog 1982; Hale 1984).

However, in Nipponoparmelia the pseudocyphellae are marginal on lobes

and lobules, not laminal as in Punctelia and Flavopunctelia. Parmelia subg.

Nipponoparmelia Kurok. was recognized by a grey to grey-brown thallus with

marginal punctiform pseudocyphellae and simple to furcate rhizines. Crespo

38 ... Kondratyuk & al.

et al. (2010), who raised the subgenus to generic rank, included four species:

Nipponoparmelia isidioclada (Vain.) K.H. Moon et al., N. laevior (Nyl.) K.H.

Moon et al., N. pseudolaevior, and N. ricasolioides (Nyl.) A. Crespo & Divakar.

We have discovered a new Nipponoparmelia species, which we describe here

as N. perplicata. We also present an improved description of N. pseudolaevior

and provide a key to Korean Nipponoparmelia species.

Taxonomy

Nipponoparmelia perplicata S.Y. Kondr., Tschab., Elix & Hur, sp. nov. Figs 1, 2

MycoBAank 806127

Differs from Nipponoparmelia laevior by its smaller thalline lobes, helmet-shaped

marginal lobes, larger and mainly inconspicuous pseudocyphellae on the upper side,

and wider ascospores.

Type — South Korea: Mt. Jiri, 35°19’30.0°N 127°42’45.7”E, 1755 m, on bark of Malus

baccata (L.) Borkh. (Rosaceae), 23.iv.2004, J.-S. Hur 040372 (Holotype, KoLRI 001148).

EtryMoLocy — The species epithet refers to the inconspicuous pseudocyphellae.

Thallus 5-8(-10) cm broad, adnate to loosely adnate, dark greenish, olive-

grey to whitish mineral gray; lobes to 4(-7) mm long and 1-2(-3) mm

wide, subirregular, somewhat subconvex to convex due to the downwards

folded edges, contiguous to subimbricate, non-ascending and non-sinuate,

without isidia and soredia, upper surface matt to shiny, rarely foveolate, upper

surface mainly entire even on older lobes, pseudocyphellae from punctiform

and rounded or elongate to fissure-like to 0.6-1(-1.5) mm in diam., usually

inconspicuous, usually marginal but rarely laminal on the underside; medulla

white; lower surface black, often with pale or dark brownish marginal zone,

moderately to densely rhizinate, the rhizines simple or furcate.

Apothecia sometimes abundant, more or less stipitate, 2-7 mm in diameter,

thalline margin more or less foveolate, pseudocyphellae punctiform, highly

elevated, pseudocyphellae often present on the inner side of the thalline

margin, disc brown, reddish brown to dark brown. Ascospores (10—)13-17 x

(7-)8-11(-12) um.

CHEMISTRY— ‘Thallus K+ yellow becoming brownish, C-, Pd+ yellow;

medullar K+ yellow becoming red, C-, Pd+ deep yellow, slowly becoming

orange. Atranorin, chloroatranorin, salazinic acid, and traces of consalazinic

and protocetraric acid.

ECOLOGY & DISTRIBUTION— On trunk or twigs of trees. Known from

scattered localities in South Korea and far eastern Russia.

OTHER SPECIMENS EXAMINED — SOUTH KOREA: JEOLLANAM-DO, Gurye-gun, Jiri

Mt., 35°1749.5"N 127°33’36.2”E, 1281 m, on bark of Quercus, 27.ix.2006, J.-S. Hur

20060783 (KoLRI 005162); Mt. Jiri, 35°18’56.0”N 127°41’29.7”E, 1690 m, on bark of

Quercus, 23.iv.2004, J.-S. Hur 040354 (KoLRI 001130); KANGwon-Do, Shokcho city,

Nipponoparmelia perplicata sp. nov. (Russia & South Korea) ... 39

an a Ee

Doe BA wee

Fic. 1. Nipponoparmelia perplicata: general habit. Scale bars: a-d = 2 mm; e-f = 1 mm.

Seorak-dong, Mt. Sorak, 38°09’47.3”N 128°27'32.2”E, 810 m, on bark, 17.vi.2005, J.-S.

Hur 050284 (KoLRI 003188); JEOLLABUK-DO, Jeongup-si, Mt. Naejang, 35°28’57.8”N

126°52’59.6”E, 711 m, on bark of Quercus, 8.viii.2003, J.-S. Hur 030621 (KoLRI

000428); GYEONGSANGNAM-DO, Hapcheon-gun, Gaya-myeon, Mt. Gaya, 35°49’05.6”N

128°07’33.2”E, 1340 m, on twigs of Rhododendron, 5.v.2006, J.-S. Hur 20060152

(KoLRI 004524); GANGWon-Do, Seon-gun, Gohan-eup, Mt. Hambaek, 37°11’47.4”’N

128°54’53.6”E, 1355 m, on bark of Quercus, 19.vi.2007, J.-S. Hur 070652 (KoLRI 007511),

Mt. Hambaek, 37°11’43.0”N 128°54’51.6”E, 1403 m, on bark of Quercus, 19.vi.2007, J.-

S. Hur 070661 (KoLRI 007534); [Mt. Suk Byeng], 37°34’29.9’N 128°51’21.8”E, 686 m,

on bark, growing together with Flavoparmelia caperata, 24.v.2008, J.-S. Hur 080187

(KoLRI 008433, sub Flavoparmelia caperata); Mt. Deogyo, 35°51'15.8”N 127°44’55.2”E,

1601 m, on bark of Quercus, growing together with Biatora sp., 10.viii.2006, J.-S. Hur

20060457 (KoLRI 004825). RUSSIA: SHIKOTAN ISLAND, near mountain Notoro, on

AO ... Kondratyuk & al.

Fic. 2. Nipponoparmelia perplicata: general view of pseudocyphellae on underside.

Scale bars: a-d, f= 1 mm; e = 2 mm.

Betula ermanii, 43°47’21.1”N 146°42’26.7”E, 229 m alt., 14.viii.2010, S. Tschabanenko

(SAKH).

TAXONOMIC NOTES — Nipponoparmelia perplicata is distinguished from the four

other Nipponoparmelia species by the irregular lobes with the characteristically

down-turned lobe ends that produce a helmet-shaped impression. The

pseudocyphellae of N. perplicata are inconspicuous, seen mainly on the

underside where they are marginally (rarely laminally) positioned, and the lack

of marginal lobules and isidia.

Nipponoparmelia isidioclada is densely supplied with branched cylindrical,

slightly dorsiventral, eventually 1 mm high coralloid-isidioid structures. It

further differs from N. perplicata by smaller pseudocyphellae and presence of

gyrophoric and 4-O-methylgyrophoric acids (Park 1990; Kurokawa 1994).

Nipponoparmelia laevior, one of the commonest lichens in montane and

subalpine areas in Japan and Korea, is characterized by marginal punctiform

pseudocyphellae, a shiny and smooth upper surface, and simple to furcate

Nipponoparmelia perplicata sp. nov. (Russia & South Korea) ... 41

rhizines and sublinear lobes with distinctly ascending edges. Furthermore, the

ascospores of N. laevior are 12-16 x 6-9 um, slightly narrower than those of

N. perplicata (Park 1990; Kurokawa 1994).

Nipponoparmelia pseudolaevior differs from N. perplicata by numerous

subascending to suberect isidia along the margin and on the surface of the

lobes (Park 1990; Kurokawa 1994).

Similar to N. perplicata, Cetreliopsis asahinae (M. Sat6) Randlane & A. Thell

possesses pseudocyphellae on the underside. However, C. asahinae also

produces conspicuous pseudocyphellae on the upperside (Randlane & ‘Thell

1995). The lobes of N. perplicata are smaller and much more closely attached

and the lower surface is smoother. Furthermore, C. asahinae is usually supplied

with marginal apothecia and the secondary chemistry is characterized by

fumarprotocetraric acid.

Nipponoparmelia pseudolaevior (Asahina) K.H. Moon, Y. Ohmura & Kashiw.,

in Crespo et al., Taxon 59(6): 1749 (2010). Fic. 3

Thallus to 10 cm across, adnate or subadnate, dull brownish or dull greenish

grey; lobes subirregular, continuous or more or less divaricate, 1-3 mm wide,

with more or less ascending margins, upper surface shiny and smooth towards

the tips and matt and somewhat wrinkled in the centre, usually with numerous

laminal secondary lobules and numerous finger like to cylindrical isidia in

the centre of thallus; rarely secondary lobules and isidia with white pruinae

in places, pseudocyphellae rather small and inconspicuous, 0.1-0.2(-0.3) mm

in diam./across, mainly along the margins, sometimes becoming isidiate (not

becoming granular and not giving rise to coralloid-isidioid growths). Secondary

lobules 1-2.5(-3) mm wide/across, from rounded to irregular, with distinctly

attenuated basis, laminal. Isidia 0.7-1.5(-2) mm long, finger-like to cylindrical,

0.1-0.3 mm wide, scarcely branched, usually numerous along the edges of the

main lobes and the edges of the secondary lobules, especially abundant in the

centre of the thallus (not being apically crumbling and subsorediate with age).

Upper surface shiny, pruinose in parts, plane to slightly wrinkled, lobe edges

pseudocyphellate; pores more or less punctuate. Medulla white. Lower surface

black, often with a pale of brownish narrow peripheral zone, moderately

to densely rhizinate, the rhizines simple or furcated, to 1.5(-2) mm long.

Apothecia rare, more or less stipitate, 1-8(-11) mm diam., dentate or lobate

along the margin, amphithecium smooth, pseudocyphellate, pseudocyphellae

punctuate, more or less elevated, disc buff brown, radially split with age, spores

10-12 x 15-17 um. Pycnidia common, submarginal.

CHEMISTRY — Thallus K+ yellow, C-, KC-, P+ yellow; medulla K+ yellow,

becoming red or brown, C-, P+ deep yellow, than slowly becoming orange.

Atranorin, chloratranorin, and salazinic acid; consalazinic acid, a fatty acid and

protocetraric acid as accessories.

42 ... Kondratyuk & al.

Fic. 3. Nipponoparmelia pseudolaevior: A. general habit; B. central portion of thallus;

C. lobe; D, laminal lobules; E-F. isidia. Scale bars: A, B = 2 mm; C-F = 1 mm.

SPECIMENS EXAMINED — SOUTH KOREA: Mt. Taebaek, 1016 m alt, on rock, 2.xi.2003,

J.-S. Hur 030888 (KoLRI 000678); Mt. Baegun, 1105 m alt, 5.v.2003, J.-S. Hur 030188

(KoLRI 000143). RUSSIA: SAKHALIN IsLAND, Kholmsky district, Slepikovsky cape,

47°17'59.33”N 141°59’37.7”E, mix forest (Abies + Quercus + Sorbus), on Quercus

crispula, 2006, S. Tschabanenko (SAKH); Uglegorsky district, Bolshoi Nadym River,

49°00'54.06”N 142°15’13.01”E, mix forest, on Sorbus commixta, 2003, S. Tschabanenko

(SAKH); SHIKOTAN ISLAND, 43°47’21.1”N 146°42’26.7”E, mix forest, Gorobets

River, on Alnus hirsuta, 14.viii.2010, S. Tschabanenko (SAKH); Irurup IsLAND, near

Lesozavodskoye, mix forest, on Padus ssiori [date and collector unrecorded] (SAKH);

PRYMORSKY KRAI, Lazovsky district, valley Proselochnaya River, 43°00’48.81”N

134°07'00.58”E, deciduous-coniferous forest, on Quercus mongolica and Alnus

japonica, 1985, S. Tschabanenko (SAKH); Lazovsky district, valley Valunovka River,

43°08’45.37”N 133°48’25.75”E, deciduous-coniferous forest, on Quercus mongolica,

1986, S. Tschabanenko (SAKH); Lazovsky district, Quercus forest near maritime coast

Nipponoparmelia perplicata sp. nov. (Russia & South Korea) ... 43

(Zapovednoye village), 42°50’25.74”"N 133°42’02.6”E, on Quercus mongolica and

Q. dentata, 1985, S. Tschabanenko (SAKH); Lazovsky district, Quercus forest near

maritime coast Krasnyi Olenevod, 42°52’19.1”N 133°46’51.46”E, on Quercus mongolica

and Q. dentata, 1985, S. Tschabanenko (SAKH); Ussurijsky district, Zharikov Stream,

deciduous-—coniferous forest, on Alnus hirsuta, [date unrecorded] S. Skirina (SAKH);

Khasansky district, valley Kedrovaya River, 43°06’10.64”N 131°31’48.75”E deciduous-

deciduous forest, on Quercus mongolica, [date unmentioned], S. Skirina (SAKH);

KHABAROVSKY KRAI, valley Gorin River, mix forest, on Quercus mongolica, [date

unrecorded], A. Mikulin (SAKH).

EcoLoGcy & DisTRIBUTION— On bark of Cryptomeria japonica and various

deciduous trees, rarely on Tsuga diversifolia, Abies veitchii, and A. sachalinensis

and on rocks, growing sometimes together with Menegazzia subsimilis and

Parmelia and Lepraria species. Known from eastern Asia (Japan, South Korea,

far eastern Russia). Known in South Korea only from two localities, Mt. Taebaek

and Mt. Baegun.

TAXONOMIC NOTES — Nipponoparmelia pseudolaevior is closely related to

N. laevior, from which it is clearly distinguished by the numerous subascending

to suberect isidia formed on the margin and less commonly present on the

surface of lobes. It is apparently the isidiate morphotype of N. laevior, which

has a similar distribution range in eastern Asia.

Nipponoparmelia pseudolaevior differs from N. isidioclada, a mainly

epiphytic species known from Russian Far East, Japan, and Taiwan, by presence

of secondary lobules and larger pseudocyphellae, longer isidia, and absence of

gyrophoric acid.

Some Korean specimens of N. pseudolaevior were previously misidentified

as Punctelia rudecta (Ach.) Krog, from which it differs in having only marginal

punctiform pseudocyphellae, laminal secondary lobes with numerous marginal

isidia and a black underside.

In contrast to Russian literature (Rassadina 1971), true isidia were not

mentioned for N. pseudolaevior in Korean and Japanese keys, being categorized

as secondary lobules (Park 1993; Kurokava 1994). However, isidiate material

from South Korea and Russia does represent N. pseudolaevior.

Key to Eastern Asian Nipponoparmelia species

1. Thallus isidiate, with coralloid-isidioid growths or with dense suberect, marginal

Cylindrical isidiae ssa. t.. went. Barhhe rn Beek andsghi tases oR gs oe aee,.eR as meant ae 2

IR: W MSIGarASCHIES tivasres, hoehag Dorktes tortie Doraliby oral hcwk ty sch te ae suk Ae eisek A moans seas 3

2. Isidia apically crumbling and subsorediate; pseudocyphellae becoming

granular-isidiate and giving rise to coralloid-isidioid growths to 1 mm high

penta My tgubad tial tris 4. trie gcbdl bye Mak dy Js gy Pet g Nipponoparmelia isidioclada

2. Isidia not sorediate, regularly cylindrical, dense and suberect along edges of

Tepitlescs x, octet tetecteg gig) gd seh gdrtyg e8dceat adhe Nipponoparmelia pseudolaevior

44 ... Kondratyuk & al.

3. Lobes sublinear, with ascending margins; pseudocyphellae well distinct,

conspicuous from upper surface.................04. Nipponoparmelia laevior

3. Lobes irregular, margin folded downwards; pseudocyphellae indistinct,

befterseen from Underside... oe, mass » tarcte « alben'e Nipponoparmelia perplicata

The status of N. laevior specimens with secondary lobules but without isidia

will be investigated in the future.

Acknowledgements

SK expresses his deep thanks to Prof. J.-S. Hur and Dr. S.-O. Oh for permission to

work with their Korean and Chinese collections. We would like to express our deep

thanks to Dr. A. Thell (Lund, Sweden) for kind help with some literature and herbarium

specimens. Authors are grateful to Profs. Ingvar Karnefelt and Pradeep K. Divakar

for their valuable comments on manuscript. This work was supported in parts by The

State Agency on Science, Innovations and Information of Ukraine (M317-2011-409,

M111-2012-409 and M40-2013-409) and the Korean Forest Service Program (KNA

2013) through the Korea National Arboretum and the Korean National Research

Resource Center Program (NRF, 2012M3A9B8021726).

Literature cited

Crespo A, Kauff F, Divakar PK, del Prado R, et al. 2010. Phylogenetic generic classification of

parmelioid lichens (Parmeliaceae, Ascomycota) based on molecular, morphological and

chemical evidence. Taxon 59(6): 1735-1753.

Hale ME. 1984. Flavopunctelia, a new genus in the Parmeliaceae (Ascomycotina). Mycotaxon 20:

681-682.

Krog H. 1982. Punctelia, a new lichen genus in the Parmeliaceae. Nordic Journal of Botany 2:

287-292. http://dx.doi.org/10.1111/j.1756-1051.1982.tb01191.x

Kurokawa S. 1994: Japanese species of Parmelia Ach. (sens. str.), Parmeliaceae (2). Journal Japanese

Botany 69(3): 121-126.

Park YS. 1990. The macrolichen flora of South Korea. The Bryologist 93(2): 105-160. http://dx.doi.

org/10.2307/3243619

Randlane T, Thell A, Saag A. 1995. New data about the genera Cetrariopsis, Cetreliopsis and

Nephromopsis (fam. Parmeliaceae, lichenized Ascomycotina). Cryptogamie, Bryologie-

Lichénologie 16(1): 35-60.

Rassadina KA. 1971. Family Parmeliaceae. 282-386, in: II Abramov (ed.). Handbook of the Lichens

of the U.S.S.R. 1. Pertusariaceae, Lecanoraceae and Parmeliaceae. Leningrad: Nauka, Leningrad

division.

MYCOTAXON

http://dx.doi.org/10.5248/126.45

Volume 126, pp. 45-50 October-December 2013

Buelliella, Codonmyces, and Polycoccum species new to Turkey

KENAN YAZICI*'& JAVIER ETAYO?

‘Biology Department, Faculty of Science, Karadeniz Technical University, 61080, Trabzon, Turkey

?Navarro Villoslada 16, 3° dcha, 31003 Pamplona, Navarra, Spain

*CORRESPONDENCE TO kcagri_1997@yahoo.com

ABSTRACT — Three species of lichenicolous fungi are reported as new to Turkey: Buelliella

poetschii and Codonmyces lecanorae (both also new to Asia) and Polycoccum pulvinatum.

Short descriptions are presented, including geographic distribution, host, and a comparison

with similar taxa.

Key worps — Ascomycetes, biodiversity, coelomycetes, Burdur, Dothideales

Introduction

Although studies on the lichenicolous fungi of Turkey are insufficient and

not intensive, there have been an increasing number of publications on the

subject during 2009-11 (Candan & Halici 2009, 2011, Etayo & Yazici 2009,

Halici et al. 2010a,b, Yazici et al. 2010, 2011). Recently, during our study of the

lichens and bryophytes of Burdur Province, we identified some lichenicolous

fungi. Although 139 taxa of lichenicolous fungi are known from other regions

of Turkey (Yazici et al. 2011), no lichenicolous fungi have been reported from

Burdur.

This number is small when compared with other European countries

(Faltynowicz 2003, Hawksworth 2003, Kocourkova 2000, Santesson 1993,

Scholz 2000). Thus exploration of more regions of Turkey is urgently needed.

Materials & methods

The present report is based on collections from the Burdur region made on 25 and

29 June 2012. Burdur has a continental Mediterranean climate with cold, snowy winters

and very hot, long and dry summers. The temperature ranges from -16 to 39°C with a

mean of 15°C; the annual rainfall averages 468 mm, and the average humidity is 51.2%

(Akman 1999).

The visited areas (Bucak and Altinyayla districts) are mountainous with vast forested

areas dominated by Abies, Cedrus, Ficus, Fraxinus, Juniperus, Olea, Pinus, Pistacia,

Prunus, Quercus (especially in Altinyayla district), and Rhus, and alternating streams,

46 ... Yazici & Etayo

lakes, and dams (e.g., Yaprakli Dam in Altinyayla district, and Karaca6ren dam in Bucak

district) (Baytop & Denizci 1963). Within the forests, other flora is very abundant, and

marble rock is common.

Hand-cut sections were examined microscopically in water (including all

measurements), 10% KOH, and lactophenol cotton blue. Air-dried samples were

observed using a Nikon SMZ1500 stereomicroscope and a Nikon Eclipse 80i light

microscope. Nomenclature and species concepts follow Hawksworth & Diederich

1988, Pegler et al. 1980, Hafellner et al. 2008, Calatayud & Etayo 1999, and Etayo 2010a.

Vouchers are stored in the herbarium of the Biology Department, Faculty of Science,

Karadeniz Technical University, Trabzon, Turkey (KTUB). The descriptions are based

on Turkish specimens.

The species

Fic. 1. Buelliella poetschii, habitus with apothecioid ascomata. Scale = 1mm

Buelliella poetschii Hafellner, Mitt. Naturw. Ver. Steierm. 137: 187. 2008 Fic. 1

Detailed descriptions: Hafellner et al. (2008); Etayo (2010a).

Ascomata apothecioid, dispersed over the host thallus in black spots with

1-4 tiny composite groups of apothecia, rarely single, rounded, globular when

young, pressed at the edges, later oval, up to 120-1200 um in diam., often

bleaching the host thallus, occasionally sunken. Vegetative hyphae intercellular.

Hymenium hyaline to brownish, on the outside olive-brown, c. 60-70 um high,

I-, KI-. Excipulum dark brown, K+ greenish-brown. Paraphyses branched,

anastomosing, capitate at apical and intercalary cells with pigment, cells at

tips 4-5 um thick with dark-brown pigment. Asci cylindrical, fissitunicate,

Lichenicolous fungi new to Turkey ... 47

4-6-spored. Ascospores, long, hyaline, later + brown, thin walled, 1-septate,

clearly constricted at the septa, cells rounded, the upper one sometimes tapered,

(15-)16-16.3-18.5(-20) x 6.5-7.2-8.5 um (n = 15).

SPECIMEN EXAMINED: TURKEY. Burpur: Bucak, surroundings of Karacaéren Dam,

37°21'35.14”N 30°50’05.80”E, 300 m, on the thallus of Endocarpon sp., 25.06.2012, leg.

K.Yazici (KTUB-2353).

Our specimen was found on an Endocarpon thallus, which appeared damaged

by this fungus. Buelliella poetschii has previously been found growing on the

thalli of Endocarpon and Catapyrenium species (E. adscendens, E. adsurgens,

E. loscosii, E. pusillum agg., Catapyrenium sp.) on exposed calcareous rocks.

The species was previously known from Europe, North America, and

South America (Hafellner et al. 2008, Diederich et al. 2009, Etayo 2010a,b,

Urbanavichus & Urbanavichene 2011, Flakus & Kukwa 2012). New to Turkey

and Asia.

Fic. 2. Codonmyces lecanorae, habitus with apothecia. Scale = 1mm

Codonmyces lecanorae Calat. & Etayo, Lichenologist 31(6): 594. 1999 Fic. 2

Detailed description: Calatayud & Etayo (1999).

Vegetative hyphae spanning fully or partly the apothecia and thallus of the

host lichen. Conidiophores + cylindrical, hyaline, and immersed in the host

tissue, up to 3-4 um wide. Cells near the surface of the host tissue + globular

and 6-7 um wide, colourless or + pale brown. Conidiogenous cells formed

by a globose cell with annellations, globose cells with + smooth and + dark

brown surfaces, annellations + dark brown, (8-)9-11(-12) x (4-)5-8(-9) um

48 ... Yazici & Etayo

(including the annellations). Conidia 1-septate, brown, oblong with rounded

apices, constricted at the septum, (6-)7-11(-12) x (4-)5-6 um (n = 20).

SPECIMEN EXAMINED: TURKEY. Burpur: AttTinyayia, between I[becik-Altinyayla

road, 36°56'59.32”N 29°27'40.68”E, 1373 m, on the thallus and apothecia of

Protoparmeliopsis muralis, 29.06.2012, leg. K.Yazici (KTUB-2345).

Our specimen growing on the apothecia and thallus of Protoparmeliopsis

muralis was not deformed in the affected parts, although the host was

moderately damaged. Codonmyces lecanorae, previously known from Europe,

is new to Turkey and Asia.

REMARKS — Codonmyces is similar to Xanthoriicola (Hawksworth &

Punithalingam 1973, Hawksworth, 1979) in producing effuse powdery colonies

lacking stromatic structures and sporodochia. The morphologically similar

Xanthoriicola physciae is distinguished by simple conidia that are not 1-septate.

Additionally warts in Xanthoriicola are typically spinulose (Hawksworth 1979,

Ellis 1976), and not flattened as in Codonmyces (Calatayud & Etayo 1999).

Fi. 3. Polycoccum pulvinatum, habitus with galls, including perithecioid ascomata.

Scale = 1mm

Polycoccum pulvinatum (Eitner) R. Sant., Lich. Lichenic. Fungi Sweden Norway:

175..1993 Fic. 3

Detailed description: Hawksworth (1975).

Galls wart-like, on Physcia sp., convex or hemispherical, 1-2.5 mm in diam.,

ascomata perithecioid (pseudothecium), 150-290 x 140-200 um, subglobose,

arising singly, walls of angular pseudoparenchymatous cells, hymenium

Lichenicolous fungi new to Turkey ... 49

not exposed at maturity; hamathecium of branched and anastomosing

pseudoparaphyses. Asci 60-80 x 14-18 um, cylindrical-clavate, (4—)8-spored;

ascospores l-septate, 14-20 x 6-8(-9) um (n = 18), ellipsoid to oblong-

ellipsoid, brown at maturity, perispore present.

SPECIMEN EXAMINED: TURKEY. BurpDuR: ALTINYAYLA, between Ibecik-Altinyayla

road, 36°56’59.32”N 29°27’40.68”E, 1373 m, on the thallus of Physcia sp., 29.06.2012,

leg. K. Yazici (KTUB-2352).

Widespread in Mediterranean to boreal and arctic-alpine areas, P pulvinatum

induces the gall formation and grows on Physcia spp. (e.g., P adscendens,

P. caesia, P. dubia, P. tenella). The species is known from all continents except

Africa and is widely distributed in Europe (Alstrup et al. 2009, Hawksworth

1983, Pegler et al. 1980). New to Turkey.

REMARKS— Polycoccum pulvinatum resembles P. clauzadei, which differs in

its smaller galls and ascomata. Additionally, its ellipsoid or oblong-ellipsoid

ascospores and usually 8-spored asci distinguish P pulvinatum from P clauzadei

with obovoid or soleiform ascospores and 4-6-spored asci (Navarro-Rosinés &

Roux 1998).

Acknowledgements

We are grateful to Dr. Ave Suija and Dr. Wolfgang von Brackel for revisions and

helpful comments on an earlier draft of this manuscript. This study was supported by

TUBITAK (Project 111T857).

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MY COTAXON

http://dx.doi.org/10.5248/126.51

Volume 126, pp. 51-54 October-December 2013

Passalora lepistemonis sp. nov. from China

LEI X1A', YING-LAN GUO? & Yu LI’ *

' Engineering Research Center, Chinese Ministry of Education for Edible and Medicinal Fungi,

Jilin Agricultural University, 2888 Xincheng Street, Changchun City, PR. China

? State Key Laboratory of Mycology, Institute of Microbiology Chinese Academy of Sciences,

Beijing 100101, RR. China

* CORRESPONDENCE TO: yuli966@126.com

ABSTRACT — A new species of Passalora, P. lepistemoni, is described and illustrated. The type

specimen is deposited in HMAS.

KEY worDs — taxonomy, microfungi, anamorphic fungi, tropical fungi, plant pathogen

Introduction

More than 600 species have been reported from Passalora (teleomorph

Mycosphaerella) (Kirk et al. 2008), a hyphomycete genus established by Fries

in 1849. Passalora species are common parasites of a large number of plants,

often causing spots on leaves, stems, flowers, fruits, and seeds resulting in

serious economic damage due to decreased plant production. The primary

morphological generic characters include pale olivaceous to almost black

stromata (when present); solitary to fasciculate pale olivaceous to dark brown

conidiophores; obvious conidial scars; solitary conidia that are variably shaped

(straight to curved, cylindrical, clavate, fusiform), hyaline to dark olivaceous

brown, nonseptate to multiseptate, and with an obvious hilum.

The authors found a new species from diseased leaves of Lepistemon

binectariferum, collected in Hainan Province, China. The species is described

and illustrated here.

Taxonomy

Passalora lepistemonis L. Xia, Y.L. Guo & Yu Li, sp. nov. FIG. 1

MycoBank MB802399

Differs from other Passalora species reported on convolvulaceous hosts by its paler and

shorter conidiophores, its conidial size and shape, and its lack of stromata.

52 ... Xia, Gao & Li

Fic. 1. Passalora lepistemonis (HMAS 244085, holotype).

1. Conidiophores; 2. Conidia. Bar = 25 um.

TypE—China, Hainan Province, Bawangling, from leafspots on Lepistemon

binectariferum (Wall.) Kuntze (Convolvulaceae), 11 XII 2009, coll. Guo Ying-lan,

(Holotype HMAS 244085).

ErymMo.oey: Named after the host genus Lepistemon.

LEAF spots amphigenous, angular, without definite margin, vein-limited, 1.5-5

mm wide, often confluent, at first pale green, later pale to medium yellowish

brown on the upper surface, laurel-green to pale yellowish brown on the

lower surface. CAESPITULI hypophyllous, effuse. PRIMARY mycelium internal;

secondary mycelium external, hyphae pale olivaceous to olivaceous, branched,

septate, smooth, 2-2.5 um wide. STROMATA absent. CONIDIOPHORES emerging

through the stomata, acrogenous or as lateral branches singly arising from the

external mycelia, olivaceous to very pale olivaceous brown, uniform in color,

irregular in width, wider on the base, straight to slightly curved, unbranched,

0-l1geniculate, broadly obtuse to conical at the apex, usually not septate,

2.5-10.5 x 4-5.3 um, sometimes longer conidiophores 1-septate, up to 25 um

in length. CoNIDIAL scars conspicuously thickened, 1.5-2.8 um wide. CONIDIA

cylindrical or sometimes obclavate for shorter ones, olivaceous to very pale

olivaceous brown, catenulate and in branched chains, straight to curved,

conical or broad-round to truncate at the apex, obconically truncate at the base,

Passalora lepistemonis sp. nov. (China) ... 53

shorter conidia 0-2- septate, longer conidia 3-7- septate, constricted at septa,

25-110 x 4—-6.7 um.

Discussion

More than ten species have been reported on plants of Convolvulaceae, of

which five were similar to Passalora lepistemonis in producing ceratoid and

inconspicuous or irregular leaf spots, and effused hypophyllous caespituli:

P. balansae (Speg.) U. Braun on Evolvulus sp.; P. convolvuli (Tracy & Earle)

U. Braun & Crous on Convolvulus acetosifolia, C. arvensis, and Ipomoea sp.;

P. lettsomiae (Thirum. & Chupp) Crous & U. Braun on Argyreia spp.;

P. merremiae (X.J. Liu & Y.L. Guo) U. Braun & Crous on Merremia umbellata

subsp. orientalis; and P. turbinae (Chupp) U. Braun & Crous on Ipomoea

burmannii and Turbina corymbosa (Braun 2000; Crous & Braun 2001, 2003).

Passalora balansae produces darker (medium brown) longer (50-300 um)

conidiophores and paler (very pale olivaceous) shorter and wider (25.6-60 x

5-8 um) conidia (Braun 2000).

Passalora convolvuli has stromata (brown to almost black, 20-50 um diam),

densely fasciculate, darker (pale to medium olivaceous brown), longer (10-50

um) conidiophores, and paler (hyaline to subhyaline), obclavate-cylindrical,

narrower (3-4 um) conidia (Crous & Braun 2003).

Passalora lettsomiae produces darker (fuliginous to almost black) angular

leaf spots, fasciculate (5-20) conidiophores, and cylindrical-obclavate narrower

(3-4.5 um) conidia (Crous & Braun 2001).

Passalora merremiae has darker (pale brown), wider (5.5-10 um diam),

catenulate conidia, and darker (pale brown to brown) larger (12.5-108 x 5-8.8

um) conidiophores (Crous & Braun 2003).

Passalora turbinae produces small (25 um diam) reddish brown stromata,

densely clustered longer (15-55 um) paler 0-4 septate conidiophores, and

smaller (15-60 x 3-5 um) conidia (Crous & Braun 2003).

All these species differ from the fungus found on Lepistemon binectariferum,

described and illustrated above as a new species.

Acknowledgments

This work was supported by the fund of Florarum Cryptogamarum Sinicarum from

the Chinese Academy of Science (No.KSCX2-EW-Z-9). We are grateful to Dr. Guozhong

Lu and Dr. Tony Whalley for their reviews.

Literature cited

Braun U. 2000. Annotated list of Cercospora spp. described by C. Spegazzini. Schlechtendalia

BY 57-79.

Crous PW, Braun U. 2001. A reassessment of the Cercospora species described by C. Chupp:

specimens deposited at BPI, Maryland, U.S.A. Mycotaxon 78: 327-343

54 ... Xia, Gao & Li

Crous PW, Braun U. 2003. Mycosphaerella and its anamorphs: 1. Names published in Cercospora

and Passalora. CBS Biodiversity Series: 571p.

Kirk PM, Cannon PF, Minter DW, Stalpers JA. 2008. Ainsworth & Bisby’s dictionary of the fungi,

10th ed. CAB International, Wallingford, UK.

MY COTAXON

http://dx.doi.org/10.5248/126.55

Volume 126, pp. 55-60 October-December 2013

Two new species of Spadicoides from southern China

JI-WEN X1A, L1-Guo Ma, JIAN MA & XIU-GUO ZHANG

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

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

ABSTRACT — Spadicoides bawanglingensis sp. nov. and S. longchiensis sp. nov. are described

and illustrated from plant debris in natural areas of southern China. The type specimens are

deposited in HSAUP (Herbarium of Department of Plant Pathology, Shandong Agricultural

University) and HMAS (Mycological Herbarium, Institute of Microbiology, Chinese

Academy of Sciences).

KEY worps — anamorphic fungi, taxonomy

Introduction

Spadicoides was established by Hughes (1958) with S. bina (Corda) S. Hughes

as the type species. The genus is characterized by distinct, single, erect, straight

or flexuous, branched or unbranched conidiophores with polytretic, terminal

or intercalary conidiogenous cells that produce solitary, acropleurogenous,

euseptate, variously shaped conidia (Ellis 1971, Kuthubutheen & Nawawi 1991).

Thirty-four species have been accepted in Spadicoides (MycoBank 2013), seven

of which have been described from China (Zhou et al. 1999, Wong et al. 2002,

Ho et al. 2002, Cai et al. 2004, Li et al. 2010, Ma et al. 2010, 2012a).

Fungal diversity in China is high but still mostly unexplored. We have

previously described several new species from Hainan and Sichuan Province

(Zhang et al. 2009, Ma et al. 2012b,c, Ren et al. 2012). During our ongoing

survey of anamorphic fungi associated with plant debris in tropical and

subtropical forests of southern China, two new species with morphological

characteristics of Spadicoides were collected on dead stems.

Spadicoides bawanglingensis J.W. Xia & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 807061

Differs from all other Spadicoides species with obclavate conidia by its larger conidia

with a long rostrum (beak or apical prolongation).

56 ... Xia & al.

Fic. 1. Spadicoides bawanglingensis (holotype, HSAUP H6028). A. Conidiophores with conidia.

B. Conidiogenous loci (arrowed). C-E. Conidia.

Type: China, Hainan Province: tropical forest of Bawangling, on dead stems of an

unidentified broad-leaved tree, 16 Aug 2011, J.W. Xia (Holotype, HSAUP H6028;

isotype, HMAS 243424).

EryMo_oey: in reference to the type locality.

Spadicoides spp. nov. (China) ... 57

COLONIES on natural substrate effuse, dark brown, hairy. Mycelium partly

superficial, partly immersed in the substratum, composed of septate, pale

brown, smooth hyphae, 2-2.5 um wide. ConipiopHores differentiated,

mononematous, unbranched, erect, straight or slightly flexuous, cylindrical,

smooth, thick-walled, brown, 7-14-septate, 130-210 x 3.5-6 um.

CONIDIOGENOUS CELLS polytretic, integrated, terminal and _intercalary,

cylindrical, smooth, pale brown to brown. Conip1a acropleurogenous, solitary,

verrucose, 9-12-euseptate, obclavate, pale brown to brown, 98-166 um long

(rostrum included), 8.5-12.5 um wide, 2.5-4.5 um wide at the truncate base,

apex extended into a pale brown to subhyaline, euseptate rostrum, 52-100 x

1-3 um.

TABLE 1. Morphological comparison of Spadicoides species with obclavate conidia.*

CONIDIA

CONIDIOPHORES

SPECIES WALL; SIZE SEPTA APEX;

(um) corer (um) (#) BASE

S. bambusicola’ 150-350 x Verrucose; 30-72.5x 5-7 Subacute;

Meier ey iinet ee, aati ene.) one color ee eee obconically truncate

S. bawanglingensis 130-210x Verrucose; 98-166 x 9-12 Rostrate;

be ee ee, Fk 2 2:

S. heterocolorata' <420 x Smooth; 16-25 x Mostly Slightly tapering;

I coe G10 een eESiColored | 3.575. slightly protruding

S. obclavata’ <550 x Smooth; 16-22 x Mostly

5-8 (base), versicolored 4-6 2 Tapering; rounded

A ee, ee ESS alk LAs, fen coe RI oy Mil int Oa PR A i

S. palmicola' <780 x Smooth; 25-70 x Mostly — Subacute, often

7-15 (base), versicolored 5-7 4or5 rostrate;

4-5.5 (apex) obconically truncate

* Data from ' Goh & Hyde (1999), ? Kuthubutheen & Nawawi (1991), and ? Zhou et al. (1999).

Comments - The conidiophores and conidia morphologies of S. bawanglingensis

and four other Spadicoides species with obclavata conidia are compared in

TABLE 1. Spadicoides bambusicola D.Q. Zhou et al., S. heterocolorata (R.F.

Castaneda et al.) Goh & K.D. Hyde, S. obclavata Kuthub. & Nawawi, and

S. palmicola Goh & K.D. Hyde can all be separated from S. bawanglingensis by

their smaller conidia with fewer septa.

Spadicoides longchiensis J.W. Xia & X.G. Zhang, sp. nov. FIG. 2

MycoBank MB 807062

Differs from all other Spadicoides species by its oval to reniform conidia.

Type: China, Sichuan Province: national forest park of Longchi, on dead stems of

unidentified broad-leaved tree, 17 Apr 2012, J.W. Xia (Holotype, HSAUP H6246;

isotype, HMAS 243425).

EryMo_oey: in reference to the type locality.

Co.Lonigs effuse on natural substrate, dark brown, hairy. Mycelium partly

superficial, partly immersed. Hyphae branched, septate, pale brown, smooth.

58 ... Xia & al.

Fic. 2. Spadicoides longchiensis (holotype, HSAUP H6246). A. Conidiophore with conidia. B.

Conidiogenous cells, conidiogenous loci (arrowed) and conidia. C. Conidia. D. Conidiophore

basal cell.

Spadicoides spp. nov. (China) ... 59

CONIDIOPHORES macronematous, mononematous, single, unbranched,

erect, straight or slightly flexuous, cylindrical, smooth, thick-walled, brown,

9-14-septate, 200-300 x 5-8.5 tm. CONIDIOGENOUS CELLS polytretic,

integrated, terminal and intercalary, cylindrical, smooth, pale brown to brown.

ConlipiA acropleurogenous, solitary, rounded at both ends, oval to reniform,

2-(3)-euseptate, mostly 2-euseptate, subhyaline to pale brown, smooth, 16-23

um long, 6.5-8 um wide in the broadest part.

ComMENTS - Spadicoides longchiensis resembles S. klotzschii S. Hughes in

having smooth 2-septate conidia, but S. klotzschii differs by its smaller, clavate

to ellipsoidal, versicolored conidia (10-13 x 4-5.5 um; Hughes 1973).

Acknowledgments

The authors express gratitude to Dr. W.B. Kendrick and Dr. R.F. Castaneda Ruiz for

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

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

31093440, 31230001) and the Ministry of Science and Technology of the People’s

Republic of China (No. 2006FY 120100).

Literature cited

Cai L, McKenzie EHC, Hyde KD. 2004. New species of Cordana and Spadicoides from decaying

bamboo culms in China. Sydowia 56: 222-228.

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey,

England. 608 pp.

Goh TK, Hyde KD. 1999 [“1998”]. Spadicoides palmicola sp. nov. on Licuala sp. from Brunei, and a

note on Spadicoides heterocolorata comb. nov. Can. J. Bot. 76(10): 1698-1702.

Ho WH, Yanna, Hyde KD. 2002. Two new species of Spadicoides from Brunei and Hong Kong.

Mycologia 94: 302-306. http://dx.doi.org/10.2307/3761808

Hughes SJ. 1958. Revisiones hyphomycetum aliquot cum appendice de nominibus rejiciendis. Can.

J. Bot. 36: 727-836. http://dx.doi.org/10.1139/b58-067

Hughes SJ. 1973. Spadicoides klotzschii. Fungi Canadenses No 8. Ottawa, Canada.

Kuthubutheen AJ, Nawawi A. 1991. Two new species of Spadicoides from Malaysia. Mycol. Res. 95:

163-168. http://dx.doi.org/10.1016/S0953-7562(09)81006-X

Li DW, Chen JY, Wang YX. 2010. Two new species of dematiaceous hyphomycetes from Hubei,

China. Sydowia 62: 171-179.

Ma LG, 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

Ma LG, Ma J, Zhang YD, Zhang XG. 2012a. Spadicoides camelliae and Diplococcium livistonae, two

new hyphomycetes on dead branches from Fujian Province, China. Mycoscience 53: 25-30.

http://dx.doi.org/10.1007/s10267-011-0138-z

Ma J, Zhang YD, Ma LG, Ren SC, Castafieda Ruiz RF, Zhang XG. 2012b. Three new species of

Solicorynespora from Hainan, China. Mycol Progress 11: 639-645.

http://dx.doi.org/10.1007/s11557-011-0775-9

MaJ, Zhang YD, Ma LG, Castaneda Ruiz RF, Zhang XG. 2012c. Three new species of Sporidesmiella

from southern China. Mycoscience 53: 187-193. http://dx.doi.org/10.1007/s10267-011-0152-1

60 ... Xia & al.

MycoBank. 2013. MycoSearch database search. International Mycological Association,

http://www.mycobank.org/Biolomics.aspx? Table=Mycobank&Page=200&ViewMode=Basic

[ Accessed: 22 Nov. 2013]

Ren SC, Ma J, Ma LG, Zhang YD, Zhang XG. 2012. Sativumoides and Cladosporiopsis, two new

genera of hyphomycetes from China. Mycol Progress 11: 443-448.

http://dx.doi.org/10.1007/s11557-011-0759-9

Wong MKM, Goh TK, McKenzie EHC, Hyde KD. 2002. Fungi on grasses and sedges: Paratetraploa

exappendiculata gen. et sp. nov., Petrakia paracochinensis sp. nov. and Spadicoides versiseptatis

sp. nov. (dematiaceous hyphomycetes). Cryptog. Mycol. 23: 195-203.

Zhang K, Ma J, Wong Y, Zhang XG. 2009. Three new species of Piricaudiopsis from southern China.

Mycologia 101: 417-422. http://dx.doi.org/10.3852/08-147

Zhou DQ, Goh TK, Hyde KD, Vrijmoed LLP. 1999. A new species of Spadicoides and new records

of bambusicolous hyphomycetes from Hong Kong. Fungal Divers. 3: 179-185.

MY COTAXON

http://dx.doi.org/10.5248/126.61

Volume 126, pp. 61-70 October-December 2013

Entocybe haastii from Watagans National Park,

New South Wales, Australia

SARAH E. BERGEMANN ', DAvID L. LARGENT ”’,, & SANDRA E. ABELL-Davis?

‘Biology Department, Middle Tennessee State University,

PO Box 60, Murfreesboro TN 37132, United States

*Biological Sciences, Humboldt State University, 1 Harpst St, Arcata CA 95521, United States

School of Marine and Tropical Biology, Australian Tropical Herbarium and Centre

for Tropical Environmental & Sustainability Science, James Cook University,

PO Box 6811, Cairns QLD 4870 Australia

“CORRESPONDENCE TO: mrp@humboldtl.com

ABSTRACT — Entocybe haastii comb. nov. ( Entoloma haastii) is distinguished by isodiametric

minutely rounded pustulate-angular basidiospores, a dark blue black to nearly black pileus

that lacks brown tones, dark blue grey lamellae, an appressed fibrillose blackish blue stipe,

intracellular pigment in the pileipellis and inflated hyphae in the outer pileal trama, and the

faintly parietal pigment on narrow pileal tramal hyphae.

Key worps — Basidiomycota, Entolomataceae, new combination

Introduction

Entocybe T.J. Baroni et al. (Basidiomycota, Agaricales, Entolomataceae) was

erected to accommodate a monophyletic group of species with the following set

of morphological features: a slender tricholomatoid or mycenoid to collybioid

habit, a relatively fragile, appressed fibrillose stipe, small, obscurely angular

basidiospores with 6-10 facets (angles) in polar view and undulate-pustulate or

rounded pustulate surface ornamentation overall, and a pileipellis subtended

by inflated hyphae in the outer pileal trama (Baroni et al. 2011). An agaricoid

fungus with this character set recently collected in Watagans National Park in

central New South Wales was determined to be Entoloma haastii, a new report

for mainland Australia. Herein, we recombine E. haastii in Entocybe.

Materials & methods

Macromorphological and micromorphological features

Techniques and equipment for collecting and describing basidiomata in the field,

GPS coordinates, microscopy of dried collection including how basidiospores were

62 ... Bergemann, Largent & Abell-Davis

TABLE 1. Collections used in the phylogenetic analyses.

COLLECTION GENBANK ACCESSION NUMBERS

SPECIES FROM GENBANK IDENTIFIER mtSSU LSU RPB2

Calliderma sp. DLL 10025 JQ793648 JQ793642 =

DLL 10054 — JQ793649 —

DLL 10088 JQ793643 JQ793650 JQ793657

Clitocybe dealbata HC95cp AF357138 AF223175 DQ825407

Clitopilopsis hirneola 8490TJB GU384587 GU384611 GU384646

Cl. hirneola [Clitopilus hirneolus] 263 GQ289352 GQ289211 GQ289278

Clitopilus cystidiatus 26 GQ289287 GQ289147 GQ289220

C. prunulus 8229TJB — GU384615 GU384650

9663TJB GU384586 GU384614 GU384648

C. “cinerascens” 8024TJB GU384585 GU384613 GU384647

Entocybe haastii DLL 9868 JQ793644 JQ793651 JQ793658

DLL 10087 JQ793645 JQ793652 JQ793659

Entc. haastii [Entoloma haastii| 126 GQ289307 GQ289167 GQ289238

Entec. nitida [E. nitidum] 7526TJB GU384602 GU384626 GU384655

s GQ289315 — GQ289246

Entc. nitida [E. alcedicolor] 210 GQ289292 GQ289152 GQ289224

Entc. trachyospora [E. trachyosporum] 414 GQ289339 GQ289199 —

405 GQ289338 GQ289198 —

Entc. trachyospora [R. trachyospora] 5856TJB GU384605 GU384629 GU384658

Entc. turbida [E. turbidum] 27. GQ289341 GQ289201 GQ289269

6949TJB GQ289341 GQ289201 GQ289269

Entc. vinacea [E. vinaceum| 8870TJB GU384598 GU384631 GU384651

E. bloxamii 219 GQ289294 GQ289154 GQ289226

E. caesiolamellatum [E. bloxamii] 6117TJB = AF261289 —

E. coeruleogracilis [E. haastii] 216 GQ289308 GQ289168 GQ289239

217 GQ289309 GQ289169 GQ289240

E. cretaceum 213 GQ289302 GQ289162 GQ289233

E. gelatinosum 212 GQ289305 GQ289165 GQ289236

E. griseolazulinum ill GQ289306 GQ289166 GQ289237

E. indigoticoumbrinum 83 GQ289318 GQ289178 GQ289249

E. kermandii 222 GQ289313 GQ289173 GQ289244

E. perbloxamii 71 GQ289318 GQ289178 GQ289249

E. prunuloides 40 GQ289324 GQ289184 GQ289255

Entoloma sp. DLL 10217 JQ793646 JQ793653 JQ793660

BY21 — AF261309 —

DLL 10055 — JQ793655 JQ793662

Lepista ovispora 442 GQ289347 GQ289207 GQ289274

Rhodocybe sp. [Clitopilus sp.] 211 GQ289353 GQ289212 GQ289279

R. aureicystidiata PBM1902 _ AY380407 AY337412

R. caelata [C. caelatus] 6995TJB GU384600 GU384625 GU384652

R. fallax CBS129.63 AF357083 AF223166 EF421051

R. fallax [C. fallax] 262 GQ289349 GQ289210 GQ289275

37 GQ289350 GQ289276 GQ289276

R. mundula 7599TJB — AY700182 DQ474128

R. nitellina [C. nitellinus] 265 GQ289354 GQ289214 GQ289281

400 GQ289355 GQ289215 GQ289282

R. pallidogrisea [C. pallidogriseus] 118 GQ289356 GQ289216 GQ289283

R. popinalis [C. popinalis] 260 — GQ289213 GQ289280

R. popinalis 6378TJB GU384601 AF261285 GU384654

R. pruinosostipitata MCA1492 GU384608 GU384627 GU384653

R. pseudopiperita [C. pseudopiperitus] 162 GQ289357 GQ289217 GQ289284

R. spongiosa MCA2129 GU384604 GU384628 GU384657

Tricholoma vaccinum 446 GQ289358 GQ289219 GQ289286

T. viridiolivaceum PBM3093 — JF706317 JF706319

New sequences generated for this study are shown in bold. Square bracketed annotations indicate

species names applied in GenBank that differ from those in the phylogram (Fie. 1).

Entocybe haastii comb. nov. (Australia) ... 63

Fae

- - 7

. a _

: ~~ = S *&

—_e

~ ae

PLaTE 1. Entocybe haastii {DLL 10087}. A: Densely appressed-fibrillose pileus surface; B: Dark

bluish-grey lamellae and stipe apex. Bar (A, B) = 5 mm.

measured, and digital microphotographs have been described in Largent et al. (201 1a,b),

while techniques for describing, coding, naming colors from Kornerup & Wanscher

(1978) and factors determined from mathematical analyses in the descriptions are

covered in Largent et al. (2013). Generic abbreviations used are: E. = Entoloma; Entec.

= Entocybe.

The holotype collection and one additional collection are deposited in The Plant

Pathology Herbarium, Orange Agricultural Institute (DAR). The herbarium acronym

is from Thiers (2012).

DNA sequences

Sequences were obtained for three loci (mtSSU, LSU, RPB2) for two E. haastii

collections from Australia (TABLE 1). The extraction and Polymerase Chain Amplification

(PCR) protocols for the mitochondrial small subunit of the ribosomal DNA (mtSSU),

variable domains (D1, D2) of the nuclear large subunit (LSU) and the second largest

subunit of the RNA polymerase gene (RPB2) follow Largent et al. (2011b). The primers

rpb2-i6f and rpb2-i7r or rpb2-EntF2 and rpb2-EntR4 were used to amplify the RPB2

(Co-David et al. 2009, Largent et al. 2013). Sequences were generated on an Applied

64 ... Bergemann, Largent & Abell-Davis

Biosystems 3130xl Genetic Analyzer at Middle Tennessee State University using the

sequencing protocols outlined in Largent et al. (2011b).

Sequences were assembled and edited using Sequencher ver. 4.2.2 (Gene Codes

Corporation, Ann Arbor, MI) and multiple sequence alignments were generated using

MAFFT version 7.1 (Katoh & Standley 2013) and adjusted manually using Se-Al ver.

2.0a11 Carbon (Rambaut 2002) after exclusion of introns from the RPB2 alignment. The

combined mtSSU, LSU and RPB2 sequences were aligned with closely related sequences

from GenBank within the Prunuloides clade, representatives of the basal Rhodocybe-

Clitopilus clade, and appropriate outgroups (Matheny & Baroni 2011) (TABLE 1). The

alignment lengths were 1909 bp (mtSSU), 1472 bp (LSU), and 972 bp (RPB2).

The sequence alignment was analyzed using a Maximum Likelihood (ML) search

GTRGAMMA model using RAxML-HPC v. 7.2.8 ALPHA (Stamatakis 2006). The

dataset was partitioned using 4 partitions: RPB2 (across each codon position), mtSSU

and LSU. Bootstrap proportions were estimated based on 1000 rapid ML replicates and

1000 bootstrap replicates (Stamatikas et al. 2008).

Results

The best likelihood tree in RaxML showed that species identified as Entocybe

(Baroni et al. 2011) formed a well-supported clade (BS = 97, Figure 1) and

lends further support for the generic concept as defined by Baroni et al. (2011).

Additional comments on Entc. haastii are provided in the Comments section.

Taxonomy

Entocybe haastii (G. Stev.) Largent, comb. nov. PLATES 1-3

MycoBank MB 804015

= Entoloma haastii G. Stev., Kew Bull. 16(2): 229 (1962).

Habit mycenoid but with a broad stipe base. PILEUs 24-33 mm broad, 10-11

mm high, umbo obscurely broad, opaque, not hygrophanous, not translucent-

striate, dull, moist but not sticky, conic- or convex-campanulate, densely

appressed- or matted-fibrillose, rivulose, dark blackish blue (19F3 or 19F3-4)

to nearly black, lacking any brown color, becoming silky or satiny appressed

fibrillose with maturity; margin decurved, entire becoming slightly eroded;

context 0.75-2.0 mm thick above the stipe, + violaceous. ODoR mild or faintly

fragrant. Taste mild, at times suggestively sweet. LAMELLAE 11-15 mm long,

5.5-11.0 mm broad, narrowly adnate or adnate, close, at first dark bluish grey

(19E-F2-3) with basidiospore maturity pale orange (6A3) then orange grey

(6B2); margin smooth and concolorous. STIPE 40-65 3-5 (apex) - 4-10 mm

(middle) - 6-12 (base) mm, clavate, shiny and glossy, appressed-fibrillose,

blackish blue but somewhat lighter than the pileus (19E5 or 20F6), white or

yellowish white at the base, hollow and splits longitudinally with ease; context

cartilaginous, white at least at the base; basal tomentum absent. BRUISING

REACTIONS none.

Entocybe haastii comb. nov. (Australia) ... 65

PLATE 2. Entocybe haastii {DLL 9868 A, B; DLL 10087 C}. A: Stature; B: Mature lamellae; C: Mature

pileal surface. Bar for A, B= 15 mm; C= 10 mm.

BASIDIOSPORES in squashed mounts of lamellae in 3% KOH often collapsed

and thin-walled, some distinctly angular and some indistinctly to faintly

rounded pustulate, in mounts from stipe surface decidedly rounded pustulate

with 6 or more facets, in profile view isodiametric to subisodiametric, rarely

heterodiametric, in polar view isodiametric 6.4-9.5(-10.2) x 5.5-7.9(-9.0) um

(x = 7.7 + 0.7 x 6.9 + 0.6 um; E = 0.96-1.31, Q = 1.11 + 0.07 (isodiametric);

n = 90/2). Basrp1a cylindro-clavate, hardly tapered, 26.2-39.8 x 7.7-11.7 um

r= 32.1 +328: K-95 411.0 pm; ES 2.51-4,.42,,0 93 Al + 0:48; in -=926/2);

2- or 4-sterigmate. HYMENIAL CYSTIDIA absent. LAMELLAR TRAMAL HYPHAE

subparallel, moderately short and broad, 48.2-131.3 x 7-30.2 um. PILEIPELLIS

mostly a multi-layered cutis with a few semi-erect terminal cells, subtended

by inflated cells of the pileal trama. PILEOCysTIDIA (terminal cells) cylindric

to narrowly obclavate, 54.1-81.1 x 2.5-7.8 um (n = 7/1). PILEAL TRAMAL

HYPHAE beneath the pileipellis, short, broad and inflated, 47-129 x 8-22 um

66 ... Bergemann, Largent & Abell-Davis

enlarging up to 188 um long and 33 um wide near the hymenium (n = 14/2).

STIPITIPELLIS a cutis. CAULOCYSTIDIA and HYMENIAL CLUSTERS absent.

OLEIFEROUS HYPHAE rare to scattered in the pileal trama. L1poID GLOBULES

absent. BRILLIANT GRANULES absent. PIGMENTS cytoplasmic, brownish blue,

soluble in 3% KOH in pileipellis and inflated hyphae of outer pileal trama;

faintly parietal or possibly encrusted on the narrow hyphae of the pileal trama.

CLAMP CONNECTIONS present on hyphae of all tissues.

EcoLoey and DISTRIBUTION — Solitary in soil among leaf litter or in mosses

on a rock, warm gallery rainforest, Watagans National Park, New South Wales.

COLLECTIONS EXAMINED — AUSTRALIA. NEw SouTH WALES, Central Hunter District,

the Watagans National Park, Boardinghouse Dam Track, 32°59’58.6”S 151°24’15.5”E, 13

April 2010, DL Largent 9868 (DAR); 32°59’56”S 151°24’15”E, 12 April 2011, DL Largent

10087 (DAR).

ComMENtTs — Entocybe haastii from New South Wales lacks the brown

color in the pileus reported for this species from Tasmania and New Zealand

(Noordeloos & Gates 2012). The lamellae from Tasmania are whitish while those

from Australia and New Zealand are bluish-gray (Horak 2008, Noordeloos

& Gates 2012). This variability suggests that fine color distinctions are not

diagnostic for Entc. haastii.

Similar to Entc. haastii, both Entc. nitida (Quel.) TJ. Baroni et al. (= Entoloma

nitidum Queél.) from Europe and Entoloma alcedicolor Arnolds & Noordel.

from Holland have small isodiametric basidiospores and abundant clamp

connections and lack cheilocystidia (Noordeloos 2004).

Entocybe nitida is distinguished from Entc. haastii by a glabrous or radially

fibrillose pileus, white 30-100 mm long immature lamellae, <100 mm long

cylindrical stipe, slightly farinaceous or raphanoid odor, and shorter and

broader tramal cells subtending the pileipellis (Noordeloos 1981, 1992, 2004).

Although Noordeloos (1992) identified basidiomata from Germany with bluish

lamellae as Entc. nitidum, identification remains speculative as no collection

was retained.

Entoloma alcedicolor differs from Entc. haastii by its steel blue pileus and

stipe, garlic odor, squamulose pileus center, and trichodermial pileipellis on

the center of the pileus.

Entoloma gracilior Noordel. & G.M. Gates from Tasmania, which shares with

Entc. haastii black pilei, lilac grey lamellae, a dark blue stipe as well as a similar

pileipellis and basidiospores, differs in its more mycenoid stature and more

slender (2-4 mm across the apex) equal stipe, a weakly hygrophanous smooth

pileus with a translucent-striate margin, and heterodiametric basidiospores

(Noordeloos & Gates 2012).

Entoloma coeruleogracilis G.M. Gates & Noordel., also from Tasmania, is

distinguished by its sky blue stipe and pileus with a violet hue and the 1-3 mm

broad cylindrical stipe.

Entocybe haastii comb. nov. (Australia) ... 67

- a? o-

: ‘a = Qe:

a ss +

= sis

PLATE 3. Entocybe haastii {DLL 10087}. A: Basidiospores from stipe surface; B: Basidia and

basidioles; C: Pileipellis and subtending inflated cells in pileal trama. Bars A = 7.0 um; B = 10 um;

C= 22 um.

Our phylogenetic analysis (Fic. 1) clearly supports (BS = 97) Entc. haastii

within Entocybe as described by Baroni et al. (2011). Entocybe haastii,

E. coeruleogracilis, Entc. nitida, and E. alcedicolor segregate as a sub-clade with

68 ... Bergemann, Largent & Abell-Davis

moderate support (BS = 77). Noordeloos & Gates (2012) classify these taxa in

Entoloma sect. Nitida (Romagn.) Noordel. based on a number of morphological

features.

Acknowledgments

Fieldwork in Australia was supported by the Largent family trust and we are

particularly grateful for the support of Pamela Largent. Fieldwork and logistical support

were provided by the Australian Tropical Herbarium and the School of Marine and

Tropical Biology, James Cook University. Comments by the two reviewers, Dr. Timothy

J. Baroni and Dr. Terry W. Henkel, and Ms. Fernanda Karstedt and Nomenclature Editor

Dr. Shaun Pennycook are also appreciated. We wish to thank Pam O'Sullivan as well as

Skye Moore for their assistance in providing descriptions and sharing the location of

collecting sites in New South Wales.

Literature cited

Baroni TJ, Hofstetter V, Largent DL, Vilgalys R. 2011. Entocybe is proposed as a new genus in

the Entolomataceae (Agaricomycetes, Basidiomycota) based on morphological and molecular

evidence. North American Fungi 6 (12): 1-19. http://dx.doi.org/10.2509/naf2011.006.012

Horak E. 2008. Agaricales of New Zealand. 1: Pluteaceae (Pluteus, Volvariella) Entolomataceae

(Claudopus, Clitopilus, Entoloma, Pouzarella, Rhodocybe, Richoniella). Fungi of New Zealand

Volume 5. Fungal Diversity Research Series 19: 1-305.

Katoh K, Standey D.M. 2013. MAFFT Multiple Sequence Alignment Software Version 7:

Improvements in Performance and Usability. Molecular Biology and Evolution: 30: 772-780.

http://dx.doi.org/10.1093/molbev/mst010

Largent DL, Abell-Davis SE, Cummings GA, Ryan KL, Bergemann SE. 201la. Saxicolous

species of Claudopus (Agaricales, Entolomataceae) from Australia. Mycotaxon 116: 253-264.

http://dx.doi.org/10.5248/116.253

Largent DL, Bergemann SE, Cummings GA, Ryan KL, Abell-Davis SE, Moore S. 2011b. Pouzarella

(Agaricales, Entolomataceae) from New South Wales (Barrington Tops National Park) and

northeastern Queensland. Mycotaxon 117: 435-483. http://dx.doi.org/10.5248/117.435

Largent DL, Bergemann SE, Abell-Davis SE, Kluting KL, Cummings GA. 2013. Five Leptonia

species from central New South Wales and Queensland, Australia. Mycotaxon 125:11- 35.

http://dx.doi.org/10.5248/125.11

Noordeloos ME. 1981. Introduction to the taxonomy of the genus Entoloma sensu lato (Agaricales).

Persoonia 11: 121-151.

Noordeloos ME. 1992. Entoloma s.l. in Fungi Europaei vol. 5. Ed. Candusso: Alassio, Italy.

Noordeloos ME. 2004. Entoloma s.l. in Fungi Europaei vol. 5A. Ed. Candusso: Alassio, Italy.

Noordeloos ME, Gates, GM. 2012. The Entolomataceae of Tasmania. Fungal Diversity Research

Series 22: 1-399. http://dx.doi.org/10.1007/978-94-007-4679-4

Rambaut A, 2002. Sequence alignment editor. Version 2.0. Department of Zoology, University of

Oxford, Oxford England.

FIGURE 1. Maximum likelihood tree (LSU+mtSSU+RPB2) showing the position of Entocybe haastii

from Australia (AUS*) and Tasmania (TAS) relative to other Entocybe collections. Each sequence

is labeled with the GenBank-listed isolate number. Branches with >70% support from 1000 rapid

bootstraps provide statistical support. Clitocybe dealbata, Lepista ovispora, Tricholoma vaccinum,

and T: virdioviolaceum serve as outgroups. Three slanted lines denote truncated branches.

0.08 substitutions/site

————————————— 99

100

hoor ENtoloma perbloxamii 71

Entoloma indigoticoumbrinum 83

Entocybe trachyospora 414

41 Entocybe trachyospora 405

9a|LEntocybe trachyospora 5856TJB

Entocybe turbida 27

Entocybe vinacea 8870TJB

Entocybe turbida 6949TJB

93, Entocybe haastii DLL10087*

P96 Entocybe haastii DLL 9868*

Entoloma haastii 126 TAS

Entocybe coeruleogracilis 216

Entocybe coeruleogracilis 217

Entocybe nitida 210

oof Entocybe nitida 7526TJB

Entocybe nitida s

1007 Rhodocybe fallax 262

100

100 __[f]

77 |

100 97

100

100;

80}! Calliderma sp. DLL10054

100

Rhodocybe nitellina 2 el

Rhodocybe nitellina 400 Rhodophana

Entocybe haastii comb. nov. (Australia) ...

Entoloma bloxamii 219

Entoloma prunuloides 40

Entoloma gelatinosum 212

Entoloma caesiolamellatum 6117TJB

Entoloma sp. DLL10217

Entoloma cretaceum 213

95, Calliderma sp. DLL10025

Calliderma sp. DLL10088

Entoloma griseolazulinum i11

Entoloma sp. BY21

Entoloma kermandii 222

Entoloma sp. DLL10055

Prunuloides

Entocybe

Rhodocybe fallax CBS129.63

Rhodocybe fallax 37

Rhodocybe popinalis 6378TJB

Rhodocybe popinalis 260

Rhodocybe mundula AFTOL-ID 521

Clitopilus cystidiatus 26

Clitopilus prunulus 8229TJB

Clitopilus prunulus 9663TJB

Clitopilus “cinerascens” 8024TJB

Clitopilopsis hirneola 8490TJB |

Clitopilopsis hirneola 263 Clitopilopsis

Rhodocybe pseudopiperitus 162

Rhodocybe pallidogrisea 118

Rhodocybe sp. 211

Rhodocybe spongiosa MCA2129

Rhodocybe caelata 695TJB

Rhodocybe pruinosostipitata MCA1492

Rhodocybe aureicystidiata PBM1902

100

Clitopilus-Rhodocybe p.p.

Rhodocybe s.s.

Outgroups

70 ... Bergemann, Largent & Abell-Davis

Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with

thousands of taxa and mixed models. Bioinformatics 22: 2688-2690.

http://dx.doi.org/10.1093/bioinformatics/btl446

Stamatakis A, Hoover P, Rougemont J. 2008. A rapid bootstrap algorithm for RAxML web servers.

Systematic Biology 57: 758-771. http://dx.doi.org/10.1080/10635 150802429642

Thiers B. 2012. Index Herbariorum: a global directory of public herbaria and associated staff. New

York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg.org/ih/[accessed May 2012]

MY COTAXON

http://dx.doi.org/10.5248/126.71

Volume 126, pp. 71-75 October-December 2013

Craterium corniculatum sp. nov. from northwestern China

Bo ZHANG? & Yu Lr'*

‘Engineering Research Center of Chinese Ministry of Education for Edible and Medicinal Fungi,

Jilin Agricultural University, 2888 Xincheng Street, Changchun City, P. R. China

* CORRESPONDENCE TO: yuli966@126.com

ABSTRACT —Craterium corniculatum is a new species collected in Gansu Province, China,

characterized by horned sporophores and warted spores. We provide a morphological

description, illustrations, and a key to the 17 Craterium species. The specimens were

deposited in the Herbarium of Mycological Institute of Jilin Agricultural University

(HMJAU), Changchun, China.

KEY worDs — taxonomy, SEM, slime molds, Myxogastrea, Physaraceae

Introduction

Myxomycetes are common inhabitants of decaying plant materials in nature,

such as decaying logs, stumps, and dead leaves (Stephenson & Stempen 1994).

Trentepohl established the genus Craterium in 1797 for myxomycetes with

stalked sporophores, a persistent basal cup, and subglobose or turbinate obovate

sporocarps that show a distinct or indistinct lid during dehiscence (Martin &

Alexopoulos 1969). In 2012 we found a new species on the bark of a dead log

on Jilian Mountain, Gansu Province, China, that differs morphologically from

all other Craterium species.

Materials & methods

The fruiting bodies and microscopic structures were examined by light and scanning

electron microscopes (Martin & Alexopoulos 1969, Zhang & Li 2013). Permanent

slides were mounted in Hoyer'’s reagent (Martin & Alexopoulos 1969). We prepared

them according to Robbrecht (1974) by spreading capillitia in a drop of 94% alcohol,

determining colour after one minute, and then mounting in Hoyer’s reagent. Colour

terms are given according to Flora of British Fungi (Royal Botanic Garden Edinburgh

1969).

We observed more than ten sporocarps under a stereomicroscope (20x) and more

than 20 spores under an optical microscope (100x). The sporophores, capillitia, and

72... Zhang & Li

spores were measured by using a Nikon DM1000 microscope and photographed with

a Canon G15 camera. For ultrastructural observation, the sporophores were attached

to the holder, coated with gold using a Hitachi E-1010 sputter, and examined with a

Hitachi S-4800 scanning electron microscope at 10 kV at Changchun Institute of

Applied Chemistry, Chinese Academy of Sciences. The specimens were deposited in the

Herbarium of Mycological Institute, Jilin Agricultural University (HMJAU).

Taxonomy

Craterium corniculatum B. Zhang & Yu Li, sp. nov. PLATE 1

MycosBank MB 806043

Differs from Craterium aureomagnum and C. aureum by its horned sporophores and

warted spores.

Type: China, Gansu Province, Jilan Mountain, on bark of a dead log, 6 August 2012,

Zhang Bo 20122203, (Holotype, HMJAU10550).

EryMo_oey: corniculatus (Latin), referring to the ‘horned’ sporophores.

SPOROPHORES gregarious, in small groups, shortly stalked or nearly sessile,

0.25-0.75 mm tall. Sporotheca subglobose or pyriform to irregular, 0.2-0.5

mm in diam. HyPOTHALLUM conspicuous, membranous, pale yellow, veined

with ribs radiating from the sporocarp base. STALK either lacking, or very short

and consisting of updrawn hypothallus, strongly ribbed, yellowish brown, pale

brown in transmitted light. Lrp paler than cup, usually with a paler calcareous

deposit, yellowish pale by transmitted light, with pale dehiscence lines and

circumscissile area. PERrDIUM thick, single, cartilaginous, containing small

lime. CAPILLITIUM rather dense, lime nodes rounded or angular, large, pale

yellowish brown by transmitted light, 40 x 70-80(-100) um. Sporss free,

fuscous black in mass, dark brown by transmitted light, subglobose, 8-14 um

diam., densely warted.

ADDITIONAL SPECIMEN EXAMINED: CHINA, GANSU PROVINCE, Jilan Mountain, on

bark of a dead log, 26 August 2010, Zhang Bo 20101201 (HMJAU10551).

ComMENts: Of the 16 species reported for Craterium Trentep. (Kirk et al.

2008, Lado 2005-13), seven have been found in China: C. aureum (Schumach.)

Rostaf., C. concinnum Rex, C. leucocephalum (Pers. ex J.F. Gmel.) Ditmar,

C. microcarpum H.Z. Li et al., C. minutum (Leers) Fr., C. obovatum Peck, and

C. rubronodum G. Lister (Li & Li 1989, Li et al. 1993). Among the 16 accepted

species, only C. aureomagnum Hooff & Nann.-Bremek. and C. aureumare similar

to C. corniculatum in having yellow sporocarps, but C. aureomagnum has larger

sporocarps (0.7-1.5 mm tall), double peridia, and darker and prominently

warted spores that are 10-12 um in diam. (Hooff & Nannenga-Bremekamp

1996), while C. aureum has stalked, turbinate, or obovoid sporocarps that are

0.7-1.0 mm tall and lemon-yellow to almost white (Rostafinski 1974).

Craterium corniculatum sp. nov. (China) ... 73

Acc.¥ Spot Magn

12.0kV 3.0 594x

PLaTE 1. Craterium corniculatum (Holotype, HMJAU 10550): 1-2, fruiting bodies; 3, capillitium; 4,

densely warted spores. Scale bar: 1 = 2 mm.

Key to species of Craterium

L -sporophoresdehiscing by-adistinct- lids fn pled sles leans eye ae cat koe ace 2

leSporophores not deliscine; by-adistinct Lae. wicks ah ees Baws bale ARS A 8

Ze SPOMES TEMCULALES, he Pues ee Sn, We ee ane, ee AV Bs C. retisporum

AUSPOLES SPUNULOSESANIG WANE, 5 5. sctag dt: chon Sch geben Sch peben dh dean Sih peteg Sia de deh adage tees 3

SOS POLCS NOMI PAT a RAT sh.) 4 sare 5 4a eB 4 tart ¢ 4 bet 4.4! ore FG ebet-Z4'Meete 4 Dee A bebe 4

SvSporesws Ouran dias 3 lore da Sealed. cS. Uda. ced al ada d ye etand ag enahdledags 6

4. Lid dome shaped, fragmenting by preformed lines at dehiscence ............... 5

4. Lid convex or fat, not veined, remaining whole at dehiscence .................. vi

5. Sporocarps 0.15-0.41 mm high; spores 8-10 um indiam. ........ C. microcarpum

5. Sporocarps 0.6-1 mm high; spores 7-9 um in diam. ............... C. reticulatum

6. Sporocarps short (0.5-0.8 mm high); spores 9-10 um in diam. ....... C. concinnum

6. Sporocarps taller (<1.5 mm high); spores 8-10 um in diam. .......... C. minutum

74 ... Zhang & Li

7. Spores 10-12 um in diam. with lum tall spines................... C. rubronodum

7. Spores 13-15 um in diam. with groups of confluent spines ............ C. costatum

8. Sporophores obconic; becoming cup-shaped after dehiscence .................. 9

8. Sporophores obovoid, turbinate, corniculate; dehiscence irregular ............. 10

9. Sporocarps usually stalked; spores 7-10 um in diam.............. C. leucocephalum

9. Sporocarps sessile or short-stalked; spores 9-10 um in diam..... C. aureonucleatum

POS pondcarpissyellow Fx iu tacts oS settee + hlgn Bard te ole Sete ate see als Fete. glt ae big ele Bele lt 11

10. Sporocarps bright pink, purplish pink, red brown .................0..00000. 13

Mi Sporocat pe stalked: PF 1st tates ics ats ig a door 8 gee oe oe EOE eo C. aureum

Ll Sporocarpssesstleor veryahort-stalked an ccs y stomcte x simi y stoncla matemelr aforatet 4 12

12. Sporocarps subcylindric; spores 10-12 umin diam. ............ C. aureomagnum

12. Sporocarps corniculate; spores 8-14 um in diam................. C. corniculatum

13. Sporocarps bright pink, purplish pink ....................00.. C. paraguayense

13. Sporocarps red brown, purplish brown, dark brown to black ................. 14

14 Spores warted,< ES Mae Cia. anccetg sl anewtte! sacs seo anes 4+) aed’ ds onese bate C. atrolucens

14. Spores coarsely warted, subreticulate or reticulate, >13um in diam. ........... 15

lSespores-withdaris prominent walktsy.£ nay). kame ek eek ee ess C. obovatum

1 SO POLES SUBKEMCU ALE LOL LOU CUM ALE ¢ ctriro.deg-ds pone dite se eeadeh Aebag arp de bon dre sehen dh aedeed ples 16

16. Spores marked with an incomplete net of broad spines fusing into short ridges,

eS SING EROT CARN 1tC Pad gD aa CM eM ee i a hat emcee C. dictyosporum

16. Spores loosely reticulate by ridges about 1 um high, 13-16 um in

OPEN TEA Dk Pe Ren eee Ren Pees eek an whee Ry ME Pon OR Pn ee C. muscorum

Acknowledgments

We express our deep appreciation to Prof. Guozhong Li (Dalian Nationalities

University, PR. China) and Prof. A.J.S. Whalley for their valuable suggestions in peer-

reviewing this manuscript. We thank Tianhao Li, Jilin Agricultural University for

correcting the manuscript. The study was supported by the fund from the Ministry of

Agriculture of China.

Literature cited

Hooff JPM van, Nannenga-Bremekamp NE. 1996. Additions to the myxomycetes of the

Netherlands. Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen,

Section C, 99: 45-53.

Kirk PM, Cannon PF, Minter DW, Stalpers JA. 2008. Ainsworth & Bisby’s dictionary of the fungi,

10th ed. Wallingford, CAB International. 771 p.

Lado C. 2005-13. An online nomenclatural information system of Eumycetozoa.

http://www.nomen.eumycetozoa.com (accessed: November 19, 2013).

Li HZ, Li Y, Chen SL. 1993. Myxomycetes of China XI: a new species of Craterium. Myxosystema

62113-1135,

Li Y, Li HZ. 1989. Myxomycetes from China I: a checklist of myxomycetes from China. Mycotaxon

35(2): 429-436.

Craterium corniculatum sp. nov. (China) ... 75

Martin GM, Alexopoulos CJ. 1969. The myxomycetes. Iowa, University of Iowa Press. 561 p.

http://dx.doi.org/10.2307/1218569

Robbrecht E. 1974. The genus Arcyria Wiggers in Belgium. Bulletin du Jardin Botanique National

de Belgique 44: 303-353. http://dx.doi.org/10.2307/3667676

Rostafinski JT, 1874. Sluzowee (Mycetozoa). Monografia. Paryz. 1-215

Royal Botanic Garden Edinburgh. 1969. Flora of British fungi: colour identification chart.

Edinburgh, H.M. Stationery Office. 6 p.

Stephenson SL, Stempen H. 1994. Myxomycetes: a handbook of slime molds. Portland, OR, Timber

Press.

Zhang B, Li Y. 2013 [“2012”]. Myxomycetes from China 16: Arcyodes incarnata and Licea retiformis,

newly recorded for China. Mycotaxon 122: 157-160. http://dx.doi.org/10.5248/122.157

MYCOTAXON

http://dx.doi.org/10.5248/126.77

Volume 126, pp. 77-81 October-December 2013

Embryonispora, a new genus of hyphomycetes from China

KAI-NING ZHAO’, GUO-HUA YIN?, GUO-ZHU ZHAO & AI-XIN CAO3

‘College of Biological Sciences and Biotechnology, Beijing Forestry University,

Beijing 100083 PR China

College of Agriculture & Life Sciences, The University of Arizona, Tucson, AZ 85721 USA

°State Key Laboratory of Urban & Regional Ecology, Research Center for Eco-environmental

Sciences, Chinese Academy of Sciences, Beijing 100085 PR China

* CORRESPONDENCE TO: zhaogz@im.ac.cn

AsstraAct — Embryonispora bambusicola gen. & sp. nov., found on dead culms of bamboo

and dead branches of an unidentified tree in China, is described and illustrated. The

genus is characterized by sporodochial conidiomata and muriform, reniform or flattened,

dorsiventrally curved, pale brown to brown conidia. The new taxon is compared with

morphologically close related genera and species.

KEY worps — anamorphic fungi, dictyosporous fungi, taxonomy

Introduction

In a study of dictyosporous hyphomycetes from China, several fungal

specimens were collected between 2002 and 2003. A fungus with sporodochial

pulvinate or effuse conidiomata and flattened dorsiventrally curved muriform

embryoniform conidia was found among them. It cannot be assigned to any

previously described genus and therefore is described here as new.

Materials & methods

Decayed waste wood was collected in the field of Zhejiang and Hainan Province, China.

Samples were placed separately into plastic or paper bags and stored in a refrigerator

at 4°C before microscopic study. They were incubated to induce sporulation in moist

containers (plastic bags or boxes) at room temperature. Humidity was maintained by

adding moistened paper towels. The incubated samples were examined microscopically

for the presence of microfungi after 4-5 days and up to one month. Sporulated colonies

were removed from the surface of the wood with a needle and transferred to a drop

of lactophenol on a microscope slide. Extensive attempts to culture fungi on artificial

media (water agar, 2% malt extract agar) failed. Digital photomicrographs were taken

using an Olympus BX-51 microscope. Herbarium specimens and slides were deposited

78 ... Zhao & al.

in HSAUP (Herbarium of Shandong Agricultural University: Plant Pathology) and

HMAS (Herbarium of Mycology, Chinese Academy of Sciences).

Taxonomy

Embryonispora G.Z. Zhao, gen. nov.

MycoBank MB 802690

Differs from other sporodochial genera by its reniform and dorsiventrally curved

conidia.

TYPE SPECIES: Embryonispora bambusicola G.Z. Zhao

Erymo toecy: Embryonispora, referring to the conidia, which are shaped like a human

embryo.

Conidiomata on natural substrate sporodochial, superficial, olivaceous,

brown to dark brown. Mycelium partly superficial and partly immersed.

Conidiophores undifferentiated, reduced to conidiogenous cells.

Conidiogenous cells monoblastic, integrated, terminal, determinate. Conidial

secession schizolytic. Conidia solitary, acrogenous, embryoniform or reniform,

curved dorsiventrally, flattened from side to side, sometimes variable in shape,

muriform, with longitudinal, transverse and oblique septa, with 1-2 layers of

cells, holoblastic, olivaceous, pale brown to brown.

Embryonispora bambusicola G.Z. Zhao, sp. nov. FIGS 1, 2

MycoBank MB 802691

Differs from other sporodochial species by its reniform, dorsiventrally curved, muriform

conidia usually composed of 1-2 layers of marginal cells and some irregular protruding

core cells.

Type: China. Zhejiang Province, Hangzhou Botanical Garden, on dead bamboo culms,

12 August 2002, coll. G.Z. Zhao ZGZII,,172 (Holotype, HSAUP0972).

Erymo.oey: bambusicola, referring to the host on which the taxon was first found.

Conidiomata on natural substrate sporodochial, scattered to confluent,

pulvinate, brown to dark brown. Mycelium superficial and immersed,

composed of septate, pale brown to brown, smooth-walled, branched hyphae.

Conidiophores undifferentiated. Conidiogenous cells monoblastic, integrated,

pale brown, determinate, cylindrical. Conidial secession schizolytic. Conidia

solitary, curved dorsiventrally, embryoniform or cerebriform, muriform,

with longitudinal, transverse and oblique septa, often constricted at the septa,

usually composed of 1-2 layers of marginal cells and some irregular protruding

core cells, holoblastic, olivaceous, pale brown to brown, 16-22 x 12-16 um,

9-11 um thick. Basal cell protruding, obconical to cylindrical, subhyaline to

pale brown, truncate, 1.5-2.5 um wide.

ADDITIONAL SPECIMEN EXAMINED: CHINA. HAINAN PROVINCE, Wuzhishan, on dead

branches of an unidentified tree, 14 December 2003, coll. G.Z. Zhao ZGZII,,106-2

(HMAS 90279).

Embryonispora bambusicola gen. & sp. nov. (China) ... 79

Fic. 1. Embryonispora bambusicola (holotype HSAUP0972).

A-D. Conidia; arrows show pale brown basal cells and conidia in side view. Bars = 10 um

Discussion

Several genera with sporodochial conidiomata, less differentiated

conidiophores, and muriform conidia are morphologically similar to

Embryonispora. They include Canalisporium Nawawi & Kuthub., Dictyosporium

Corda, Turturconchata J.L. Chen et al. (= Venustusporium R.F. Castaneda &

Iturr., now corrected to “Venustisporium” in accordance with Art. 60.8 of the

International Code of Nomenclature, McNeill & al. 2012), Oncopodiella G.

Arnaud ex Rifai, Petrakia Syd. & P. Syd., Oncopodium Sacc., and Hermatomyces

Speg. (Ellis 1971, 1976; Nawawi & Kuthubutheen 1989; Chen at al. 1999;

Castahfeda & Iturriaga 1999; Chung & Tzean 2000). Canalisporium and

80 ... Zhao & al.

Fic. 2. Embryonispora bambusicola (holotype HSAUP0972).

A. Sporodochium with conidia; B. Mature conidia with pale brown basal cells. Bars = 10 um

Dictyosporium can be separated from Embryonispora by having cheiroid, fan-

shaped, regularly muriform, flattened conidia, which are usually composed of

columns or rows of cells. The conidia of Turturconchata are borne horizontally

flattened, muriform, margin irregular, somewhat crenate, eccentric protuberate

pedicel, while those of E. bambusicola are vertically dorsiventrally curved and

muriform. Oncopodiella and Petrakia differ from Embryonispora by producing

muriform conidia with hyaline horn-like protruding cells. Oncopodium is

distinct from Embryonispora in having conidia that commonly contain swollen

basal cells. Hermatomyces differs from Embryonispora in the presence of

lenticular conidia.

Key to Embryonispora and morphologically similar genera

LS@GHicia USHA Tattene ls seule apdtagne orsaielee otauils oases oaauaeles casei a Meese a Ba ad gZ

1. Conidia not flattened, with protruding hyaline horn-like cells .................. 3

2 CONIAAAIORSIVENTEAL CUEVERS #8 9% F888. 0 888 TR e FRA me kN 28 te Ee 4

2; @Onidia- shaped: Giikeremty 5 df a disse a dcyace dh dshaeed d dypey 8 diane d deen a Boeke dB setae 6

Embryonispora bambusicola gen. & sp. nov. (China) ... 81

3. Mornclike cells S83 iy, cies. te 5.25 S Pein Gh Gis ee Gin, ine Wales ale 28 Petrakia

3. Horn-like:cells St vers hai Wed was Rew ste Mew ate Mod Wigley Wd Grate Wiad boat odes Oncopodiella

4. Conidia subglobose or hemispherical, clathrate or muriform......... Oncopodium

A CHNTCIA SWAP EC, GMTERETIELe-5 toas arate tranasaln Seewaccte tearacte boark aee teawacsee town eecwanncce ukeyneatitas 5

5. Conidia embryoniform or cerebriform, muriform ................ Embryonispora

5. Conidia horizontally, flattened, muriform, margin irregular, somewhat crenate,

eccentriceprotuberate pedigels, Ax ot Biel y thesis + stumain » allen Turturconchata

6. Conidia lenticular, muriform, with pale peripheral cells surrounding the central

CElS aE) ate htoe sandy LAO e Dee Reece en Gre Uy ay Sen STG Eee S Hermatomyces

6eGonidia shaper dutetentiyg 14.98 ee F Moo 200 we Aer) oe ART wn. SEL wheel eg i) om Gh 8 7

7. Conidia usually cheiroid, composed of columns or rows of cells .... Dictyosporium

7. Conidia broadly ellipsoidal, with cell lumen connected by narrow

CAMTCUGS Fe. gees: Peres lose Medes lose Big esse Regt Ege oP eto Regt te ag Ess Canalisporium

Acknowledgments

We are deeply grateful to Drs. G. Delgado (EMLab P&K, USA) and M. Zhang

(Henan Agricultural University, China) for kindly serving as pre-submission reviewers

and their useful suggestions. This project was supported by the National Natural Science

Foundation of China (Project No. J1103516 & 31093440) and the Fundamental Research

Funds for the Central Universities (No.TD2012-03).

Literature cited

Castaneda RF, Iturriaga T. 1999. Venustusporium, a new genus of hyphomycetes from Venezuela.

Mycotaxon 72: 455-459.

Chen JL, Huang TL, Tzean SS. 1999. Turturconchata, a new genus of hyphomycetes from Taiwan.

Mycological Research 103: 830-832. http://dx.doi.org/10.1017/S0953756298008016

Chung CH, Tzean SS. 2000. Notes on the genera Turturconchata and Venustusporium (mitosporic

fungi). Mycotaxon 75: 311-313.

Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey.

Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew,

Surrey.

McNeil J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth DL, Herendeen PS, Knapp S,

Marhold K, Prado J, Prudhomme Van Reine WE, Smith GF, Wiersema JH. 2012. International

code of nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Vegetabile 154.

Koeltz Scientific Books. [Online: www.iapt-taxon.org/nomen/main.php]

Nawawi A, Kuthubutheen AJ. 1989. Canalisporium, a new genus of lignicolous hyphomycetes from

Malaysia. Mycotaxon 34: 475-487.

MY COTAXON

http://dx.doi.org/10.5248/126.83

Volume 126, pp. 83-90 October-December 2013

The genus Neopaxillus (Crepidotaceae)

Roy WATLING”! & M. CATHERINE AIME?”

' Caledonia Mycological Enterprises, Edinburgh EH4 3HU UK

? Department of Botany and Plant Pathology, Purdue University, West Lafayette, IN, 47901 USA

* CORRESPONDENCE TO: caledonianmyc@blueyonder.co.uk

Asstract — Neopaxillus (Agaricomycotina, Basidiomycota) was formerly placed in the

Paxillaceae (Boletales) by Singer. Herein we provide additional evidence that the genus is

referable to the Crepidotaceae (Agaricales) and a redescription of N. echinospermus from

recently collected material.

Key worps — fungal systematics, Melanomphalia, neotropical fungi, Paxillus, Simocybe

Introduction

Neopaxillus Singer (1948) was erected as a monotypic genus, typified by

N. echinosporus. The description was based on a collection by J. Rick from sandy

soilin Southern Brazil that is deposited in the Farlow Herbarium, Massachusetts,

USA. Neopaxillus was characterized as being morphologically intermediate

between the Cortinariaceae (Agaricales) and the Paxillaceae (Boletales). The

habit and trama were likened to that of the Paxillaceae in having a clitocyboid-

omphalinoid facies reminiscent of a small Phylloporus rhodoxanthus

(Schwein.) Bres. (Boletaceae) or Paxillus species. The basidiospores, however,

were described as having characters more reminiscent of Cortinarius, although

their ornamentation was not identical to that of any known Cortinarius species

(Singer 1948).

Soon after his 1948 publication, Singer (1951) determined that Neopaxillus

echinosporus was in fact conspecific with the Argentine Naucoria echinosperma

(Spegazzini 1889), which he recombined as Neopaxillus echinospermus.

Because this name has priority, Neopaxillus echinospermus is the correct name

for the combined taxon; however, the genus Neopaxillus remains typified by

N. echinosporus. In later works, Singer included two other South American

species, Flammula echinospora (Spegazzini 1902) and Flammula papillosispora

(Spegazzini 1919), in the synonymy of Neopaxillus echinospermus (Singer 1952,

1964; Singer et al. 1990).

84 ... Watling & Aime

Singer (1948) likened Neopaxillus echinosporus in size to Deconica crobula

(Fr.) Romagn., with which Rick apparently confused his collection. One

distinguishing feature was the ornamented spores, which Singer regarded as

a link to Ripartites P. Karst., a genus also then placed in the Paxillaceae. Their

ornamented basidiospores separated both genera from all the other taxa that

Singer accepted in the Paxillaceae (Singer 1990). Neopaxillus is separable from

Ripartites by its larger basidiospores, lamellae that anastomose towards the

stipe, and an almost continuous trichodermial pileipellis composed of chains

of hyphae with broadly clavate terminal cells. Furthermore, according to

Singer (1948), the hymenophoral trama in Neopaxillus is not characteristically

divergent but rather weakly or vaguely bilateral in immature specimens.

It has more recently become apparent that Paxillaceae sensu Singer (1986) is

an artificial family with many disparate constituent elements, e.g. Omphalotus

Fayod (now placed in the Marasmiaceae; Kirk et al. 2008), Hygrophoropsis

(J. Schrét.) Maire ex Martin-Sans (now placed in the Hygrophoropsidaceae; Kirk

et al. 2008), and Phyllobolites Singer (now placed in the Boletaceae; Kirk et al.

2008). The genus Paxillus sensu Singer itself has been subdivided with saprobic

species [e.g., Paxillus panuoides (Fr.) Fr.] now placed in Tapinella E.-J. Gilbert

(Tapinellaceae), southern hemisphere species now in Austropaxillus Bresinsky

& Jarosch (Serpulaceae), and the ectomycorrhizal members remaining in

Paxillus (Kirk et al. 2008, Watling 2008).

Ripartites, which has been placed in several agaric families, including the

Tricholomataceae where it is presently placed, was linked to Neopaxillus by

general appearance, with the pair placed in proximity to Paxillus by virtue of

the open-pored hilum of the basidiospore. Results from numerical analyses

conducted by Machol & Singer (1971) supported this placement. However, in

his review of the boletes and their relatives, Watling (2008) deemed it necessary

to reexamine Neopaxillus, which became possible with collection of new

material in the Atlantic rainforests of Brazil by André de Meijer. Watling’s initial

examination of this material indicated a possible link to the Crepidotaceae due

to the echinulate basidiospores reminiscent of similar ornamentation seen

in some Crepidotus taxa. This observation led us to attempt to infer the true

position of Neopaxillus within the Agaricomycotina using DNA sequence

analyses.

Materials & methods

Collections examined

New collections of N. echinospermus were made by André de Meijer on forest humus

in the Mata Atlantica in Parana, Brazil. Material is deposited in the Royal Botanic

Garden, Edinburgh, Scotland, UK (E). The field colors were matched to Methuen’s

Handbook (Kornerup & Wanscher 1978). Microscopic characters were observed using

a Leitz Ortholux microscope and examined in a 10% aqueous ammonia solution.

Neopaxillus (Crepidotaceae) ... 85

25 spores of each collection in five separate mounts were measured; Melzer’s reagent

(a mixture of chloral hydrate, potassium iodide and elemental iodine) and cotton blue

in lactophenol supplemented the observations; material examined using a Jeol T200

scanning electron microscope were suspended first in the above ammonia solution

before applying to the stub, drying, and coating with gold.

Molecular analyses

DNA was extracted from dried tissue from the two herbarium Neopaxillus

echinospermus specimens with the UltraClean Plant DNA Isolation Kit (MoBio

Laboratories, Inc., Solana Beach, CA) following the manufacturer's instructions. Primers

LSU4-B (Aime & Phillips-Mora 2005) and LR6 (Moncalvo et al. 1995) were used to

amplify and sequence the first ~1200 bp of the 28S ribosomal DNA using previously

described parameters (Aime & Phillips-Mora 2005). Preliminary phylogenetic analysis

was conducted by analyzing LSU sequences of N. echinospermus within the 877 taxa

dataset of Moncalvo et al. (2002). This dataset contains representatives of all major

agaricalean lineages, as well as exemplar taxa from the other major Agaricomycotina

clades, including Boletales, with Auricularia polytricha (Auriculariales) as the outgroup

(Moncalvo et al. 2002). Based on the results of the preliminary analyses (not shown) a

new dataset was constructed with additional sampling from within the Crepidotaceae, to

which N. echinospermus showed affinity. A total of 21 species were chosen including the

type species of Crepidotus [C. mollis (Schaeff.) Staude] and Simocybe [type, S. centunculus

(Fr.) P. Karst.] from previously published Crepidotaceae sequences (Aime & Miller 2002,

Aime et al. 2002, Aime et al. 2005, Aime et al. 2009, Matheny 2005, Moncalvo et al.

2002, and unpublished), exemplar sequences from the genus Inocybe (Matheny 2005,

Moncalvo et al. 2002, Ryberg et al. 2008, and unpublished), which has been suggested as

the sister family to the Crepidotaceae sensu Aime (Matheny et al. 2006), with Agaricus

aff. campestris and Deconica montana (=Psilocybe montana) (Matheny et al. 2006, and

unpublished) selected as outgroups.

New sequences were edited and assembled in Sequencher v.4.1.4 (Gene Codes

Corp., Ann Arbor, MI) and aligned visually in Se-Al v2.0a11 (Andrew Rambaut, Dept.

Zoology, University of Oxford, U.K.; http://evolve.zoo.ox.ac.uk/) over a total of 1074

bp. Maximum likelihood (ML) analyses were conducted in RAxML-HPC2 v. 7.2.7

(Stamatakis 2006) via the CIPRES portal (Miller et al. 2011) using default parameters

and 1000 bootstrapping replicates. Maximum parsimony (MP) analyses were

conducted in Paup* v4.0b10 (Swofford 2002) as heuristic searches with 1000 random

addition replicates and TBR branch swapping; bootstrap analysis consisted of 1000 MP

replicates. Sequences have been deposited in GenBank as JQ404442 (AM 1424) and

JQ404443 (AM 1690).

Results

Based on prior molecular studies, the members of Paxillaceae sensu

Singer have been segregated into three groups, the Paxillineae, Serpulaceae

(Coniophorineae), and Tapinellineae, all of which belong to the Boletales

(Binder & Hibbett 2006). Preliminary results from 28S analyses of Neopaxillus

echinospermus within an inclusive dataset of Agaricomycotina placed

86 ... Watling & Aime

N. echinospermus firmly within the Crepidotaceae, with no affinity to the

Boletales (data not shown). A second dataset was constructed with exemplars

from all Crepidotaceae genera and representative outgroups (Fig. 1), strongly

supporting the placement of N. echinospermus within the Crepidotaceae. The

28S locus, alone, did not provide strong support separating the three genera

(Crepidotus, Simocybe, and Neopaxillus), although each forms a monophyletic

clade within Crepidotaceae under all searching algorithms used, with

Neopaxillus consistently resolved as the sister lineage to Crepidotus.

Taxonomy

Neopaxillus echinospermus (Speg.) Singer, Lilloa 22: 633. 1951 [“1949”].

= Naucoria echinosperma Speg., Bol. Acad. Nac. Cienc. Cordoba 11: 424. 1889.

= Flammula echinospora Speg., An. Mus. Nac. B.Aires 8: 51. 1902.

= Flammula papillosispora Speg., Bol. Acad. Nac. Cienc. Cordoba 23:396. 1919.

= Neopaxillus echinosporus Singer, Mycologia 40: 262. 1948.

PiLEus 12-40 mm broad, center depressed even in youth, slightly hygrophanous,

non-striate when fresh, golden brown, brownish orange when dry, smooth, dry,

glabrous; margin often decurved, sometimes undulate. LAMELLAE strongly

decurrent, distant, margin straight or concave, 5.5 mm broad, clay-brown at

first, dark brown when fully mature, neither interveined, nor forked. STIPE

17-40 x 3.5-7 mm, cylindrical or slightly attenuated downwards, solid, paler

and more yellow than pileus, minutely innately brown-striate, smooth, dry;

basal mycelium white. Context in pileus when fresh yellowish brown, drying

pale ochraceous to white; in stipe drying white, but in stipe base often becoming

brown; odor fungoid or none.

BASIDIOSPORES 7-8(-8.5) um subglobose, strongly warty-echinulate,

ornaments <0.7 um high, inamyloid. CHEILo- & PLEUROCYSTIDIA absent.

PILEIPELLIS composed of irregularly arranged, rarely encrusted hyphae 5-7 um

broad. CLAMP-CONNECTIONS absent.

MATERIAL EXAMINED: BRAZIL, Parana, General Caneiro, in forest humus, 20 xii 1989,

Meijer 1424 (E); 12 v 1990, Meijer 1690 (E).

Notes: There is some confusion as to where Joao Rick collected his material,

which Watling & de Meijer (1997) have addressed. This fungus has been

previously illustrated (e.g., de Meijer 2008, fruiting body; Watling & de Meijer

1997, basidiospores) and described in detail (e.g., Singer 1948, 1952, Watling

& de Meijer 1997).

Fic. 1. Phylogenetic tree based on maximum parsimony (MP) analyses of 28S ribosomal DNA

sequences of two isolates of Neopaxillus echinospermus with representative crepidotaceous and

inocybaceous taxa. Agaricus aff. campestris and Deconica montana chosen as outgroups. Support at

nodes shown as MP bootstrapping/maximum likelihood bootstrapping values.

Neopaxillus (Crepidotaceae) ...

rae ~ Crepidotus subfulviceps FJ947116

-— Crepidotus mollis AF205677

83/100 |

S2l--

--/52) — Crepidotus fraxinicola AF205697

— Crepidotus lundellii AF205705

— | — Crepidotus versutus AF205695

—— Crepidotus subverrucisporus AF367948

~~ Crepidotus betulae AF205679

Crepidotus cf. applanatus AY380406

| Neopaxillus echinospermus AM1424

97/100

' Neopaxillus echinospermus AM1690

72159

, — Simocybe cf. serrulata GQ89298

88/100

--/54 Simocybe serrulata AY745706

[sd

a /64 | —— _ ,Simocybe sp. GQ892979

— ———~ Simocybe centunculus AF205707

L Simocybe americana AF205709

a -— Inocybe sp. GQ892974

75/50 |

91/-- | ~~ Inocybe asterospora AM882897

_ 95/94 ~~ Inocybe lilacina AY380385

83/72 _ | Inocybe petiginosa AF261510

| |

Inocybe (as Mallocybe) dulcamara AY380372

Agaricus aff. campestris DQ110871

Deconica montana DQ470823

———— 10 changes

88 ... Watling & Aime

Discussion

Previous work showed the Crepidotaceae to consist of two saprobic

agaricoid genera — the predominantly pleurotoid genus Crepidotus (Fr.)

Staude and Simocybe P. Karst. (which contains both pleurotoid and centrally

stipitate forms; Aime et al. 2005) — and several cyphelloid genera (Singer 1986,

Bodensteiner et al. 2004). Our results support the findings of Vizziniet al. (2012)

in placing Neopaxillus as a sister genus to Crepidotus. However, that study was

inconclusive in its treatment of Simocybe, whereas the latter is supported within

Crepidotaceae in our analyses. The inclusion of Neopaxillus expands the familial

concept to include a genus composed entirely of centrally stipitate, terrestrial

species with unknown ecology. Although some species of Crepidotus and

Simocybe can be found on bare or moss-covered soil, they generally fruit on

well decomposed large diameter logs, as well as fallen branches and sometimes

herbaceous and twiggy material. One question that has not been resolved is

whether Neopaxillus is ectomycorrhizal, although its present affinities would

not suggest this life-strategy, and at least one other species (i.e., N. plumbeus

Singer & Lodge) was not collected near any ectomycorrhizal host trees (D. Jean

Lodge, pers. comm.). However, the sister-relationship between Crepidotaceae

and the ectomycorrhizal Inocybaceae (Matheny et al. 2006) admits the

possibility that Neopaxillus is also ectomycorrhizal. A placement in Paxillaceae

might have hinted at an ectomycorrhizal role, although in the sense of Singer

(1951) this family contained both saprobes and ectomycorrhizal associates.

Singer (1952) wrote “on the ground in woods accompanying rivers usually in

small groups but never densely clustered.” Our present collections are from the

Atlantic rainforest in mixed ombrophilous forest with Araucaria angustifolia at

800-900 m above sea level, which basically agrees with Singer’s observation for

Argentina. He also noted, “fruiting in summer and fall” Watling & de Meijer

(1997) indicated that de Meijer’s collections occurred between December

(summer) and May (fall) in Brazil.

A hint to the relationship of Neopaxillus and the Crepidotaceae may be sought

in the observations of Singer, who noted “an actual affinity between Neopaxillus

and Tubaria section Thermophila, but even so, they are not congeneric, but can

be put into the same family” (Singer 1951). Indeed, Tubaria thermophila has

subsequently been shown to belong to Crepidotaceae, along with several other

stipitate species formerly placed in Melanomphalia M.P. Christ. (Aime et al.

2002, 2005, 2009). This refuted Christiansen’s suggestion that Melanomphalia

might be related to the Gomphidiaceae (Christiansen 1936), which, like

Paxillaceae, is now also placed within Boletales. Horak (1980) also hinted

at a relationship between Neopaxillus and Crepidotus in his descriptions of

N. bryogenus E. Horak and C. velutinoaffinis Singer, both agarics from South

America, although spore ornamentation is very different between the former

Neopaxillus (Crepidotaceae) ... 89

and the majority of Crepidotus spp. Indeed the illustrations in Horak (1980)

for both species resemble those depicted for Melanomphalia inocyboides

Singer and M. viscosa Singer, respectively, which leaves one to suspect that

N. bryogenus may not be assignable to Neopaxillus based on the type species.

Three additional Neopaxillus species are known: N. plumbeus from Puerto Rico

(Singer & Lodge 1989), N. reticulatus (Petch) Pegler from Sri Lanka (Pegler

1986), and N. dominicanus Angilini & Vizzini from the Dominican Republic

and Mexico.

Acknowledgments

Our thanks are offered to Andre de Meijer without whose field work and collections

this study could not have been undertaken. We thank Drs. Terry Henkel and D. Jean

Lodge for critical edits to a previous version of this manuscript. Molecular data were

supported in part by NSF DEB 0732968 to MCA.

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http://dx.doi.org/10.1017/S0960428600004042

MY COTAXON

http://dx.doi.org/10.5248/126.91

Volume 126, pp. 91-96 October-December 2013

New records of Aspicilia species from China

SHU-XIA LI, XING-RAN Kou & QIANG REN"

College of Life Sciences, Shandong Normal University Jinan, 250014, P. R. China

* CORRESPONDENCE TO: rengiang@sdnu.edu.cn*

ABSTRACT — As a result from our studies on the lichenized ascomycete genus Aspicilia, the

species A. persica, A. verrucigera, and A. subdepressa are reported for the first time from

China.

Key worps — lichen, taxonomy

Introduction

Aspicilia A. Massal. (Megasporaceae, Pertusariales, Ascomycota), a lichen

genus with worldwide distribution, includes over 200 species (Kirk et al. 2008).

Forty-one species have previously been reported from China (Wei 1991, Abbas

& Wu 1998, Sohrabi et al. 2010a). Our recent studies on Aspicilia establish

that three additional species occur in China: A. persica, A. verrucigera, and A.

subdepressa.

Materials & methods

The specimens studied are preserved in SDNU (Lichen Section of the Botanical

Herbarium, Shandong Normal University). Specimen morphology and anatomy

were examined using a dissecting microscope (Olympus SZ51) and a compact light

microscope (Olympus CX41). The lichen substances were detected by standardized thin

layer chromatography techniques (TLC) with solvent C (Orange et al. 2010). Photos

of the morphology were taken with an Olympus SZX16 stereomicroscope and of the

anatomy with an Olympus BX61 compound microscope with DP72.

New records

Aspicilia persica (Mill. Arg.) Sohrabi, Sauteria 15: 467. 2008. FIG. 1

THALLUS crustose, rimose to areolate, + orbicular, subradiate, 2—5.5(—7) cm

in diam., + 0.5 mm thick; areoles angular to irregular, 0.7-1.2(-1.5) mm in

diam., convex, continuous, fissured, cracks lighter towards the thallus edge;

92 ... Li, Kou & Ren

FicurE 1. Aspicilia persica (Z.L. Huang 20106405). A. Habitus; B. Apothecia; C. Apothecial

anatomy; D. Ascus and ascospores after I treatment; E. Conidia. Scale bars: A = 2 mm; B = 250 um;

C =50 um; D = 20 um; E= 10 um.

prothallus sometimes present, black. UPPER SURFACE white-gray to white,

matt, densely pruinose. APOTHECIA prominent, aspicilioid, usually solitary in

the areoles, 0.3-0.9 mm in diam.; thalline margin prominent, usually forming

a black rim, or sometimes somewhat paler due to the dense pruina; pisc black,

with dense white pruina; EPITHECIUM olive-green to olive-brown, K+ brown,

N+ green; HYMENIUM hyaline, I+ blue, 125-167.5 um high; pARAPHYSES

separating in KOH, moniliform; suBHYMENIUM and HYPOTHECIUM colorless,

I+ blue, together 62.5-87.5 um thick, without algae below the hypothecium;

ASCI clavate, Aspicilia-type, 8-spored; AscosporEs hyaline, simple, ellipsoid,

17.5-22.5x13-15 um; ConrpIA filiform, 15-25x 0.8-1 um.

SPOT TESTS — Medulla K+ yellow, C-, I-, P+ orange.

SECONDARY METABOLITES — Stictic acid (major), norstictic acid (minor).

SPECIMENS EXAMINED — CHINA. XINJIANG. UrRumai city, Mt. Tianshan, Glacier

No.1, alt. 3800 m, on siliceous rock, 27 Aug. 2011, Z.L. Huang 20106405 (SDNU); L. Li

20125895 (SDNU).

ComMENTS — Aspicilia persica is characterized by convex areoles, medium-

sized ascospores, prominent thalline margins, and black disks covered by dense

white pruina. Aspicilia boykinii Owe-Larss. & A. Nordin, which also has a white

Aspicilia spp. new to China... 93

or gray-white thallus and prominent thalline margins, looks like A. persica but

differs in its smaller spores, thin thalline margin, flat or (sometimes) convex

areoles, and white or gray apothecial rims. Aspicilia candida (Anzi) Hue,

another species with white upper surface, differs from A. persica by having a

rather low (75-85 um) hymenium and submoniliform paraphyses. Circinaria

calcarea, a subradiate and calciferous species with white thalli, also resembles

A. persica but has 4-spored asci, shorter conidia, and a K- thallus.

Aspicilia persica is a saxicolous species, known previously only from Iran

(Szatala 1957, Seaward et al. 2004, Sohrabi et al. 2010b).

Aspicilia verrucigera Hue, Nouv. Arch. Mus. Hist. Nat., 5 Sér., 2: 48. 1912. FIG. 2

THALLUS crustose, distinctly verrucose-areolate in the center, sometimes

rimose at the edge, 2-5 cm in diam., + 0.5 mm thick; AREOLEs angular to

irregular, 0.5-1 mm in diam., contiguous, convex, rough; UPPER SURFACE gray

to gray-brown, matt or somewhat shiny; prothallus present, black to brown-

black, 0.1-0.5 mm wide. APOTHECIA 0.2-0.5(-0.7) mm in diam., aspicilioid,

1-3 per areole, round or angular; thalline margin thin, slightly raised,

concolorous with the thallus, sometimes forming a white rim when young;

FiGuRE 2. Aspicilia verrucigera (J.Z. Zhao 20050703-9). A. Habitus; B. Apothecia; C. Apothecial

anatomy; D. Ascus and ascospores after I treatment; E. Conidia. Scale bars: A = 2 mm; B = 500 um;

C =50 um; D = 20 um; E= 10 um.

94 ... Li, Kou & Ren

pisc black, bare or with slight pruina; EPITHECIUM olive-green to olive-brown,

K+ brown, N+ green; HYMENIUM hyaline, I+ blue to green-blue, 87.5-125

uum high; PARAPHYSES separating in KOH, moniliform; suBHYMENIUM and

HYPOTHECIUM colorless, I+ blue, together 37.5-50 um thick, without algae

below the hypothecium. Ascr clavate, 8-spored, Aspicilia-type; ASCOSPORES

hyaline, simple, ellipsoid, 12.5-17.5x7.5-10 um; conip1a filiform, 15-22.5x

0.8-1 um.

SPoT TESTS — Medulla K+ yellow, C-, I-, P+ orange.

SECONDARY METABOLITES — Stictic acid (major), norstictic acid (minor).

SPECIMENS EXAMINED — CHINA. XINJIANG, KANAst, alt. 1350 m, on siliceous rock,

3 Jul. 2005, J.Z. Zhao 20050703-9 (SDNU); QINGHAI, MENYUAN COUNTY, Fengxiakou,

alt. 3300 m, on siliceous rock, 3 Aug. 2007, Y.D. Du 20070248 (SDNU).

ComMENTs — Aspicilia verrucigera is characterized by its numerous small

apothecia and the presence of stictic acid. Morphologically, this species is easily

confused with Aspicilia cinerea (L.) Kérb., which has a gray K+ red thallus and

norstictic acid as major metabolite. Circinaria caesiocinerea, which is similar to

A. verrucigera in thallus color and thalline margin, differs by its larger spores,

short conidia, and the presence of aspicilin. Aspicilia dudinensis (H. Magn.)

Oxner also resembles A. verrucigera in its verrucose or prominent areoles but

is easily separated by the K+ orange thallus reaction, a different exciple color,

and smaller spores.

Aspicilia verrucigera is distributed on siliceous rocks from sea level to the

mountains and has been reported from Europe (Magnusson 1939) and North

America (Owe-Larsson et al. 2007).

Aspicilia subdepressa (Nyl.) Arnold, Verh. Ges. Wien 19: 611. 1869. FIG. 3

THALLUS crustose, rimose, areolate, areoles 0.2-1.2 mm in diam., + 0.3 mm

thick, angular to irregular, or sometimes round, contiguous, smooth, somewhat

convex, indistinctly lobed, and the edge of the areoles is often slightly lifted

upward; UPPER SURFACE gray to green-gray, sometimes brownish gray, matt;

prothallus absent. APoTHEcIA 0.3-0.7 mm in diam., aspicilioid, ordinarily

solitary, or 2 per areole; pisc brown or black, not pruinose; thalline margin

prominent when old, concolorous with the thallus; EprrHectum brown or

olive-brown, K+ brown, N+ green; HYMENIUM hyaline, colorless, 100-150 um

high, I+ blue; PARAPHYSEs submoniliform to moniliform, separating in KOH;

SUBHYMENIUM and HYPOTHECIUM colorless, I+ blue, together 25-37.5(-50)

um thick, with algae below the hypothecium. Asc clavate, 8-spored, Aspicilia-

type; ascospores hyaline, simple, ellipsoid, (12.5-)15-20(-22.5) x (7.5-)10-

12.5(-15) um, conrpia filiform, straight or slightly curved, 7.5-12 x 0.8-1 um.

SPOT TESTS — Medulla K+ yellow, C-, I-, P+ orange.

SECONDARY METABOLITES — Stictic acid.

Aspicilia spp. new to China... 95

FiGuRE 3. Aspicilia subdepressa (Q. Ren 2011304). A. Habitus; B. Apothecia; C. Apothecial

anatomy; D. Ascus and ascospores after I treatment; E. Conidia. Scale bars: A = 2 mm; B = 250 um;

C= 100 um; D = 20 um; E= 10 um.

SPECIMEN EXAMINED — CHINA. HEILONGJIANG. WUDALIANCHI CITY, Wudalianchi,

Shihai, alt. 4400 m, on rock, 14 Jul. 2011, Q. Ren 2011304 (SDNU).

ComMENTs — Aspicilia subdepressa is characterized by its smooth grayish

thallus, short conidia, and the presence of stictic acid. Aspicilia cupulifera

(H. Magn.) Oxner is morphologically similar to A. subdepressa but has a

K+ nebulous yellow exciple, intensely branched paraphyses, and a lower

hymenium. Aspicilia proluta (Nyl.) Hue has several features in common with

A. subdepressa, such as a smooth thallus, short conidia, and the same major

metabolite, but it is distinguished by its gray or bluish thallus, different habitat,

and lack of prominent thalline margin.

Aspicilia subdepressa has been reported from France, from Pyrénées-

Orientales (Roux et al. 2011) and Massif Central (Magnusson 1939), where it

grows on rocks.

Acknowledgements

The project was financially supported by the National Natural Science Foundation of

China (31100011) and the Program for Scientific Research innovation team in Colleges

and Universities of Shandong Province. The authors are grateful to Dr. A. Nordin

96 ... Li, Kou & Ren

(Museum of Evolution, Botany, Uppsala University) and Dr. M. Sohrabi (Department

of Biotechnology, Iranian Research Organization for Science and Technology) for the

professional advice and great help during the study. The authors thank Dr. Harrie J. M.

Sipman (Botanischer Garten & Botanisches Museum, Freie Universitat Berlin, Konigin-

Luise-Str. 6-8, D-14195 Berlin, Germany) and Dr. A. Aptroot (ABL Herbarium, Soest,

The Netherlands) for presubmission reviews.

Literature cited

Abbas A, Wu JN. 1998. Lichens of Xinjiang. Urumgi, Sci-Tech & Hygiene Publishing House of

Xinjiang.

Kirk PM, Cannon PF, Minter DW, Stalpers JA (eds). 2008. Dictionary of the fungi, 10th edition.

Wallingford: CAB International.

Magnusson AH. 1939. Studies in species of Lecanora, mainly the Aspicilia gibbosa group. K.

Svenska Vetensk.- Akad. Hand. Ser. 317(5): 1-182.

Orange A, James PW, White FJ. 2010. Microchemical methods for the identification of lichens. 2nd

edition. London: British Lichen Society.

Owe-Larsson B, Nordin A, Tibell L. 2007. Aspicilia. pp. 61-108, in: TH Nash III et al. (eds). Lichen

flora of the greater Sonoran desert region, Vol. 3. Tempe, Lichens Unlimited, Arizona State

University.

Roux C, Nordin A, Tibell L, Sohrabi M. 2011. Quelques espeéces d’Aspicilia peu connues ou

nouvelles des Pyrénées-Orientales (France). Bull. Soc. linn. Provence, n° spécial 14: 177-227.

Seaward MRD, Sipman HJM, Schultz M, Maassoumi AA, Haji Moniri Anbaran M, Sohrabi M.

2004. A preliminary lichen checklist for Iran. Willdenowia 34: 543-576,

Seaward MRD, Sipman HJM, Sohrabi M. 2008. A revised checklist of lichenized, lichenicolous and

allied fungi for Iran. Sauteria 15: 459-520. http://dx.doi.org/10.3372/wi.34.34218

Sohrabi M, Owe-Larsson B, Nordin A, Obermayer W. 2010a. Aspicilia tibetica, a new

terricolous species of the Himalayas and adjacent regions. Mycological Progress 9: 491-499.

http://dx.doi.org/10.1007/s11557-010-0656-7

Sohrabi M, Sipman H, Toghranegar Z, Nejadsattari T. 2010b. A contribution to the lichenized

mycota of Zanjan province, Iran. Iran. J. Bot. 16(1): 125-129.

Szatala O. 1957. Prodromus einer Flechtenflora des Irans. Annales Historico-Naturales Musei

Nationalis Hungarici, ser. nov. 8: 101-154.

Wei JC. 1991. An enumeration of lichens in China. Beijing: International Academic Publishers.

MY COTAXON

http://dx.doi.org/10.5248/126.97

Volume 126, pp. 97-108 October-December 2013

Ambomucor gen. & spp. nov. from China

RU-YONG ZHENG* & XIAO-YONG LIU

State Key Laboratory of Mycology, Institute of Microbiology,

Chinese Academy of Sciences, Beijing 100101, China

* CORRESPONDENCE TO: Zhengry@im.ac.cn

ABSTRACT — Ambomucor is described as a new genus containing A. seriatoinflatus sp. nov.

(type), A. seriatoinflatus var. brevior var. nov., and A. clavatus sp. nov. Morphological

characteristics for recognizing and differentiating these new taxa are discussed, and full

descriptions, line drawings, and a key to taxa are provided.

KEYWORDS — taxonomy, morphology, Mucorales, new species, new variety

Introduction

In the summer of 2003, an interesting mucoraceous fungus was isolated

from a soil sample collected in Inner Mongolia. Morphological examination

revealed the fungus to represent a new species Ambomucor seriatoinflatus,

described here as the type species of a new genus, Ambomucor. One month

later, a second strain of this new genus was isolated from Xinjiang Province

and is described here as a new variety of the type species, A. seriatoinflatus

var. brevior. In addition, a fungus from Inner Mongolia, originally identified

as Zygorhynchus moelleri Vuill., is described here as new species, A. clavatus.

Since then, another new species (from a dung sample collected in Tibet) and

another new variety of A. seriatoinflatus (eleven strains from Tibet, and one

from Shaanxi Province) have been found by the second author, to be described

elsewhere.

Materials & methods

Isolations

Fungi were isolated from soil samples using the soil plate method of Warcup (1950).

Cultures

Living cultures of the three taxa studied are preserved in the Culture Collection of

the State Key Laboratory of Mycology (with an “Am-” prefix) and the China General

98 ... Zheng & Liu

Microbiological Culture Collection Center (with a “CGMCC-” prefix). Dried cultures

of the type strains are deposited in the Herbarium Mycologicum Academiae Sinicae

(HMAS). These culture collections and herbarium all belong to the Institute of

Microbiology, Chinese Academy of Sciences, Beijing, China.

Media & cultivation

Modified SMA (Hesseltine’s modified synthetic Mucor agar: dextrose 20 g, asparagine

2 g, KH,PO, 0.5 g, MgSO,-7H,O 0.25 g, thiamine chloride 0.5 mg, agar 20 g in 1000

mL distilled water, pH 7; Hesseltine & Ellis 1973) and PDA adjusted to pH 7 were

used for morphological studies. PDA adjusted to pH 7 was also used for establishing

temperature-growth relationships and for mating experiments. Cultivation period and

temperature were (i) 5-7(-10) days at 18-20°C for morphological studies; (ii) 4-7 days

at 25-37°C for determining the temperature maximum; and (iii) 7-14 days at 18-25°C

for mating experiments.

Results

Maximum growth temperature

The three Ambomucor strains described in this paper were tested twice for

their maximum growth temperature. The maximum growth temperatures of

both A. seriatoinflatus var. seriatoinflatus and A. seriatoinflatus var. brevior are

30°C and that of A. clavatus is 33°C. As shown by our studies on Cunninghamella

(Zheng & Chen 2001) and Rhizopus (Zheng et al. 2007), different varieties of

the same species have a similar maximum growth temperature, while different

species have different ranges of maximum growth temperature.

Mating experiments

The 16 living cultures of Ambomucor were crossed in all combinations, but

zygospores were not produced.

Morphological studies

Within the Mucorales, the new genus is morphologically unique in producing

two kinds of sporangia (fertile and non-fertile) on the same or different

sporangiophores. The fertile sporangia are multispored and columellate, similar

to those in the Mucoraceae. Aborted sporangia, which occur in short to long

chains, are vesicle-like, produce no spores, and lack a columella.

Taxonomy

Ambomucor is distinguished from all other genera of the Mucorales

(Alexopoulos et al. 1996, von Arx 1982, Benny & Benjamin 1991, Hesseltine

1955, Hesseltine & Ellis 1973, Tai 1979) in having simultaneously two kinds

of sporangia: fertile and aborted. We include Ambomucor in the Mucoraceae

based mainly on the characteristics of the fertile sporangia as well as its close

relationship with the genus Zygorhynchus, also in Mucoraceae.

Characteristics for dividing species or varieties in Ambomucor include 1) growing

or not growing above 30°C, 2) sporangiophore branching patterns, 3) sporangial

Ambomucor gen. & spp. nov. (China) ... 99

size and shape, 4) number of aborted sporangia in a chain, 5) presence or

absence, place of origination, and sterile outgrowths that may or may not be

spirally twisted from the sporangiophores bearing aborted sporangia, 6) size

and shape of the columellae in the fertile sporangia, 7) sporangiospore size and

shape, and 8) chlamydospore presence or absence.

Ambomucor R.Y. Zheng & X.Y. Liu, gen. nov.

MycoBank MB 518101

Differs from all known genera in the Mucorales by possessing both fertile and aborted

sporangia simultaneously on the same or different sporangiophores.

TYPE SPECIES: Ambomucor seriatoinflatus R.Y. Zheng & X.Y. Liu

EryMoLocy: Ambomucor, referring to the presence of two kinds of sporangia.

Myce. branching, nonseptate when young, septate in age, with aerial

hyphae. Stotons absent. RHIzo1Ds scanty, simple or branched, not opposite

sporangiophores. SPORANGIOPHORES arising directly from substrate mycelia

or from aerial hyphae, simple or more usually branched, of 3 main types:

(1) bearing normal fertile sporangia only, (2) bearing aborted sporangia only,

(3) bearing both fertile and aborted sporangia on the same sporangiophore.

ABORTED SPORANGIA either borne terminally on the main axes or branches

of the sporangiophores or repeatedly proliferating terminally or laterally

many times, transferring their contents forward to form chains of aborted

sporangia until a fertile sporangium is formed. FERTILE SPORANGIA borne

terminally on the main axes or branches of the sporangiophores, globose to

somewhat depressed-globose, non-apophysate, breaking or deliquescing, dark

brown after mature. COLUMELLAE well developed but only in fertile sporangia.

SPORANGIOSPORES also forming only in fertile sporangia. CHLAMYDOSPORES

present. ZyGOsPoREs unknown.

Ambomucor seriatoinflatus R.Y. Zheng & X.Y. Liu, sp. nov. FIGs 1, 2

MycoBAnk MB 518102

Differs from all known species of Mucorales by simultaneously possessing both fertile

and aborted sporangia on the same or different sporangiophores.

TyPE: People’s Republic of China. Inner Mongolia, Taipusiqi, 41.934593°N 115.282116°E,

soil, 16 July 2003, Xiao-yong Liu (Holotype HMAS 84258—1; ex-holotype culture

CGMCC 3.6665 = Am-1; GenBank AY743664. Isotypes HMAS 84258—2-8).

EryMo_oey: seriatoinflatus, referring to the chains of aborted sporangia.

Cotonigs on SMA and PDA at first subfloccose, then granulate, filling the

Petri dish in 6-7 (SMA) or 5-6 (PDA) days at 18°C, about 1-3 mm high, at

first white, then ‘Drab Gray’ to ‘Light Gray’ (Ridgway XLVI), reverse cream

to light yellowish brown. HypHae branching, nonseptate when young, septate

100 ... Zheng & Liu

FiGurE 1. Ambomucor seriatoinflatus var. seriatoinflatus (AS 3.6665, = Am-1, ex-holotype).

1. Upper portions of sporangiophores showing various kinds of constructions: (a) a long chain of

aborted sporangia with intercalary sporangiophores formed between two aborted sporangia; (b)

both fertile and aborted sporangia are formed on the same sporangiophore, note that the aborted

sporangia are in short chains; (c) same as (b) except that the aborted sporangial chains are longer,

also that rhizoids are formed at the base of a sporangial branch; (d) similar to (b) and (c) by forming

both fertile and aborted sporangia on the same sporangiophore except that aborted sporangia are

much more abundant than the fertile sporangium and the sporangiophore branches usually do not

originated from the tip of the former aborted sporangium; (e) a branched sporangiophore similar

to (b), (c), and (d) by forming both fertile and aborted sporangia on the same sporangiophore

except that rhizoids are formed at the apex of one of the aborted sporangia. 2. Chlamydospores

formed on the substrate mycelium.

Ambomucor gen. & spp. nov. (China) ... 101

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FIGURE 2. Ambomucor seriatoinflatus var. seriatoinflatus (AS 3.6665, = Am-1, ex-holotype).

1. Upper portions of sporangiophores, with fertile sporangia only. 2. Columellae of various shapes,

all with a small collar. 3. Sporangiospores.

in age, (1.5-)7-14(-18.5) um diam. SToLons absent. RHIzoIDs scanty, not

opposite sporangiophores; arising from various parts of the sporangiophores

or the aborted sporangia. SPORANGIOPHORES erect or recumbent, arising

directly from substrate or aerial hyphae, of 3 main types: (1) bearing only

102 ... Zheng & Liu

normal fertile sporangia, (2) bearing only aborted sporangia, (3) bearing

both fertile and aborted sporangia on the same sporangiophore; main axes

of the sporangiophores rarely simple, usually branched, 9-18.5(-22) um

diam.; primary branches 1-4(-10), single, in pairs, remaining simple, more

often branching monopodially or pseudo-verticillately, sometimes zigzag

and branching up to 7 times, (35-)98-437(-1000) um long, 4.5-13(-18)

um diam., rarely subcurved, usually equal or subequal in width throughout,

sometimes slightly subequal in width throughout, sometimes slightly narrowed

just beneath the terminal sporangia, hyaline to pale greenish brown, with or

without granular contents, becoming vacuolated in age. SEPTA in main axes

and branches of the sporangiophores usually present at the place of branching,

sometimes without definite position. ABORTED SPORANGIA may also form

terminally on the main axes and branches of sporangiophores, or may repeatedly

proliferate terminally or laterally, forming intercalary sporangiophores

between two aborted sporangia up to 20 times or more, transferring their

contents forward to form a short or long chain of aborted sporangia until

formation of a fertile and normal sporangium, globose and (18.5-)32-69 um

diam., or slightly elongate when proliferated, granular when young, vacuolate

in age, subhyaline. FERTILE SPORANGIA borne terminally on the main axes or

branches of the sporangiophores, globose, (14—)26-64(-88) um diam.; non-

apophysate, breaking or slowly dissolving, thin-walled, dark brown when

mature, broken pieces of wall pale grayish brown. COLUMELLAE formed only

in fertile sporangia, well developed, very regular in shape, larger ones mostly

globose and 16-51(-60) um diam., or ovoid to ellipsoid-ovoid and (21-)37-55

x (19-)32-48.5 um diam., smaller ones depressed-globose to sub-applanate

and 11.5-51 x 15-64.5 um, hyaline, some with grayish to brownish contents,

smooth, regularly with a distinct collar. SpPORANGIOSPORES forming only in

fertile sporangia, ellipsoid to oblong-ellipsoid, (2.5-)3.5-7 x (1.5—)2-2.5(-3.5)

um, (1-)2 guttulate, hyaline, grayish in mass, smooth, becoming vacuolated

in old cultures. STERILE OUTGROWTHS usually absent. CHLAMYDOSPORES rare,

single, in short chains, or in small masses, subglobose, ovoid or irregular, 5-9 x

4-7 um, hyaline to yellow. ZyGosporEs unknown.

Ambomucor seriatoinflatus var. brevior R.Y. Zheng & X.Y. Liu, var. nov. Fics 3, 4

MycoBank MB 518103

Differs from Ambomucor seriatoinflatus var. seriatoinflatus by its shorter sporangiospores

that in turn have much shorter, curved, verticillate branches.

TyPeE: People's Republic of China. Xinjiang, a glacial valley, 43.648994°N 85.117264°E,

soil, 3 August 2001, Xing-zhong Liu (Holotype HMAS 89521—1; ex-holotype culture

CGMCC 3.6784 = Am-2; GenBank KC108739. Isotypes HMAS 89521—2-8).

Erymo_oey: brevior, referring to the shorter sporangiospores.

Ambomucor gen. & spp. nov. (China) ... 103

FiGuRE 3. Ambomucor seriatoinflatus var. brevior (AS 3.6784, = Am-2, ex-holotype). 1. Upper

portions of sporangiophores, with both fertile and aborted sporangia on the same sporangiophore.

Note that the aborted sporangia are mostly in short chains, also that rhizoids are formed at the

apical portions of some of the aborted sporangia. 2. Short and finger-like rhizoids formed at the

base of the sporangiophore. 3. Simple or branched rhizoids formed on one side of a sporangiophore.

104 ... Zheng & Liu

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FiGuRE 4. Ambomucor seriatoinflatus var. brevior (AS 3.6784, = Am-2, ex-holotype). 1. Upper

portions of sporangiophores, mostly with fertile sporangia, some with aborted sporangia only.

Note that sterile outgrowths as well as septa may be formed on some of the sporangiophores.

2. Columellae of various shapes, all with a small collar. 3. Sporangiospores.

Cotontgs on SMA and PDA at first subfloccose, then granulate, filling the Petri

dish in 6-7 (SMA) or 5-6 (PDA) days at 18°C, about 1-3 mm high, at first

Ambomucor gen. & spp. nov. (China) ... 105

white, then “Drab Gray’ to ‘Light Gray’ (Ridgway XLVI), reverse cream to light

yellowish brown. HypHae branching, nonseptate when young, septate in age,

(1.5-)7-14(-18.5) um diam. SToLons absent. RHIZOIDs scanty, not opposite

sporangiophores; arising from various parts of the sporangiophores or the

aborted sporangia. SPORANGIOPHORES erect or recumbent, arising directly from

substrate or aerial hyphae, of 3 main types: (1) bearing normal fertile sporangia

only, (2) bearing aborted sporangia only, (3) bearing both fertile and aborted

sporangia on the same sporangiophore; main axes of the sporangiophores rarely

simple, usually branched, curved to subcurved, 9-18.5(-22) um diam.; primary

branches 1-4(-10), verticillately branched, (35-)98-437(-500) um long,

4.5-13(-18) um diam., curved to subcurved, usually equal or subequal in width

throughout, sometimes slightly subequal in width throughout, sometimes

slightly narrowed just beneath the terminal sporangia, hyaline to pale greenish

brown, with or without granular contents, becoming vacuolated in age. SEPTA

in main axes and branches of the sporangiophores usually present at the place

of branching, sometimes without definite position. ABORTED SPORANGIA may

also borne terminally on the main axes and branches of sporangiophores,

or may repeatedly proliferated terminally or laterally, forming intercalary

sporangiophores between two aborted sporangia, up to 20 times or more,

transferring their contents forward to form a short or long chain of aborted

sporangia until formation of a fertile and normal sporangium, globose and

(18.5-)32-69 um diam., or slightly elongate when proliferated, granular when

young, vacuolated in age, subhyaline. FERTILE SPORANGIA borne terminally on

the main axes or branches of the sporangiophores, globose, (14—)26-64(-88)

um diam.; non-apophysate, breaking or slowly dissolving, thin-walled, dark

brown when mature, broken pieces of wall pale grayish brown. COLUMELLAE

formed only in fertile sporangia, well developed, very regular in shape, mainly

depressed globose to applanate, sometimes oblong-ovoid, rarely globose,

hyaline, some with grayish to brownish contents, smooth, regularly with a

distinct collar. SPORANGIOSPORES forming only in fertile sporangia, ellipsoid

to oblong-ellipsoid, (2.5-)3.5-5 x (1.5-)2-2.5(-3.5) um, (1-)2-guttulate,

hyaline, grayish in mass, smooth, becoming vacuolated in old cultures.

STERILE OUTGROWTHS absent or rarely present and zigzag when present.

CHLAMYDOSPORES rare, single, in short chains, or in small masses, subglobose,

ovoid or irregular, 5-9 x 4-7 um, hyaline to yellow. ZyGospores unknown.

Ambomucor clavatus R.Y. Zheng & X.Y. Liu, sp. nov. FIG 5

MycoBank MB 518402

Differs from Ambomucor seriatoinflatus by its ellipsoid aborted sporangia, mostly

applanate sporangial columellae, frequent spirally twisted sterile outgrowths, and higher

maximum growth temperature.

106 ... Zheng & Liu

Type: People’s Republic of China. Inner Mongolia, Aer Mountain, 47.152486°N

119.976711°E, soil under a larch tree, 16 August 1991, Z.M. Sun (Holotype HMAS

240173—1; ex-holotype culture CGMCC 3.5877 = Am-3; GenBank KC108740. Isotypes

HMAS 240173—2-8).

EryMo_oey: clavatus, referring to the shape of the aborted sporangia.

Cotontgs on SMA and PDA at first subfloccose, then granulate, filling the Petri

dish in 6-7 (SMA) or 5-6 (PDA) days at 18°C, about 2-4 mm high, at first white

then slightly grayish white, reverse dirty white. HypHAE branching, nonseptate

when young, septate in age; 2-4.5 um diam. STOLons absent. RHIZOIDs scanty,

not opposite sporangiophores or arising from the aborted sporangia, scarcely

arising from the lower part of the sporangiophores. SPORANGIOPHORES erect

or recumbent, arising directly from substrate or aerial hyphae, of 2 main

types: (1) bearing aborted sporangia only, (2) bearing both fertile and aborted

sporangia on the same sporangiophore; main axes of the sporangiophores

substraight to recurved, rarely simple, usually branched, equal or subequal

in width throughout, sometimes slightly narrowed just beneath the terminal

sporangia, grayish to light brownish-gray, sometimes with granular contents in

the upper part, 5.5-14 um diam. and reaching a length of 380-762 um; primary

branches 1-3, remaining simple, repeatedly branching 1-2 times, or forming

aborted sporangia in a chain of 2-6. SEPTA usually absent, very rarely present at

the branches 40-60 um below the sporangia. ABORTED SPORANGIA also borne

terminally on the main axes or branches of the sporangiophores, or repeatedly

proliferating terminally or laterally forming intercalary sporangiophores

between two aborted sporangia, usually not exceeding 7 in number, transferring

their contents forward to form a chain of aborted sporangia until formation of

a fertile sporangium, globose and 18.5-41.5 um diam., or ovoid to ellipsoid

and 23-74 x 14-32 um, granular when young, vacuolate in age, subhyaline

to gray in color. FERTILE SPORANGIA borne terminally on the main axes or

branches of the sporangiophores, globose, 27.5-78 um diam., non-apophysate,

breaking and usually not dissolving, thin-walled, at first gray, then grayish-

brown when mature. COLUMELLAE formed only in fertile sporangia, well

developed, mostly applanate to depressed globose and (4-)9-25 x (6-)12.5-34

um, rarely subglobose and 14-33 um diam., light grayish-brown, often with a

large and distinct collar. SPORANGIOSPORES forming only in fertile sporangia,

oblong-ellipsoid, 2.5-4.5(-5.5) x 2-2.5 um, (1-)2-guttulate, hyaline, grayish

in mass, smooth, becoming vacuolate in old cultures. STERILE OUTGROWTHS

abundant, mycelium-like, growing out from the tip of the aborted sporangia

or from the branches of sporangiophores, spirally twisted. CHLAMYDOSPORES

absent. ZyYGOsPOREs unknown.

Ambomucor gen. & spp. nov. (China) ... 107

FiguRE 5. Ambomucor clavatus (AS 3.5877, = Am-3, ex-holotype). 1. Upper portions of

sporangiophores, mostly with both fertile and aborted sporangia on the same sporangiophore, a

few with aborted sporangia only. Note that the aborted sporangia are either subglobose or more

frequently ovoid to elliptic-ovoid and spirally twisted sterile outgrowths are formed from the

upper portion of the aborted sporangia or less frequently from the branches of sporangiophores.

2. Columellae of various shapes, each with a large collar. 3. Sporangiospores.

108 ... Zheng & Liu

Key to the taxa of Ambomucor

1. Aborted sporangia globoid, ovoid to ellipsoid; columellae of the fertile sporangia

mostly applanate, rarely globose to subglobose; sterile outgrowths frequent,

often spirally twisted; maximum growth temperature 33°C.......... A. clavatus

1. Aborted sporangia globose to subglobose; columellae of the fertile sporangia

mostly globose to subglobose, or ovoid-ellipsoid, smaller ones applanate;

sterile outgrowths absent or rare, not spirally twisted when present;

Maxinuny srowth temperature SOO C aan. « accor a Ath gerd dt ooo a dt eoe-g aia td areas y

2. Aborted sporangia in chains of 10(-20); sporangiophores never verticillately

branched; sterile outgrowths not in zigzags, usually absent; when globose

columellae reaching 60 um diam.; sporangiospores (2.5—)3.5-8 x

O15 =) 22,503.75) HT Swiss Bye, Deets A. seriatoinflatus var. seriatoinflatus

2. Aborted sporangia usually not exceeding chains of 6; sporangiophores sometimes

verticillately branched; sterile outgrowths when present often in zigzags; when

globose columellae not exceeding 42 um diam., sporangiospores (2.5-)3.5-5 x

DS (AVE No rene retlna te lS hte oa teins SEN A. seriatoinflatus var. brevior

Acknowledgements

The project was partially supported by the National Science Foundation of China

(Nos. 31070016, 31070019 and 31370068) and the Foundation of the Knowledge

Innovation Program of the Chinese Academy of Sciences (No. KSCX2-EW-J-6). Prof.

Xing-zhong Liu, Director of the State Key Laboratory of Mycology, who supplied us with

collections, is greatly appreciated. We also thank (both of this Laboratory) Ms Xiang-fei

Zhu for inking the line drawings and Ms Hong-mei Liu for testing the maximum growth

temperatures. Prof. Gerald L. Benny (University of Florida, USA) and Prof. Paul M. Kirk

(Kew, UK) are thanked for reviewing this paper.

Literature cited

Alexopoulos CJ, Mims CW, Blackwell M. 1996. Introductory mycology, 4th ed. John Wiley & Sons,

Inc., New York.

von Arx JA. 1982. On Mucoraceae s. str. and other families of the Mucorales. Sydowia 35: 10-26.

Benny GL, Benjamin RK. 1991. The Radiomycetaceae (Mucorales; Zygomycetes): II]. A new

species of Radiomyces, cladistic analysis and taxonomy of the family; with a discussion

of evolutionary ordinal relationships in Zygomycotina. Mycologia 83: 713-735.

http://dx.doi.org/10.2307/3760429

Hesseltine CW. 1955. Genera of Mucorales with notes on their synonymy. Mycologia 47: 344-363.

http://dx.doi.org/10.2307/3755457

Hesseltine CW, Ellis JJ. 1973. Mucorales. 187-217, in: GC Ainsworth et al. (eds). The Fungi IVB.

Academic Press, New York.

Tai FL. 1979. Sylloge Fungorum Sinicorum. Science Press. 1527 p.

Warcup JH. 1950. The soil-plate method for isolation of fungi from soils. Nature 166: 117-118.

http://dx.doi.org/10.1038/166117b0

Zheng RY, Chen GQ. 2001. A monograph of Cunninghamella. Mycotaxon 80: 1-75.

Zheng RY, Chen GQ, Huang H, Liu XY. 2007. A monograph of Rhizopus. Sydowia 59: 27-372.

MY COTAXON

http://dx.doi.org/10.5248/126.109

Volume 126, pp. 109-120 October-December 2013

Two new species of Rhytismataceae on fagaceous trees

from Anhui, China

Li CHEN’, D.W. MINTER’, SHI-JUAN WANG? & YING-REN LIN”

' School of Forestry & Landscape Architecture, Anhui Agricultural University,

West Changjiang Road 130, Hefei, Anhui 230036, China

? CAB International, Bakeham Lane, Egham, Surrey, TW20 9TY, UK

*CORRESPONDENCE TO: yingrenlin@yahoo.com

ABSTRACT — Lophodermium circinatum sp. nov. on Castanopsis eyrei and Terriera nitens

sp. nov. on Cyclobalanopsis myrsinifolia and C. pentacycla from China are described,

illustrated, and discussed. Type specimens are deposited in the Reference Collection of

Forest Fungi of Anhui Agricultural University, China (AAUF).

KEY worps — conservation, foliicolous fungi, morphology, parasitism, taxonomy

Introduction

During fieldwork in China studying leaf-inhabiting fungi on members of

the Fagaceae, three specimens of the ascomycete family Rhytismataceae were

collected. Examination of that material and comparison with species already

described led to the conclusion that one specimen represents a new species of

Lophodermium Chevall. and the other two represent a new species of Terriera

B. Erikss.

Materials & methods

For microscopic examination, pieces of leaf material with fruitbodies were selected

from each collection. The external appearance of the fungal colonies, zone lines,

ascomata, and conidiomata were observed with a dissecting microscope at 10—50x

magnification. After 10-15 min. in water, fruitbodies were cut into 10-15 um thick

vertical transverse sections using a freezing microtome (YD-202, China). These were

mounted in lactic acid or cotton blue (with pretreatment in water) onto microscope

slides, and examined. Squash mounts in water, lactic acid or cotton blue were made to

observe asci, ascospores and paraphyses. Gelatinous sheaths surrounding ascospores

and paraphyses were examined in water or 0.1% (w/v) cotton blue in lactic acid. The

colour of internal structures and ascus contents were observed in water. Measurements

were made from more than 30 asci, ascospores, paraphyses, conidiogenous cells and

110... Chen &al.

conidia for each specimen using material mounted in 5% KOH or Melzer’s reagent. Line

and point integrated drawings of external shapes and internal structures of fruitbodies

were prepared using a microscope drawing tube (Panasoianic XSJ-2, Japan).

Taxonomy

Lophodermium circinatum Li Chen bis & Y.R. Lin, sp. nov. FIGs 1-7

FUNGAL NAME FN570066

Differs from Lophodermium yangii by subcuticular ascomata, well-developed lips, much

shorter asci and ascospores, and paraphyses with circinate apices.

Type: China, Anhui, Qingliangfeng, Zhujiashe, alt. 180 m, on fallen leaves of Castanopsis

eyrei (Champ. ex Benth.) Hutch. (Fagaceae), 4 July 2006, T. Zhang & Y.R. Lin 2129

(Holotype AAUF 68937).

EryMoLoey: circinatum, referring to the paraphyses, which are circinate at the apex.

Cotontgs on both sides of leaves, forming subcircular or irregular, light yellow

to grayish-yellow spots, 10-18 mm diam., which tend to coalesce into larger

irregular shapes.

ZONE LINES frequent, gray-black and thin, entirely or partly surrounding

the paler areas, not always conspicuous.

ConipioMaTA on both sides of leaves, predominantly on the upper side,

crowded, sometimes with several coalescing. In surface view, conidiomata

90-175 x 80-135 um, almost circular to elliptical, slightly raising the

substratum surface, concolorous with the substratum surface or light yellow-

brown, dark-brown in the centre and at the perimeter line, often universally

brown after spore discharge which occurs through an apical ostiole. In vertical

section, conidiomata subcuticular. UPPER WALL 3-6 um thick, blackish brown,

composed of tiny angular cells. BAsaL WALL poorly developed, brown to dark

brown, consisting of 1—2 layers of angular and globose, thick-walled cells 2.5—4.5

um diam. SUBCONIDIOGENOUS LAYER 3.5—8 um thick, consisting of colourless,

thin-walled angular cells. TRICHOGYNES 20—28 x 2—3 um, cylindrical, tapering

towards the rounded apex, 4-6-septate. CONIDIOGENOUS CELLS 8-12 x 2-2.5

um, flask-shape, holoblastic, proliferating sympodially. Conrp1a colourless,

aseptate, smooth, cylindrical or elliptical, 3-4.5 x ca 1 um.

Ascomara in similar positions to conidiomata on the substratum, scattered,

occasionally with 2 coalescing in the paler areas. In surface view, ascomata

900-1750 x 360-650 um, elliptical to elongate-elliptical, straight or curved

to one side, ends obtuse or slightly acute, black, shiny, with a clearly marked

outline, moderately raising the substratum surface but clearly sunken near

the split, opening by a single longitudinal split which extends almost to the

ascomatal edge. Lips present. In median vertical section, ascomata subcuticular.

COVERING STROMA 20—28 um thick near the opening, slightly thinning towards

the edge, connecting to the basal stroma, composed mainly of textura angularis

Lophodermium & Terriera spp. nov. (China) ... 111

me m0,

Zea 29

At ¥

Fics 1-7. Lophodermium circinatum on Castanopsis eyrei. 1. A leaf bearing zone lines and

fruitbodies. 2. A zone line and fruitbodies observed under a dissecting microscope. 3. Ascoma in

median vertical section. 4. Conidioma in vertical section. 5. Portion of ascoma in median vertical

section. 6. Paraphyses, asci, and ascospores. 7. Trichogynes, conidiogenous cells, and conidia.

112 ... Chen & al.

with dark brown to blackish brown, thick-walled cells 3-5.5 um diam. Lip

CELLS well developed, arranged somewhat radially, 10-24 x 1.5-2.5 um,

cylindrical, the apical cells sometimes slightly wider, nearly colourless, with

black and markedly brittle tissue sticking to the outer side. BASAL STROMA dark

brown, composed of 1—2 layers of thick-walled, angular to globose cells 4-6

uum diam. The triangular area, visible in sections between the covering stroma

and the basal stroma at each edge of the ascoma, is filled with colourless,

gelatinous, thin-walled angular cells 4—9.5 um diam. SUBHYMENIUM 10-16

um thick, rather flat, consisting of colourless textura porrecta. PARAPHYSES

90-120 x 1.2—1.5 um, filiform, usually curved into uncinate or circinate shapes,

sometimes gradually widening to 2—2.5 um at the apex, and immersed in a ca

1.5 um thick gelatinous matrix. Asci ripening sequentially, 70-95 x 7—9 um,

cylindrical-clavate to cylindrical, somewhat long-stalked, thin-walled, apex

rounded, without circumapical thickening, J—, 8-spored. AscosPorEs arranged

in a fascicle, 40-55 x 1.2—1.6 um, filiform, colourless, aseptate, slightly tapered

towards the rounded base, and immersed in a gelatinous sheath ca 1 um thick.

ASSOCIATED ORGANISM, HABITAT, AND DISTRIBUTION: Castanopsis eyrei, on leaves.

Known only from the type locality, Anhui Province, China.

ComMENTS—Lophodermium is the largest genus of the Rhytismataceae.

IndexFungorum (2013) lists 282 specific epithets for the genus, of which

SpeciesFungorum (2013) accepts 135. Although exceptions exist, in the

Rhytismataceae individual species tend to occur on only conifers, only

dicotyledons, or only monocotyledons, while species on dicotyledons tend to

occur on either herbs or woody plants but not both. These patterns help limit

the taxa for comparison when assessing possible new species.

No Lophodermium has been described with type material on Castanopsis,

but 14 species from IndexFungorum have been recorded on trees of the

Fagaceae (Cannon & Minter 1986; Farr & Rossman 2013; Johnston 1989a, b,

2001; Lin et al. 2002). Lophodermium echinophilum Speg., L. gamundiae P.R.

Johnst., L. mahuianum P.R. Johnst., L. medium P.R. Johnst., L. petiolicola Fuckel,

L. punctiforme (Fr.) Fuckel, L. quercus S.K. Bose & E. Mull., and L. yangii Y.R.

Lin & C.L. Hou have type material on other fagaceous genera and are known

mainly in association with the Fagaceae. Lophodermium agathidis Minter

& Hettige, L. eucalypti (Rodway) P.R. Johnst., L. foliicola (Fr.) PE Cannon &

Minter, L. maculare (Fr.) De Not., L. mangatepopense P.R. Johnst., and L. minus

(Tehon) P.R. Johnst. were originally described from leaves of other woody plants

but have subsequently also been recorded in association with the Fagaceae.

None of the Lophodermium species with type material on the Fagaceae have

paraphyses with circinate apices. Lophodermium gamundiae, L. mahuianum,

and L. medium were all described on Nothofagus species from New Zealand or

southern South America. No anamorph or zone lines have been observed for

Lophodermium & Terriera spp. nov. (China) ... 113

L. gamundiae, and lengths of asci (160-190 um) and ascospores (60-80 um)

are much greater (Johnston & Park 2007) than for our new species. Johnston

(1989b) described L. mahuianum as never having zone lines or an anamorph,

and lengths of asci (130-175 um) are also much greater. Ascomata of L. medium

are often triangular in outline and its asci (100-140 um; Johnston 1989b) are

longer than those of L. circinatum. Recent evidence suggests that Nothofagus

should be placed in its own family separate from the Fagaceae and the natural

distribution of Nothofagus in South America and Australasia apparently

represents a single population divided when continental drift made Antarctica

too cold for plants. Similar distributions might be expected for species of the

Rhytismataceae evolving with Nothofagus. The present fungus is thus clearly

different from these three species.

Their protologues described both L. petiolicola and L. punctiforme as

occurring only on petioles and leaf nerves of deciduous species of Quercus

in Europe. In the north and west of the UK, L. petiolicola is common in this

habitat, and there are over 100 records of that species throughout Europe

and into Georgia (the most western part of Asia) mostly on Quercus but

also on Castanea. [One record on Sorbus is probably a misidentification of

L. aucupariae (Schleich.) Darker.] There are very few records of L. punctiforme,

all from Europe (Cybertruffle’s Robigalia 2013; GBIF 2013). It is possible that

these two names represent the same fungus. If so, L. punctiforme has priority

and would be the correct name.

Lophodermium echinophilum, described from Italy on spines of Castanea

cupules, may also be a synonym. The paraphyses of these species are not

circinate (Tehon 1935). Lophodermium quercus, described from India on leaves

of the evergreen Quercus leucotrichophora A. Camus, has septate ascospores

(Cannon & Minter 1986). Lophodermium yangii, described from China on

leaves of Cyclobalanopsis glauca (Thunb.) Oerst., has asci 95-130 um long. All

these species therefore also differ significantly from the present fungus.

Lophodermium agathidis, originally described from leaves of Agathis australis

(D. Don) Loudon (a conifer) has also been reported from a wide range of other

plants including Castanopsis eyrei (Lin et al. 2012), apparently an exception

to the general patterns described earlier. Developing ascomata of L. agathidis

have a characteristic pale central band on their surface, the asci are 120-170

uum, and the paraphyses do not have circinate apices. Lophodermium eucalypti,

which has been recorded on conifers, dicotyledons and monocotyledons, is

another apparent exception to that general pattern (Johnston 2001). While it

has paraphyses with circinate apices, it does not occur on paler areas of the

leaf, nor has it been observed with zone lines or conidiomata. It seems to be an

Australasian fungus that has also been recorded, perhaps as an introduction, in

North America.

114... Chen & al.

Lophodermium foliicola, a species of Europe and North America, typically

associates with members of the Rosaceae but has occasionally been recorded

from other dicotyledonous families. At least some records on the Fagaceae are

likely to represent early misidentifications by European mycologists attempting

to describe fungi on Nothofagus and other plants in the southern hemisphere.

Specimens on Crataegus monogyna Jacq. from the UK (e.g., IMI 296407) have

ascomata that are shorter than 1 mm, asci about 80 um long, ascospores about

55 um long, and paraphyses with complex curved and sometimes branched

apices. Lophodermium maculare, another species of Europe and North America,

typically occurs on dead leaves of Vaccinium species and has ascomata less than

1 mm long, asci 60-120 um long, ascospores 55—95 um long, and paraphyses

bent, but not circinate, at the apex. An anamorph has not been observed.

In the protologue, L. mangatepopense, a predominantly southern

hemisphere species, was described as lacking zone lines and conidiomata.

Asci were said to be 100-135 um long and paraphyses to have clavate, not

circinate, apices. Lophodermium minus has been observed on a wide range of

woody plants, mostly in the southern hemisphere, but with some records from

China. Paraphyses are uniformly narrow, sometimes branched at the apex, but

not circinate. Asci are 100-130 um long (Cabarroi & Minter 2005). Although

accepted in Lophodermium by SpeciesFungorum, L. minus was moved to

Terriera by Johnston (Ortiz-Garcia et al. 2003) and is discussed again under

that name later in the present work.

The other possible generalist Lophodermium species not yet recorded in

association with the Fagaceae is L. mangiferae Koord. As in L. agathidis, its

developing ascomata have a characteristic pale band on their surface. Asci

are 95-110 um long and ascospores 70—75 um long, while paraphyses tips

are widened but not circinate. Lophodermium circinatum clearly differs from

L. mangiferae and all other generalist species considered in the foregoing

paragraphs.

Terriera nitens Y.R. Lin, sp. nov. Figs 8-13

FUNGAL NAME FN570067

Differs from Terriera illiciicola by smaller ascomata, a well-developed excipulum,

and swollen paraphyses and from T: minor by the almost circular to broadly elliptical

ascomata, strongly sunken subhymenium, and the presence of conidiomata and

frequent zone lines.

Type: China, Anhui, Shitai, Dayan, alt. 1100 m, on leaves of Cyclobalanopsis myrsinifolia

(Blume) Oerst. (Fagaceae), 6 July 2006, T. Zhang & Y.R. Lin 2077 (Holotype AAUF

68885).

ETyMoLoey: nitens, referring to the shiny appearance of ascomata.

Cotonigs on both sides of leaves, forming irregular, light yellow bleached

spots, 4-10 mm diam.

Lophodermium & Terriera spp. nov. (China) ... 115

ZONE LINES frequent, grey-brown to black, thin, surrounding the paler

areas.

Conrp1omarta on both sides of leaves, mostly on the upper side, scattered,

occasionally confluent in groups of two or three. In surface view, conidiomata

150-210 um diam., circular or almost circular, applanate, dark brown in the

centre and on the perimeter line of the conidioma, gray-brown or almost

concolorous with the substratum surface elsewhere, discharging spores through

an apical ostiole. In vertical section, conidiomata subepidermal. UPPER WALL

composed of 3-4 layers of nearly colourless, angular cells 3—5.5 um diam.,

the cells around apical ostiole blackish brown. BAsAL WALL 7—12 um thick,

consisting of 2—3 layers of brown to dark brown thick-walled cells forming a

textura angularis. TRICHOGYNES 18—30 x 1.5—2 um, cylindrical, tapering to the

apex, 2-4-septate. CONIDIOGENOUS CELLS and CONIDIA not observed.

AscoMaTA in similar positions to conidiomata on the substratum,

crowded in the paler areas. In surface view, ascomata 240-320 x 190-250 um,

suborbicular or broadly elliptical, straight or slightly curved, black, shiny, with a

clearly marked outline, strongly raising the surface of the substratum, opening

by a single longitudinal split */,—’/, the length of the ascoma. In median vertical

section, ascomata subepidermal. CovERING STROMA 10-16 um thick near the

opening, becoming slightly thinner towards the edge, extending to the basal

stroma, composed of textura angularis with dark brown to blackish brown,

thick-walled cells 2—3.5 um diam. Along the edge of the ascomatal opening,

there is a 5—8 um thick, flat extension adjacent to the covering stroma which is

comprised of markedly black and brittle tissue with no obvious cellular structure.

BASAL STROMA dark brown, comprised of 2—3 layers of 2.5—5 um diam., thick-

walled angular cells. A layer of textura prismatica 8-17 um thick, composed

of nearly colourless or light brown cells 2.5—4.5 um diam. occurs between the

covering and basal stromata. ExcipuLuM 12-16 um thick near the opening,

becoming slightly thinner towards the base, arising from the inner layer of the

basal stroma, consisting of colourless textura porrecta. SUBHYMENIUM 5—9 um

thick, strongly concave to form a cup shape, consisting of textura porrecta.

PARAPHYSES colourless, thin-walled, smooth, filiform, aseptate, 95-150 x

1—1.2 um, sometimes swollen at the apex. Asci ripening sequentially, 85-128

x 4.5—6 um, narrow cylindrical, somewhat short-stalked, apex round, without

circumapical thickening, J—, 8-spored. Ascospores arranged in a fascicle,

68-115 x 0.8-1.2 um, filiform, colourless, aseptate, round at the apex, slightly

tapered towards the acute base, with a thin gelatinous sheath.

ADDITIONAL SPECIMEN EXAMINED — CHINA, Huser: Shengnongjia, on leaves

of Cyclobalanopsis pentacycla (Y.T. Chang) Y.T. Chang ex Y.C. Hsu & H.W. Jen

(Fagaceae), 10 July 2010, Y.R. Lin & G.J. Jia (AAUF 69308).

COMMENTS — Terriera was established for T. cladophila (Lév.) B. Erikss.

(Eriksson 1970). IndexFungorum (2013) lists 21 specific and two subspecific

116... Chen &al.

epithets in this genus, of which SpeciesFungorum (2013) accepts 16 species

and two varieties in Terriera, one epithet in Lophodermium, and expresses

no opinion about the remaining four epithets, all recent. At least three other

species have very recently been described in or transferred to this genus. Most

Terriera species were originally described in Lophodermium or other genera

before transfer to Terriera by Johnston (Johnston 2001; Ortiz-Garcia et al.

2003). Four Terriera species have been recorded associating with trees of the

Fagaceae: T. coacervata Y.R. Lin & Q. Zheng, T: illiciicola (S.J. Wang et al.)

Q. Zheng & Y.R. Lin, T! minor (Tehon) P.R. Johnst., and T! rotundata C.L. Hou

(Lin et al. 2005, 2012; Song et al. 2012; Zheng et al. 2012).

Terriera coacervata, described on leaves of Lithocarpus cleistocarpus

(Seemen) Rehder & E.H. Wilson, has ascomata at least 650 (mostly 1000-1800)

long, compared with our new species for which the maximum ascomal length

is 320 um. Ascomata of T: coacervata tend to coalesce to form complex multiple

structures, whereas those of T. nitens tend to remain single. There are no records

of T: coacervata on Cyclobalanopsis species.

Terriera illiciicola, described originally on [Illictum verum Hook. f.

(Schisandraceae), has also been recorded from the Fagaceae on Lithocarpus

cleistocarpus (Lin et al. 2012). Zheng et al. (2012) reported that T. illiciicola

does not produce conidiomata, which are abundant in T! nitens. Ascomata of T.

illiciicola are 280-360 x 260-300 um (Lin et al. 2012) or even larger (300-380 x

280-330 um; Zheng et al. 2012) while those of T’ nitens are 240-320 x 190-250

um so that, despite some overlap, those of T. illiciicola are distinctly larger. In

comparison with T. nitens, the excipulum of T! illiciicola is less well developed

and paraphyses are not swollen (Zheng et al. 2012). There are no records of

T. illiciicola on Cyclobalanopsis species.

Terriera minor (discussed above under the name Lophodermium minus) was

originally described from Desecheo Island (Puerto Rico) on Clusia rosea Jacq.

The holotype is lost, and the species has been neotypified with a collection on

Clusia sp. from Venezuela (Johnston 1989a). A recent description exists based

on Clusia specimens from Cuba (Cabarroi & Minter 2005). It differs from T.

nitens in having no conidiomata in vivo (although conidiomata have been

produced in pure culture) and in having more elongated ascomata (0.3-1 mm

long), which sometimes coalesce to form more complex structures. When

examined in vertical section, the ascoma of T. minor has a flat lower wall, a

very thin excipulum, and a distinctive flat black area adjacent to both sides

of the opening slit, whereas the lower wall in T. nitens is strongly curved, the

excipulum is thicker, and there is no flat black area adjacent to the opening slit.

Paraphyses of T: minor are unswollen and sometimes branched, where those of

T. nitens are often swollen but remain unbranched. There are no records of T.

minor on Cyclobalanopsis species.

Lophodermium & Terriera spp. nov. (China) ... 117

BANS

BBE NY

ENN

Fics 8-13. Terriera nitens on Cyclobalanopsis myrsinifolia. 8. A leaf bearing zone lines and

fruitbodies. 9. Zone lines and fruitbodies observed under a dissecting microscope. 10. Ascoma in

median vertical section. 11. Portion of ascoma in median vertical section. 12. Paraphyses, asci, and

ascospores. 13. Conidioma in vertical section.

118 ... Chen & al.

Terriera rotundata was described from Yunnan, China on dead fallen leaves

of Quercus sp. The protologue stated that conidiomata were not observed

while they are present in T: nitens. Ascomata are mostly 600-900 um long and

thus much larger than those of T. nitens. The two species are similar in having

ascomata that are more or less globular in vertical section, but ascospores

of T: nitens have thin gelatinous sheaths, which have not been observed for

T. rotundata. There are no records of T: rotundata on Cyclobalanopsis species.

Terriera nitens thus differs from all other species of Terriera that have been

observed on the Fagaceae.

Discussion

Lophodermium circinatum is known only from a single collection, and

Terriera nitens only from two collections augmented by some field observations.

As a result, very little can be said with confidence about their biology.

The associated plants, Castanopsis eyrei, Cyclobalanopsis myrsinifolia, and

C. pentacycla are evergreen trees with leathery leaves that decay slowly. This

type of substratum very often carries species of the Rhytismataceae. The main

leaf fall is in September and October, and by July of the following year these

fungi have produced pale areas on the leaf bearing zone lines, conidiomata with

trichogynes, conidiogenous cells and conidia (observed only in L. circinatum

but probably also produced by T! nitens), and ascomata with paraphyses,

asci, and ascospores. In the case of T’ nitens, ascomata with fully formed

ascospores are also sometimes observed on dead leaves still attached to the

tree. Trichogynes are rarely reported for the Rhytismataceae (probably because

they are easily overlooked), but their presence here suggests the anamorph

has a sexual function, something compatible with the few other reports about

anamorph biology in the family.

It is likely that colonization of new substrata is effected in humid condition

through wind-dispersed ascospores with a mucous sheath, which may help

adhesion. There is not enough information to determine which substrata

are colonized by the ascospores. It is possible that they infect leaves already

destined to senesce and fall in the following autumn, but that strategy is not

typical of rhytismataceous species on evergreen leaves. It is perhaps more likely

that the ascospores colonize young leaves produced in the spring of that year,

while the leaf cuticle is still soft, and then remain in the leaves as a symptomless

endobiont.

There is not enough information to determine any possible positive or

negative impacts these fungi might have on their associated trees. It is interesting

that there are very few collections. This may simply reflect a past lack of

attention to this substratum, but other cases are known in the Rhytismataceae

where a species may be genuinely rare and only very occasionally seen to fruit.

Lophodermium & Terriera spp. nov. (China) ... 119

Lophodermium neesii Duby is a good example of this. It occurs in Europe on

dead fallen leaves of Ilex aquifolium L., a very abundant tree, but has been

collected fewer than ten times in 100 years (Minter 2005). One great value of

the present work is that it draws attention to the huge diversity of fungi, an

important biodiversity resource too often overlooked.

A good place to start when evaluating the conservation status of leaf-

inhabiting fungi is to look at the status of the associated plants. Castanopsis

eyrei, Cyclobalanopsis myrsinifolia, and C. pentacycla have not yet been

evaluated for the IUCN Red List (2013). All three species occur in a range of

arboreta, but such collections rarely if ever protect the microfungi associated

with plants in their natural environment. Castanopsis eyrei is widely distributed

in China (GBIF 2013), and there is no immediate reason to suppose this plant is

endangered. GBIF (2013) holds very little information about the distribution of

Cyclobalanopsis myrsinifolia, and no information at all about C. pentacycla, and

for these two plants it is not possible to comment on their conservation status

in the wild. It is also important to evaluate threats to the ecosystems where

these fungi were collected. All three collecting sites for these two species are in

or close to nature reserves or other conservation areas, and the status of these

sites provides some protection, but in each case the reserve or conservation

area is only a small remnant of what was once a more extensive ecosystem.

There are also new possible threats. One of the three collection sites is in the

Shengnongjia region where the impact of big hydro-electric schemes has

aroused international attention (Watts 2011). More information about such

developments is needed before proper evaluations can be made of threats to

fungi in the region. With the present information, and with only one collection

of L. circinatum and only two of T. nitens, the only possible conservation status

evaluation for these fungi is, at present, Data Deficient.

Acknowledgments

The authors are grateful to Dr PR. Johnston and Dr M. Ye for serving as pre-

submission reviewers and to Dr Y.H. He for the identification of the host plants. This

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

31270065, 31170019, 30870014).

Literature cited

Cabarroi M, Minter DW. 2005. Terriera minor. IMI Description Sheets of Fungi and Bacteria No.

1660.

Cannon PF, Minter DW. 1986. The Rhytismataceae of the Indian subcontinent. Mycological Papers

Lspe 2123;

Chen JL, Lin YR, Hou CL, Wang SJ. 2012 [“2011”]. Species of Rhytismataceae on Camellia spp.

from the Chinese mainland. Mycotaxon 118: 219-230. http://dx.doi.org/10.5248/118.219

Cybertruffle’s Robigalia. 2013. [www.cybertruffle.org.uk/robigalia (viewed online on 25 March

2013)].

120 ... Chen & al.

Darker ED. 1932. The Hypodermataceae of conifers. Contributions from the Arnold Arboretum of

Harvard University 1: 1-131.

Eriksson B. 1970. On Ascomycetes on Diapensales and Ericales in Fennoscandia. Symbolae

Botanicae Upsalienses 19: 1-71.

Farr DF, Rossman AY. 2013. Fungal Databases, Systematic Mycology and Microbiology Laboratory.

ARS, USDA [http: //nt.ars-grin.gov/fungaldatabases/fungushost/FungusHost.cfm (viewed

online on 20 March 2013)].

GBIF (Global Biodiversity Information Facility). 2013. [www.gbif.org (viewed online on 25 March

2013)].

IndexFungorum. 2013. [www.indexfungorum.org (viewed online on 25 March 2013)].

IUCN Red List. 2013. [www.iucnredlist.org (viewed online on 25 March 2013)].

Johnston PR. 1989a. Lophodermium (Rhytismataceae) on Clusia. Sydowia 41: 170-179.

Johnston PR. 1989b. Rhytismataceae in New Zealand 2. The genus Lophodermium on indigenous

plants. New Zealand Journal of Botany 27 (2): 243-274.

Johnston PR. 2001. Monograph of the monocotyledon-inhabiting species of Lophodermium.

Mycological Papers 176: 1-239.

Johnston PR, Park D. 2007. Revision of the species of Rhytismataceae reported by Spegazzini from

South America. Boletin de la Sociedad Argentina de Botanica 42(1-2): 87-105.

Kirk PM, Cannon PF, Minter DW, Stalpers JA. 2008. Ainsworth & Bisby’s dictionary of the fungi,

10" ed. CAB International. Wallingford. 771 p.

Lin YR, Hou CL, Xu ZS, Li K. 2002. Two new species of Lophodermium (Rhytismatales,

Discomycetes) 147-150, in: Anhui Soc. PI. Path. et al., Tact. Future Integrated Pest Management.

China’ Agricultural Science & Technology Press. Beijing. 311 p.

Lin YR, Yu SM, He YF, Wang SJ. 2005. A new species and two new Chinese records of Lophodermium

Chev. (in Chinese). Mycosystema 24: 1-5.

Lin YR, Liu HY, Hou CL, Wang SJ. 2012. Flora fungorum sinicorum, vol. 40, Rhytismatales

(in Chinese). Science Press. Beijing. 261 p.

Minter DW. 2005. Lophodermium neesii. IMI Description Sheets of Fungi and Bacteria No. 1657.

Ortiz-Garcia S, 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

Song JF, Liu L, Li YY, Hou CL. 2012. Two new species of Terriera from Yunnan Province, China.

Mycotaxon 119: 329-335. http://dx.doi.org/10.5248/119.329

SpeciesFungorum. 2013. [www.speciesfungorum.org (viewed online on 25 March 2013)].

Tehon LR. 1935. A monographic rearrangement of Lophodermium. Illinois Biological Monographs

13(4): 1-151.

Watts J. 2011. Shennongijia exposes reckless development of China’s water resources. Guardian

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development (viewed online on 25 March 2013)].

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from Mt Huangshan, China. Mycotaxon 118: 311-323. http://dx.doi.org/10.5248/118.311

MY COTAXON

http://dx.doi.org/10.5248/126.121

Volume 126, pp. 121-126 October-December 2013

Three newly recorded species of Parasympodiella and Chalara

from China

X1A0-X1A Lr”, LI-Guo Ma?, JI-WEN XIA? & XIU-GUO ZHANG”

' Life Science Institute, Zunyi Normal College, Zunyi, Guizhou, 563002, China

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

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

ABSTRACT — Three new Chinese records of fungi were discovered from subtropical forests

of Guangxi Province, China. Parasympodiella laxa, P. eucalypti, and Chalara unicolor are

described and illustrated from specimens collected on dead branches and compared with

closely related species.

KEY worpDs — anamorphic fungi, taxonomy

Introduction

Parasympodiella was established by Ponnappa (1975) with P. laxa as the

type species. The genus is characterized by distinct, single conidiophores with

terminal and intercalary, integrated, indeterminate, sympodially extending

conidiogenous cells that produce catenate, cylindrical, thallic conidia by

disarticulations of the fertile hyaline branches. These characters separate

Parasympodiella from similar genera such as Sympodiella W.B. Kendr.

(Kendrick 1958), Bahusakala Subram. (Subramanian 1958), Polyscytalum

Riess (Riess 1853), and Neoscytalidium Crous & Slippers (Crous et al. 2006).

Parasympodiella currently includes 10 accepted species.

Rabenhorst (1844) introduced Chalara for the single species Torula

fusidioides Corda. The genus has been defined as having distinct, solitary or

loosely aggregated conidiophores with monophialidic, integrated, terminal,

determinate apical conidiogenous cells that produce catenate, endogenous,

cylindrical phialospores. Chalara is similar to Chaetochalara B. Sutton & Piroz.

(Sutton & Pirozynski 1965), Thielaviopsis Went (Went 1893), and Fusichalara

S. Hughes & Nag Raj (Hughes & Nag Raj 1973) in having phialidic conidiogenous

cells with a deep cylindrical collarette. Of the more than 130 species described

in Chalara, C. ampullula (Sacc.) Sacc., C. aotearoa Nag Raj & S. Hughes,

L22>.. x Se alt

C. cylindrosperma (Corda) S. Hughes, C. sinensis W.P. Wu, and C. ungeri Sacc.

have previously been reported from China (Zhuang 2001, 2005).

During a continuing survey of subtropical microfungi from the forests of

Guangxi Province, China, three new Chinese records of Parasympodiella and

Chalara were collected on decaying branches. They are described and illustrated

below.

Fic. 1. Parasympodiella laxa.

A. Colonies on natural substratum; B. Conidiophores and conidiogenous cells; C, D. Conidia.

Parasympodiella laxa (Subram. & Vittal) Ponnappa, Trans. Br. Mycol. Soc. 64: 344,

1975. FIG. 1

COLONIES on natural substrate effuse, brown to blackish brown, hairy.

Mycelium superficial and immersed, consisting of branched, septate, hyaline

to pale brown hyphae. ConrpiopHores distinct, solitary, unbranched,

erect, cylindrical, <780 um long, 1-9-septate; sterile part dark brown, with

New Parasympodiella and Chalara records for China ... 123

a thickened wall, 6-9 um wide, swollen base <18 um wide; fertile part grey-

brown, later becoming paler toward the apex. CONIDIOGENOUS CELLS terminal

and intercalary, integrated, indeterminate, with sympodial extensions and one

conidiogenous locus per cell, smooth, pale grey-brown, becoming hyaline

toward the apex, 42-78 x 6.3-9.4 um between conidiogenous loci. CONIDIA

thallic-arthric, forming in loose chains, hyaline, dry, smooth, thin-walled,

cylindrical, 23-40 x 7-10 um, (0-)3(-4)-septate, apex and base of intercalary

conidia truncate, apical conidia with obtuse or rounded apex.

SPECIMEN EXAMINED: CHINA, GUANGXI PROVINCE: Dayaoshan Nature Reserve, on

decaying twigs of an unidentified broadleaf tree, 7 Nov. 2012, X.X. Li (HSAUP H9021,

HMAS 243426).

Comments ~This is the first report of this species in China. The conidia of

the specimen examined are somewhat shorter and wider than the material

originally described by Ponnappa (18-50 x 6-8 um; Ponnappa 1975), but we

believe they are the same species. Parasympodiella laxa is similar to P. elongata

Crous et al. in conidial morphology, but P elongata differs by its much longer

conidiophores (<1300 um; Crous et al. 1995).

Parasympodiella eucalypti Cheew. & Crous, Persoonia 23: 70, 2009. FIG. 2

COLONIES on natural substrate effuse, brown to blackish brown, hairy.

Mycelium superficial and immersed, consisting of branched, septate, hyaline to

pale brown hyphae. ConipropHores distinct, solitary, unbranched, cylindrical,

<670 um, <13-septate; sterile part with a thickened wall, medium to dark

grey-brown, 6-8 um wide; fertile part thin-walled, pale grey-brown at basal

region, paler toward the apex. CONIDIOGENOUS CELLS terminal and intercalary,

integrated, indeterminate, extending sympodially, with one conidiogenous

locus per cell, smooth, pale grey-brown, becoming hyaline toward the apex.

Conipia thallic-arthric, hyaline to very pale brown, dry, smooth, thin-walled,

cylindrical, 28-54 x 6-9 um, (0-)1(-2)-septate, somewhat swollen in the

apical cells, apex and base of intercalary conidia truncate, with a punctiform

septal plug at each end, apical conidia with obtuse or round apex, occurring in

unbranched chains.

SPECIMEN EXAMINED: CHINA, GUANGXI PROVINCE: Dayaoshan Nature Reserve, on

decaying twigs of an unidentified broadleaf tree, 10 Nov. 2012, X.X. Li (HSAUP H9038,

HMAS 243427).

ComMENTS - Parasympodiella eucalypti is reported for the first time from

China. The conidia of the Chinese specimen are narrower than those of the

type material (width 8-11 um; Cheewangkoon et al. 2009), but other features

of our material closely match those of the original description. Parasympodiella

eucalypti resembles P. elongata in conidial shape, but P elongata has slightly

shorter and wider conidia (20-40 x 6-12 um; Crous et al. 1995).

124 ... Li &al.

Fic. 2. Parasympodiella eucalypti.

A. Colonies on natural substratum; B. Conidiophores and conidiogenous cells; C. Conidia.

Chalara unicolor S. Hughes & Nag Raj, New Zealand J. Bot. 12: 121, 1974. FIG. 3

COLONIES on natural substrate superficial, effuse, brown, hairy, with a thin

white bloom of conidial chains. Mycelium mostly immersed in the substrate.

CONIDIOPHORES solitary or loosely aggregated, erect, straight, subcylindrical,

uniformly brown, smooth, walls thickened somewhat below and thinner above,

85-130 um long, composed of a short basal stalk cell, 8.6-9.5 um wide with an

apical phialide. CONIDIOGENOUS CELLS phialides with a slightly inflated venter

about 11-14 um wide merging into a long, cylindrical collarette 8-9 um wide.

New Parasympodiella and Chalara records for China... 125

Conrpi cylindrical, barely rounded at the apex and flattened at the marginally

frilled base, hyaline, 3-septate, smooth, 24-34 x 7-8 um, sometimes observed

in readily seceding chains.

SPECIMEN EXAMINED: CHINA, GUANGXI PROVINCE: Guposhan National Forest Park,

on decaying twigs of an unidentified broadleaf tree, 9 Nov. 2012, X.X. Li (HSAUP

H9026, HMAS 243428).

Comments - This species has not been previously reported in China. The

Chinese collection has shorter conidia than those of the type material ((20-)

34-40(-45) um; Nag Raj & Hughes 1974), but otherwise fits well with the

original description. Chalara unicolor is similar to C. magnispora Matsush. in

conidial shape, but C. magnispora conidia are larger and sometimes 4-septate

(25-45(-56) x 7.5-10.5 um; Matsushima 1993).

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Fia. 3. Chalara unicolor.

A. Colonies on natural substratum; B. Conidiophores and conidiogenous cells; C. Conidia.

126 ... Li & al.

Acknowledgments

The authors express gratitude to Dr. Rafael E Castafeda-Ruiz and Dr. Bryce

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, 31230001) and the Ministry of Science and Technology of the

People’s Republic of China (Nos. 2006FY120100).

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Myrtaceae, a cache of fungal biodiversity. Persoonia 23: 55-85.

http://dx.doi.org/10.3767/003158509X474752

Crous PW, Wingfield MJ, Kendrick WB. 1995. Foliicolous dematiaceous hyphomycetes from

Syzygium cordatum. Can. J. Bot. 73(2): 224-234. http://dx.doi.org/10.1139/b95-025

Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasa WFO, Philips AJI, Alves A, Burgess T,

Barber P, Groenewald JZ. 2006. Phylogenetic lineages in the Botryosphaeriaceae. Stud. Mycol.

55: 235-253. http://dx.doi.org/10.3114/sim.55.1.235

Hughes SJ, Nag Raj TR. 1973. New Zealand fungi 20. Fusichalara gen. nov. New Zealand J. Bot. 11:

661-671. http://dx.doi.org/10.1080/0028825X.1973.10430307

Kendrick WB. 1958. Sympodiella, a new hyphomycete genus. Trans. Br. Mycol. Soc. 41(4): 519-521.

http://dx.doi.org/10.1016/S0007-1536(58)80077-7

Matsushima T. 1993. Matsushima Mycological Memoirs No. 7. Published by the author, Kobe,

Japan.

Nag Raj TR, Hughes SJ. 1974. New Zealand fungi 21. Chalara (Corda) Rabenhorst. New Zealand

J. Bot. 12: 115-129. http://dx.doi.org/10.1080/0028825X.1974. 10428628

Ponnappa KM. 1975. Parasympodiella gen. nov. Trans. Br. Mycol. Soc. 64(2): 344-345.

http://dx.doi.org/10.1016/S0007-1536(75)80126-4

Rabenhorst L. 1844. Deutschlands Kryptogamen-Flora 1. 614 p.

Riess H. 1853. Beitrage zur Pilzkunde. Bot. Zeitung 11: 129-140.

Subramanian CV. 1958. Hyphomycetes-V. J. Indian Bot. Soc. 37: 47-64.

Sutton BC, Pirozynski KA. 1965. Notes on microfungi. II. Trans. Br. Mycol. Soc. 48(3): 349-366.

http://dx.doi.org/10.1016/S0007-1536(63)80050-9

Went FA. 1893. Die ananaziekte van het suikerriet. Meded. Proefstat. Suikerriet W.-Java 5. 8 p.

Zhuang WY. 2001. Higher fungi of tropical China. Mycotaxon Ltd., Ithaca, New York. 485 p.

Zhuang WY. 2005. Fungi of Northwestern China. Mycotaxon Ltd., Ithaca, New York. 430 p.

MYCOTAXON

http://dx.doi.org/10.5248/126.127

Volume 126, pp. 127-132 October-December 2013

Tuber bomiense, a new truffle species from Tibet, China

Kat-ME!I Su', WEI-PING XIONG?, YUN WANG3,

SHU-HonNG LI’, RONG XIE? & DANZENG BAIMA?

" Biotechnology and Germplasm Resources Institute, Yunnan Academy of Agricultural Sciences,

Kunming 650223, Yunnan, China

? Institute of Vegetable Sciences, Tibet Academy of Agricultural and Animal Husbandry Sciences,

Lhasa, 850002, Tibet, China

° New Zealand Institute for Plant and Food Research Limited, Canterbury Agriculture & Science

Centre, Gerald St., Lincoln 7608, Christchurch, New Zealand

* CORRESPONDENCE To: pingwx_102@163.com

ABSTRACT — Tuber bomiense sp. nov. is described based on collections from the Bomi

region of Tibet, China. It differs from other Tuber species by its rust brown ascomata with a

verrucose and glabrous surface and ascospores with a regular reticulum. Molecular analysis

supports the erection of this new species.

Key worps — Pezizales, taxonomy, phylogeny

Introduction

Tibet is a unique region characterized by its high elevation and a wide

range of environments. In particular, the Linzhi region, which is located in the

Yaluzangbu River valley in southeastern Tibet, has a warm moist climate and a

vegetation known as “Tibetan Jiangnan” (‘jiangnar refers to warmth and green

scenery). Virgin biodiverse forests of pine and oaks cover the valley floors and

slopes.

Almost no research has been conducted on the hypogeous fungi of Tibet

although Tuber liui A.S. Xu, T: xizangense A.S. Xu, and T. oligospermum (Tul.

& C. Tul.) Trappe. have been recorded from the Linzhi region (Xu 1999; Chen

2007). Since 2010 the authors have systematically searched for truffles in the

Linzhi region and have made a few collections of both whitish and black

species. Here we describe and name one of these new species Tuber bomiense.

Materials & methods

The macroscopic and microscopic characters of the new species were described

based on fresh specimens following the methods of Yang & Zhang (2003). Sections

128 ... Su & al.

were made with a razor blade, mounted in water and examined under a Nikon E400

microscope. For scanning electron microscopy (SEM), spores were scraped from the

dried gleba onto doubled-sided tape, mounted directly onto an SEM stub, coated with

gold-palladium, and examined and photographed with a JEOL, JMS-5600LV SEM. The

holotype (SKM101) is deposited in the Herbarium of Yunnan Academy of Agricultural

Sciences, Kunming, China (YAAS).

DNA was extracted from ascomata using CTAB according to Doyle (1987) modified

by adding 200 uL 5M potassium acetate after the 4xCTAB treatment. The primers ITS1F

(Gardes & Bruns 1993) and ITS4 (White et al. 1990) were used to amplify the ITS-

rDNA region of the DNA extract, and the PCR reaction solution and cycling parameters

followed Chen & Liu (2007). Amplification products were electrophoresed on a 1%

agarose gel, and purified with Sangon’s purification kit. Sequencing was performed with

a BigDye Terminator v3.1 Cycle Sequencing Kit on an ABI 3730XL automatic sequencer.

TABLE 1. Origin of the fungal sequences included in the molecular analyses.

TAXON VOUCHER/CODE ORIGIN GENBANK*

Choiromyces alveolatus MES97 California, USA HM485332

Tuber bomiense SKM101 Bomi, Tibet, China KC517480

SKM106 Bomi, Tibet, China KC517481

T. borchii K(M)23814 unknown EU784423

T, foetidum B-2489 Szigetujfalu, Hungary AJ557544

T. indicum Clone C4 Unknown AF106883

Clone D3 Unknown AF106884

Clone C98-9 Unknown AF106881

T. macrosporum Clone Macrol Central Umbria, Italy AF106885

T. maculatum MTM2012 Nida’s Valley, Poland JX559773

T. magnatum Clone Ma2 Central Umbria, Italy AF106888

T. melanosporum Clone A51 Northern Piedmont, Italy AF106876

Clone A71 Spain AF106877

Clone A62 France AF106879

T. mesentericum Clone Mese4 Central Umbria, Italy AF106887

T. nitidum AH39101 Morocco JX402092

AH11906 Spain JX402090

T. rapaeodorum K(M)7705 Unknown EU784430

T. rufum GK4422 Greece JX402094

AH39102 Morocco JX402093

T. scruposum CMI-UNIBO 2207 Dilijan, Armenia DQ011847

CMI-UNIBO 2201 Dilijan, Armenia DQ011845

T. whetstonense Strain LO8B California, USA JF419244

J1T25783 Oregon, USA HM485392

SOC 756 Unknown AY830855

*Sequences produced in this study in bold.

Tuber bomiense sp. nov. (Tibet) ... 129

— 10ym amet

380kV¥ AMRAY #8082

PLATE 1. Tuber bomiense (YAAS SKM101). 1. Ascomata; 2-3. Peridial structure of one ascoma;

4. Asci (light microscope); 5. Ascospore (SEM); 6. Ascospores (light microscope).

The two T. bomiense ITS-rDNA sequences extracted in this study were compared

with 24 Tuber ITS-rDNA sequences downloaded from NCBI (TABLE 1); Choiromyces

alveolatus was selected as the outgroup. Software and sequence alignment and

phylogenetic protocols followed Chen & Liu (2007).

Taxonomy

Tuber bomiense K.M. Su & WP. Xiong, sp. nov. PLaTE 1

MycoBank MB 803258

Differs from other Tuber species by its rust brown ascomata with verrucose and glabrous

surface and subglobose to broadly ellipsoid ascospores with regular reticulum.

Type: China: Linzhi region, Bomi County, Yigong Town, 30°14’N 94°54’E, alt. 2390

m, in forest of Pinus densata Mast., 16.10.2011, Kaimei Su (Holotype, YAAS SKM101;

GenBank KC517480).

130 ... Su & al.

Erymo_oey: from the Latin bomiense referring to the location of the type collection.

AscoMarta subglobose or irregular, firm, rust-brown to brown, up to 2.0 cm

diam, surface verrucose, glabrous, cracked into small polygonal segments.

Ovok slightly aromatic when mature. PERIDIUM 230-520 um thick, two layers,

the outer layer 112.5-217.2 um thick, pseudoparenchymatous, composed

of large polygonal to subglobose cells 3-10 x 5-12 um diam, brownish to

brown, thin or thick walled 1-3 um thick, the inner layer 94.8-191.9 um thick,

composed of hyaline to pale yellow interwoven hyphae. GLEBA solid, whitish

when young, becoming rust brown at maturity, marbled with distinct, white,

meandering veins merging at many points within the peridium. Asc1 63-78 x

45-63 um, subglobose, rhombic, ellipsoid or irregular, sessile, thin walled 1-2

um thick, 1-4(-5) spored, randomly dispersed in glebal tissue. AscosPORES

subglobose to broad ellipsoid, in 1-spored asci 42.0 x 35.6 um, 2-spored

asci 35.2 x 30.4 um, 3-spored asci 34.3 x 31.0 um, 4-spored asci 28.7 x 25.1

uum, 5-spored asci 26.8 x 23.2 um; Q = (1.06-)1.07-1.29(-1.32), Q. = 1.18 +

0.08 (72/8/4); spore walls 25 um thick, brown at maturity, ornamented with

a regular alveolate-reticulum, up to 6 um deep, formed by mostly hexagonal

meshes 8-15 x (5-)7-13 um, 4-5 along the spore length and (2-)3-4 across

the spore width.

ECOLOGY & DISTRIBUTION: Hypogeous in calcareous soils with pH 7.4 under

Pinus densata at an elevation of c. 2400 m, fruiting in late October. Known only

from Cegang Village, Yigong, Bomi County, Linzhi region, Tibet, China.

ADDITIONAL SPECIMEN EXAMINED: CHINA: Linzhi region, Bomi County, Yigong

Town, 30°15’N 94°54’E, alt. 2400 m, in Pinus densata forest, 16.10.2011, Kaimei Su

(YAAS SKM106; GenBank KC517481).

Phylogenetic analysis

A total of 25 ITS Tuber sequences were used in the phylogenetic analysis

(TABLE 1) with an ITS Choiromyces alveolatus sequence as the outgroup. The

MP tree (PLATE 2) includes 776 steps (CI = 0.772, RI = 0.912).

Phylogenetic analysis of the ITS sequences revealed two well-supported

clades. Tuber borchii, T: maculatum, and T. foetidum are included in clade 1 with

99% bootstrap support. Clade 2 is composed of T. whetstonense, T. scruposum,

and T: bomiense with 100% bootstrap support. In Clade 2, T: bomiense and

T. scruposum formed a subclade with 79% bootstrap support. The result of

phylogenetic analysis shows that T: bomiense differs from all other truffle

species but is related to T. scruposum (PLATE 2).

Discussion

Tuber bomiense differs morphologically from all other truffle species

by its rust brown ascomata with a hairless verrucose surface cracked into

Choiromyces alveolatus HM485332

100

Tuber bomiense sp. nov. (Tibet) ... 131

T. foetidum AJ557544

T. foetidum AJ557543

Clade |

T. borchii EU784423

T. maculatum JX559773

T. rapaeodorum EU784430

T. whetstonense JF 419244

T. whetstonense HM485392

T. whetstonense AY830855

T. scruposum DQ011847

T. scruposum DQ011845 Clade II

Subclade |

T. bominense KC517480

T. bominense KC517481

T. macrosporum AF106885

81 T. magnatum AF106888

T. mesentericum AF106887

100

= 100

100

T. nitidum JX402092

T. nitidum JX402090

T. rufum ¥X402094

T. rufum ¥X402093

98 T. indicum AF106883

T. indicum AF106884

100

T. indicum AF106881

_10

T. melanosporum AF106876

T. melanosporum AF 106877

T. melanosporum AF106879

PLATE 2. Equally weighted parsimony tree of Tuber species based on ITS sequence.

small polygonal segments, and its ascospores are more subglobose than

ellipsoid (PLATE 1, Fics. 1, 4). It is morphologically close to T. quercicola,

T: rapaeodorum, T. scruposum, and T. whetstonense (TABLE 2). Tuber quercicola

has a similarly cracked peridium but its spiny ascospores distinguish it (Frank

et al. 2006). Tuber rapaeodorum differs in having pubescent ascomata and

ellipsoid ascospores with denser meshes. In T. scruposum the ascoma surface

is ochraceous tawny to dull ochraceous orange and its ascospores are more

ellipsoid than subglobose with narrow meshes (Ceruti et al. 2003). Finally,

T. whetstonense differs from T: bomiense by a pubescent peridium and ellipsoid

ascospores with much denser meshes (Ceruti et al. 2003; Frank et al. 2006).

132 ... Su &al.

TABLE 2. Morphological differences between Tuber bominense and similar Tuber species

SPECIES: bominense quercicola rapaeodorum scruposum whetstonense

PERIDIUM Verrucose, Minutely Smooth Slightly Minutely

SURFACE cracked verrucose to to finely warty, finely scurfy,

scaly, cracked pubescent pubescent pubescent

GLEBA Rust brown Light tan to Dark Greyish Dark

COLOR dk reddish brown brown vinaceous greyish

SPORE Subglobose / Ellipsoid Ellipsoid Ellipsoid Subglobose /

SHAPE ellipsoid ellipsoid

SPORE Reticulate; Spiny; Reticulate; Reticulate; Reticulate;

COVERING meshes broad spines curved meshes dense meshes dense meshes

very dense

The phylogenetic analysis also shows that T: bomiense is distinct from these

similar species (PLATE 2). The discovery of T: bomiense in Tibet provides

further evidence of the close phylogeographic relationship between Chinese

truffles and European and American species.

Acknowledgements

We are grateful to Dr Ian R. Hall and Associate Professor Li Fan for reviewing the

paper and providing valuable comments. This study received financial support from the

Chinese modern agricultural technology funding system (CARS-24).

Literature cited

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Torino.

Chen J. 2007. Taxonomy and phylogeny of the genus Tuber in China (Pezizales, Ascomycetes,

Ascomycotina). PhD Thesis of Graduate School of Chinese Academy of Sciences, China (in

Chinese).

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

Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure from small quantities of fresh leaf

tissues. Phytochem. Bull. 19: 11-15.

Frank JL, Southworth D, Trappe JM. 2006. NATS truffle and truffle-like fungi 13: Tuber quercicola,

and T. whetstonense, new species from Oregon, and T. candidum redescribed. Mycotaxon 95:

229-240.

Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes —application

to the identification of mycorrhizae and rusts. Mol. Ecol. 2: 113-118.

http://dx.doi.org/10.1111/j.1365-294X.1993.tb00005.x

Wang Y, Liu PG. 2009. Achievements and challenges of research on Chinese truffles. Acta Botanica

Yunnanica, Suppl. XIV: 1-9.

White TJ, Bruns T, Lee S, 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, New York.

Xu AS. 1999. Notes on the genus of Tuber from Tibet. Mycosystema 18: 361-365.

Yang ZL, Zhang LE. 2003. Type studies on Clitocybe macrospora and Xerula furfuracea var. bispora.

Mycotaxon 88: 447-454.

MY COTAXON

http://dx.doi.org/10.5248/126.133

Volume 126, pp. 133-141 October-December 2013

Lepiota brunneoincarnata and L. subincarnata:

distribution and phylogeny

A. RAZAQ”™, E.C. VELLINGA’, S. ILYAS* & A.N. KHALID’

‘Department of Botany, University of the Punjab, Lahore. 54590, Pakistan

*Department of Plant and Microbial Biology, University of California Berkeley California USA

*CORRESPONDENCE TO: ectomycorrhiza@gmail.com

ABSTRACT — An updated phylogeny of the clade of toxic Lepiota species is presented, and

new insights in the distribution of L. brunneoincarnata and L. subincarnata are given. Lepiota

brunneoincarnata is widespread in Europe and temperate Asia, and L. subincarnata is now

known from Asia, Europe, and North America. Morphological and anatomical descriptions

are provided for these two species based on material from the western Himalayan forests in

Pakistan, where they are reported for the first time.

KeEYworps — amatoxins, lepiotaceous fungi, mushroom diversity, rDNA

Introduction

Among lepiotaceous fungi the genus Lepiota (Pers.) Gray (Agaricales,

Basidiomycota) has a worldwide distribution and is highly diversified (Bon

1993; Ahmad et al. 1997; Vellinga 2003; Kirk et al. 2008; Razaq et al. 2012;

Nawaz et al. 2013). The genus is characterized by having a scaly (rarely smooth)

pileus, free lamellae, partial veil in the form of annulus, a universal veil, smooth

white dextrinoid spores (in most species), and clamp connections (present in

all but one or two species) (Vellinga 2001, Kumar & Manimohan 2009). The

nature of the pileus covering elements and the spore shape are very important

characters for infrageneric classification (Candusso & Lanzoni 1990; Bon

1993; Vellinga 2001, 2003). The pileus covering of Lepiota is a hymeniderm,

a trichoderm, an epithelium made up of long or globose elements, or cutis-

like (Vellinga 2001, 2003). Bon (1993) and Vellinga (2001) recognized two

subsections in sect. Ovisporae (J.E. Lange) Kithner, a section morphologically

characterized by ellipsoid spores and a trichodermal pileus covering based on

structure and size of the pileus elements: subsect. Helveolinae Bon & Boiffard

with a trichoderm made up of only longer pileus elements and subsect. Felininae

Bon with long and basal short elements. Vellinga (2003) discovered that sect.

134 ... Razaq & al.

Ovisporae is not monophyletic: species of subsect. Felininae clustered with

sect. Lepiota, while species of subsect. Helveolinae clustered with species with

spurred spores in sect. Stenosporae (J.E. Lange) Kihner. The clade of subsect.

Helveolinae is represented by the European species L. brunneoincarnata,

L. subincarnata, and L. farinolens Bon & G. Riousset, plus the tropical species

L. elaiophylla Vellinga & Huijser. Both L. brunneoincarnata and L. subincarnata

are infamous for amanitin poisoning (Gérault & Girre 1975; Bresinsky & Besl

1985). Such poisonings from Lepiota species result in liver and kidney failure

and, if not treated in time, death in humans (Khelil et al. 2010; Delacour et al.

2009; Donnelly et al. 2000; Haines et al. 1985).

Collected during fieldwork in the moist temperate forests of western

Himalaya dominated by coniferous vegetation, L. brunneoincarnata and

L. subincarnata have not previously been documented for Pakistan and are here

described morphologically and molecularly. Lepiota subincarnata is known

from several parts of North America, Europe, and eastern Himalaya, and

L. brunneoincarnata occurs throughout Europe and in eastern China. ITS-

rDNA sequences of these two species are now available from different regions

of the world. The objective of this paper is to provide a precise description of

the Pakistani material, to evaluate the phylogenetic relationship among all

isolates of these species from Asia, Europe, and North America, and to place

the species in a phylogenetic context.

Materials & methods

The basidiomata were carefully dug up using a knife and photographed in the field.

Material was characterized morpho-anatomically and molecularly. For microscopic

observation, sections were stained with Congo Red and Melzer’s reagent. Dimensions

were determined for 25 basidiospores, 20 basidia, 20 cheilocystidia, and 20 elements

of pileus covering from each basidioma. The following abbreviations are used: avl

for average length, avw for average width, Q for the basidiospore length-width ratio,

and avQ for average of all Q. Drawings were made using a camera lucida attached to

a compound microscope. Dried specimens were deposited in the LAH Herbarium,

Department of Botany, University of the Punjab, Lahore.

The protocol of Extract-N-Amp (XNAP-2) (Sigma, St. Louis, MO, USA) was

followed. Dried material of basidioma (approx. 1 mg) was taken in small PCR tubes to

which 10 ul of extraction solution was added. These tubes were incubated at 65°C for

10 min then at 94°C for 10 min. After the addition of 10 ul dilution solution (XNAP-

2), the tubes remained at room temperature for one hour. ITS regions of rDNA were

amplified using the universal primer pair ITS1F and ITS4 (White et al. 1990, Gardes

and Bruns 1993). PCR was performed in 25 uL reaction volume following Gardes and

Bruns (1993). PCR product of the ITS1+5.8S+ITS2 region was directly sequenced in

both directions using the same amplification primers (Macrogen, Korea).

All available ITS sequences of species in subsect. Helveolinae plus selected species in

subsect. Felininae and sect. Lepiota were downloaded from GenBank. Coprinus comatus

Lepiota brunneoincarnata and L. subincarnata new to Pakistan ... 135

Fic. 1: Lepiota brunneoincarnata. A, B. Basidioma. C. Basidiospores. D. Basidia E. Cheilocystidia.

F, Pileus covering elements. Scale bars: A, B = 1 cm; C = 3.45 um; D = 6.25 um; E = 13 um;

F = 21.25 um.

was used as outgroup. The sequences were aligned using MAFFT version 6 (Katoh et

al. 2002, Katoh & Toh 2008) with default settings. The total 34-sequence dataset was

analyzed by a maximum likelihood (ML) method with RAxML 7.2.3 (Stamatakis et

al. 2008). All free-model parameters were estimated by RAxML with a general time

reversible (GTR) substitution matrix and a proportion of invariable sites estimated. One

hundred ML bootstraps were performed.

The newly produced nucleotide sequences were submitted to the European Molecular

Biology Laboratory (EMBL) database and are available in GenBank. Numbers are listed

with the collections and in Fie. 3.

Results

Lepiota brunneoincarnata Chodat & C. Martin, Bull. Trav. Soc. Bot.

Geneve 5: 222. 1889. Fic. 1

PILEUsS 2.6 cm diam., campanulate to plano-convex, with obtuse to slightly

umbonate central disc and deflexed margins; central disc with greyish-brown

to dark brown scales on white to cream or pinkish buff background, around

center concentric crowded and prominent squamules slightly paler or lighter

136 ... Razaq & al.

Fia. 2: Lepiota subincarnata. A, B. Basidioma. C. Basidiospores. D. Basidia. E. Cheilocystidia. F. Pileus

covering elements. Scale bars: A, B = 1 cm; C = 3.45 um; D = 6.25 um; E = 13 um; F = 21.25 um.

toward margin, made up of small pyramids; margin smooth, at maturity fragile

and broken. LAMELLAE free, moderately crowded, white to cream, with 2-3

tiers of lamellulae alternating with lamellae; edges wavy to denticulate. STIPE

4.2 x 0.3 cm, centrally attached, cylindrical, dark-brown to dull brown, light

brown to pinkish brown in apical half, smooth in upper part, scaly in lower

part; scales in bands forming incomplete girdles, non-persistent annulus; white

basal mycelial strands. Taste not recorded.

BASIDIOSPORES 5-8.5 x 3.5-5.5 um, avl x avw = 6.5 x 4.5 um, Q = 1.4-1L.5,

avQ = 1.45, ellipsoid to ovoid, slightly amygdaliform, smooth, moderately

thick-walled, colourless or pale yellow in 5% KOH, reddish brown in Melzer’s

reagent and strongly dextrinoid; wall colouring in Cresyl Blue not observed.

Basip1A 21-31.5 x 8-9.5 um, 4-spored, subclavate to clavate, hyaline to pale

yellow in 5% KOH, thin-walled, with oil-like contents. PLEUROCYSTIDIA

absent. CHEILOCYSTIDIA 18.5-31 x 8-10.5 um, usually clustered, hyaline,

thin-walled, narrowly clavate to clavate. PILEUS COVERING a trichoderm with

erect or ascending, light brown to hyaline, clamped terminal elements, 90-253

x 7.5-13.5 um, cylindrical to slightly narrowly fusoid, without any shorter

elements at the base. CLAMP CONNECTIONS common in all tissues.

Lepiota brunneoincarnata and L. subincarnata new to Pakistan ... 137

MATERIAL EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA, Khanspur, 2250 m a.s.L.,

solitary, on moist ground under Abies pindrow, 23 August 2010, A. Razaq K-40 (LAH

230840; GenBank HE863668).

Lepiota subincarnata J.E. Lange, Fl. agar. dan. 5 (Taxonomic Conspectus):

V, 1940. FiG: 2

Piteus 4.0-4.2 cm diam., hemispherical to plano-convex, with obtuse to

slightly umbonate central disc, white, with thick context, with reddish brown

to pinkish brown scales on white background on central disc, fading towards

margins; margins smooth, sometimes splitting open at maturity. LAMELLAE

free, crowded, white to cream, ventricose, with 3-4 tiers of lamellulae, with

wavy to denticulate edge. Stipe 5.5 x 0.8 cm, centrally attached, cylindrical,

vinaceous red to pinkish brown, hollow; surface smooth to scaly with fibrillose

type scales below annular zone. ODouR none. TASTE not recorded.

BASIDIOSPORES 6-7.5 Xx 3.5-5 um, avl x avw = 6.6 x 4.3 um, Q= 1.5-1.7;

avQ = 1.55, oblong ellipsoid to ellipsoid to ovoid, colourless or pale yellow

in 5% KOH, reddish brown in Melzer’s reagent and strongly dextrinoid,

smooth, thin-walled. BAsip1a 19.5-28.5 x 7-8.5 um, 4-spored, occasionally

2-spored, subclavate to cylindrical, hyaline, thin-walled with oil contents.

PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 18-35.5 x 7-11.5 um, hyaline,

thin walled, subclavate to clavate. PILEUS COVERING a trichoderm with erect

or ascending, hyaline to light brown, clamped terminal elements, 134-318 x

10.5-12.0 um, cylindrical to slightly widened, with thickened apices, without

any shorter elements at the base. CLAMP CONNECTIONS present in all tissues.

MATERIAL EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA, Khanspur, 2250 m a.s.L.,

solitary, on moist ground under Abies pindrow, 23 August 2010, A. Razaq KP-09 (LAH

230809; GenBank HE863669).

Discussion

Bon (1993), Vellinga (2001), and the authors before them attached much

importance to basidiospore shape for infrageneric classification in Lepiota.

Sect. Lepiota, sect. Ovisporae, and species in sect. Stenosporae share a

trichodermal pileus covering but differ in spore shape: fusiform-amygdaliform

in sect. Lepiota, ellipsoid to oblong in sect. Ovisporae, and with a spur in sect.

Stenosporae. Vellinga (2003) discovered that sect. Ovisporae is polyphyletic

and that in general the pileus covering is a more significant character for

infrageneric delimitation than the spore shape. Bon (1993) and Vellinga (2003)

dealt mostly with European species.

In our present analysis, we included all available ITS sequences from

different parts of the world and recovered the same results (Fic. 3). Species with

a trichodermal pileus covering with long and short elements cluster together in

Clade 2 (Fic. 3). This clade corresponds to sect. Lepiota and subsect. Felininae.

138 ... Razaq & al.

AY176370 L. forquignonii NL

AY176366 L. echinella Belgium

Clade 2 97 AY176405 L. xanthophylla NL

98 U85329 “L. subincarnata” S. Korea

100 AF391005 L. magnispora Germany

90 AY176470 L. erminea NL

76 U85330 L. felina U.S.A.

AY176490 L. subgracilis NL

AY176400 Lepiota sp. MO U.S.A.

HE614898 L. himalayensis PAK

AY176368 Lepiota farinolens France

99 HE974764 L. vellingana PAK

100 HE974766 L. vellingana PAK

HE974765 L. vellingana PAK

AY176478 Lepiota sp. MI U.S.A.

Clade 1 FJ998401 “L. helveola” Italy

EU416302 L. brunneoincarnata China

AF482875 L. brunneoincarnata NL

FJ998395 L. brunneoincarnata Ital

100_1FJ481017 L. brunneoincarnata China

AY176355 L. brunneoincarnata NL

FJ481017 L. brunneoincarnata China

JN224825 Lepiota sp. Thailand

JN224828 Lepiota sp. Thailand

JN224826 Lepiota sp. Thailand

AY176402 Lepiota sp. HI U.S.A.

D AY 194534 “Galeropsis desertorum”

AY176491 L. subincarnata NL

FM999639 Uncultured OH U.S.A.

KC556779 L. subincarnata CA U.S.A.

AF391024 L. elaiophylla NL

HE819398 Coprinus comatus Pakistan

100

91} 100

100

0.05

Fic. 3. Phylogenetic relationships of Pakistani collections of L. brunneoincarnata and L. subincarnata

with other Lepiota species based on Maximum Likelihood analysis of nrITS sequences. Bootstrap

values >70, based on 100 replicates shown at each node. ‘The analysis involved 33 sequences with

Coprinus comatus as outgroup. Names in quotation marks are discussed in the text. The two new

Pakistani collections are outlined.

Lepiota brunneoincarnata and L. subincarnata new to Pakistan ... 139

Species without these short pileus covering elements cluster in Clade 1, which

corresponds to subsect. Helveolinae. The taxa in this clade are infamous for

amatoxins (Vellinga 2003). Two recently described species from Pakistan —

L. himalayensis Razaq & Khalid and L. vellingana Nawaz & Khalid — also

cluster in Clade 1.

Both L. brunneoincarnata and L. subincarnata form well supported clades

(both with 100% bootstrap support) and do not show any geographic structure,

despite the fact that the sequences originate from very widely separated areas

of Asia, Europe, and North America. It might be that ITS sequences alone are

not sufficient, and data from other gene regions are needed to differentiate

populations within the species.

Lepiota brunneoincarnata and L. subincarnata differ mainly in basidiocarp

colours: L. brunneoincarnata has reddish brown or dark brown scales while

L. subincarnata has nearly pinkish or vinaceous brown fine scales.

Lepiota brunneoincarnata from Pakistan, which clusters with isolates from

Europe and China, is morphologically similar to European collections as

described by Vellinga (2001). One GenBank sequence (FJ998401, labelled as

L. helveola Bres.) falls within the L. brunneoincarnata clade, but the sequence is

very short and it is likely that the specimen was misidentified.

The occurrence of L. brunneoincarnata in Pakistan suggests that it could

be expected in other Asian countries; it is already known from China, based

on sequence data, and from Israel (Kosakyan et al. 2008) and Turkey (Afyon

& Yagiz 2004), based on morphological descriptions. This species is not yet

known from North America, although a morphologically similar species was

found in Ann Arbor (Michigan; GenBank AY176478).

The Pakistani collection of L. subincarnata is in close agreement with

Vellingas (2001) morphological description of Dutch collections. However,

our collection is distinguished by longer pileal elements with thickened apices

(11-20 um), a difference that may be due to regional climate and geographical

isolation.

In Asia, Lepiota subincarnata was previously known from Israel (Kosakyan

et al. 2008) and from southwestern China (Yang 1990); the description of the

Chinese material is similar to that of our Pakistani collection.

Kumar & Manimohan (2009) list L. subincarnata from Kerala, India, but

their short description suggests a different species based on the dark brown

colours and the pileus covering structure with short basal elements that are up

to 400 um long. A collection from South Korea identified as L. subincarnata

(U85329; Johnson 1999) does not belong to L. subincarnata but falls among

the Clade 2 species having a pileus trichoderm composed of longer and shorter

cells (Fic. 1).

140 ... Razaq & al.

A sequence in GenBank (AY 194534, labelled as Galeropsis desertorum Velen.

& Dvorak), falls within the L. subincarnata clade. According to Hallen et al.

(2003), this sequence is from a specimen collected in 1930 in Moravia (Czech

Republic). It is possible that contamination or a laboratory mix-up might have

caused this unexpected placement.

Acknowledgements

Authors are thankful to Dr. Junfeng Liang (Research Institute of Tropical Forestry,

China) and Dr. Zai-Wei Ge (Chinese Academy of Science, China) for acting as expert

reviewers to improve the manuscript. We are really thankful to Prof. Tom Bruns

(Department of Plant and Microbial Biology, University of California, Berkeley) for

providing space to complete this article. This work was financially supported by Higher

Education Commission (HEC) of Pakistan under the “Indigenous Ph.D. Fellowship

Scheme 5000 Phase IV” and International Research Initiative Support program (IRISP).

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MY COTAXON

http://dx.doi.org/10.5248/126.143

Volume 126, pp. 143-156 October-December 2013

New combinations in Trichoderma (Hypocreaceae, Hypocreales)

WALTER M. JAKLITSCH* & HERMANN VOGLMAYR

Department of Systematic and Evolutionary Botany, Faculty Centre of Biodiversity,

University of Vienna, Rennweg 14, A-1030 Vienna, Austria

* CORRESPONDENCE TO: walter jaklitsch@univie.ac.at

ABSTRACT — Unitary nomenclature demands the use of a single name for pleomorphic fungi

determined according to priority. For this reason combinations in Trichoderma are here

provided for 46 species for which such a combination is lacking. Although many more such

species are known, only those are included here that are dealt with in more recent papers

and where some DNA data are available in GenBank, even if erroneous; for other species it is

strongly recommended to consult databases like Index Fungorum or MycoBank. Information

on types is provided for most species, and representative cultures, GenBank accessions for

tefl and rpb2, and important references are given for all species.

KEY worps — anamorph, Ascomycota, ICN, Kew rule, pyrenomycetes, teleomorph

Introduction

Hypocrea Fr. 1825 is the type and name-giving genus of the Hypocreaceae,

Hypocreales. However, its anamorphic counterpart, Trichoderma Pers. 1794 :

Fr. is older. The Melbourne Code of Nomenclature (ICN) dictates a unitary

nomenclature, i.e., the use of a single name for a pleomorphic fungus, no matter

whether ana- or teleomorphic, based either on priority or on commission-

sanctioned decisions. The species of Hypocrea/Trichoderma are monophyletic

and this is to be expressed in a single generic name. Trichoderma has priority

over Hypocrea and is commonly used in commercial applications, e.g., cellulase

production by T! reesei. A poll of individuals interested in the subject was taken

by the International Subcommission on Trichoderma and Hypocrea (ISTH),

which indicated a clear preference for adoption of Trichoderma over Hypocrea.

Thus Rossman et al. (2013) proposed this generic name for acceptance by

the Nomenclature Committee for Fungi (NCF) and the General Committee

(GC) of the International Association for Plant Taxonomy (IAPT). However,

a list of accepted names in Trichoderma has not yet been produced. There

are some difficulties associated with this task. One example is the wide use

144 ... Jaklitsch & Voglmayr

in commercial applications of Trichoderma epithets that are younger than the

associated Hypocrea epithets, e.g., T. reesei / H. jecorina and T. citrinoviride

/ H. schweinitzii. Gams et al. (2012) suggested that authors establishing lists

of protected names and the committees should apply the ‘Kew rule’ and

accordingly “not recombine older epithets from a list-demoted genus into the

list-accepted genus, when another one from pre-2013 is already available in

that genus.”

Index Fungorum lists 509 epithets in Hypocrea, including 55 names of

varieties and forms (which have no priority at the species level). Many of

these names do not represent members of this genus or are synonyms of other

species of the genus, and the vast majority has never been re-assessed since

their original description. Because of this large number of species names, it

is strongly advisable for those who intend to describe new species to consult

the Index of Fungi or databases like Index Fungorum or MycoBank, in order

to avoid the creation of homonyms. At present, 219 Trichoderma epithets

(including 9 varieties and forms) are listed in Index Fungorum. Many species

have been re-assessed by several recent workers and others have been newly

described in recent years. Jaklitsch et al. (2009, 2011), for example, established

names in both Hypocrea and Trichoderma for species that are known to form

an anamorph, but provided a name only in Hypocrea where no anamorph

is known. In another example, Chaverri & Samuels (2003) did not establish

names in Trichoderma for species earlier described in Hypocrea or for new

species, where cultures did not survive storage. Such species as described

in these works and several other recent papers (see below), are usually well

characterized and also documented by molecular data. Therefore we provide

here combinations in Trichoderma for these species as an intermediate step

towards a list of accepted names in the genus Trichoderma.

The large number of species on one hand, the large number of isolates of many

species on the other, and also the different portions of genes used as molecular-

phylogenetic markers has made it difficult to select representative isolates and

respective accessions for many species. For this reason we provide information

on types, ex-type cultures, representative cultures and representative GenBank

accessions for the most important markers, rpb2 and tefl. We indicate also

section or clade affinity for the species; when a formally established section is

also supported by molecular phylogeny, we give preference to that designation.

Materials & abbreviations

All publications on the taxonomy and phylogeny of Hypocrea and/or Trichoderma

that include molecular data were consulted and for species lacking an epithet in

Trichoderma, information on types, ex-type or representative cultures was extracted.

GenBank accessions for representative cultures were retrieved from the NCBI

homepage (http://www.ncbi.nlm.nih.gov/). The following abbreviations are used in the

New Trichoderma combinations ... 145

text: n.a. = not available, n.d. = not determined; phylog. = phylogenetic affinity; repres.

= representative; sect. = section.

Results - data arrangement

New combinations in Trichoderma are proposed for Hypocrea names in

species where such a combination is lacking. The 46 species treated here are

(1) 41 for which DNA data are available and (2) 5 that are included because of

confusing data. Additional data are given as: type information; repres. cultures;

phylog. clade or section (plus short information on (atypical) anamorphs

deviating from Trichoderma that form green conidia); repres. GenBank

sequence accessions; important (recent) references.

Trichoderma albocorneum (Yoshim. Doi) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807413

= Hypocrea albocornea Yoshim. Doi, Bull. natn. Sci. Mus., Tokyo 15: 712 (1972).

Ho.otype: TNS-F-190171, ex-type culture: n.a.; possibly repres.: G.J.S. 97-28 (tefl:

AY937440).

PHYLOG.: unknown; species with green ascospores. REFERENCES: Doi (1972), Chaverri

& Samuels (2003).

Trichoderma albofulvum (Berk. & Broome) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807414

= Hypocrea albofulva Berk. & Broome, J. Linn. Soc., Bot. 14: 113 (1873).

HOo.uoryPe: n.d.; ex-type culture: n.a. The species is similar to T. ochroleucum.

PHYLOG.: sect. Trichoderma. Note: The species was described from Sri Lanka. The

strains G.J.S. 01-234 (tefl accession: DQ846668) and/or G.J.S. 01-265 (rpb2 accession:

DQ835524) from material collected in Thailand may represent this species (see Jaklitsch

2011).

Trichoderma alcalifuscescens (Overton) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807415

= Hypocrea alcalifuscescens Overton, in Overton, Stewart

& Geiser, Stud. Mycol. 56: 62 (2006).

Ho.ortyPe: BPI 843638; isotype TAA(M) 181548; ex-type culture: CBS 122303 = TFC

2000-36.

Phylog.: unnamed clade; ‘basal’; anamorph verticillium-like, conidia hyaline. REPREs.

SEQUENCES: tefl: FJ860610; rpb2: DQ834462. REFERENCE: Overton et al. (2006a),

Jaklitsch (2011).

Trichoderma americanum (Canham) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807416

= Hypocrea citrina var. americana Canham, Mycologia 61: 320 (1969).

= Hypocrea americana (Canham) Overton, in Overton, Stewart,

Geiser & Jaklitsch, Stud. Mycol. 56: 21 (2006).

HouortyPe: NY ex CUP 38045; ex-type culture: ATCC 18574; other repres. strains: CBS

123072 = G,J.S. 92-93, G.J.S. 96-191.

146 ... Jaklitsch & Voglmayr

PHYLOG.: sect. Hypocreanum; anamorph acremonium-like, conidia hyaline. REPREs.

SEQUENCES: fef1: DQ835435 + DQ005523 + DQ835489; rpb2: DQ835455. REFERENCE:

Overton et al. (2006a).

Trichoderma andinense (Samuels & Petrini) Samuels, Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807417

= Hypocrea andinensis Samuels & Petrini, in Samuels, Petrini, Kuhls,

Lieckfeldt & Kubicek, Stud. Mycol. 41: 13 (1998).

Ho .ortyPe: BPI 1109854; ex-type culture: CBS 345.97 = G.J.S. 90-140 = ATCC 208857.

PHYLOG:.: sect. Longibrachiatum. REPRES. SEQUENCES: tefl: AY956321; rpb2: JN175531.

REFERENCES: Samuels et al. (1998, 2012).

Trichoderma atrogelatinosum (Dingley) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807418

= Hypocrea atrogelatinosa Dingley, Trans. Roy. Soc. New Zealand 83: 645 (1956).

Ho.otype: PDD 10471; ex-type culture: n.a..

REFERENCE: Chaverri & Samuels (2003). NoTE: Status unclear. Several accessions are

deposited in GenBank under the name Hypocrea atrogelatinosa; rpb2 sequences (strains

G,J.S. 88-28, 89-136, and 95-159) as well as tefl exon sequences belong to T! ceraceum

(H. ceracea); however, tefl intron sequences vary considerably among these isolates.

Trichoderma avellaneum (Rogerson & S.T. Carey) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807419

= Hypocrea avellanea Rogerson & S.T. Carey, in Carey

& Rogerson, Brittonia 28: 381 (1976).

Ho.otype: NY (MEBB 2471); isotype in K; ex-type culture: n.a.; repres. strain CBS

121667 = C.T.R. 77-155.

PHyLoG.: unknown, ‘basal’; anamorph verticillium-like, conidia hyaline. REpREs.

SEQUENCES: tef1: AY225857; rpb2: AF545562. REFERENCE: Carey & Rogerson (1976).

Trichoderma britdaniae (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807420

= Hypocrea britdaniae Jaklitsch & Voglmayr, Mycologia 104: 1216 (2012).

HOo.otyPe: K(M) 89878; ex-type culture: n.a. (ascospores not germinating).

PHYLOG.: sister to sect. Longibrachiatum; anamorph unknown. REPRES. SEQUENCES

(from stromata): tefl: JQ685865; rpb2: JQ685881. REFERENCE: Jaklitsch & Voglmayr

(2012).

Trichoderma caerulescens (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807422

= Hypocrea caerulescens Jaklitsch & Voglmayr, in Jaklitsch,

Stadler & Voglmayr, Mycologia 104: 928 (2012).

Ho otype: WU 31600; ex-type culture: CBS 130011 = S195.

PHYLOG.: sect. Trichoderma. REPRES. SEQUENCES: tefl: JN715621; rpb2: JN715604.

REFERENCE: Jaklitsch et al. (2012).

New Trichoderma combinations ... 147

Trichoderma citrinum (Pers. : Fr.) Jaklitsch, W. Gams & Voglmayr, comb. nov.

MycoBank MB807423

= Sphaeria citrina Pers., Observ. mycol. (Lipsiae) 1: 68 (1796) : Fr.

= Hypocrea citrina (Pers. : Fr.) Fr., Summa veg. Scand.,

Section Post. (Stockholm): 383 (1849)

= Sphaeria lactea Fr., Kongl. Vetensk. Acad. HandL., n.s. 37: 141 (1816) : Fr.

= Hypocrea lactea (Fr. : Fr.) Fr., Summa veg. Scand., Section Post.: 383 (1849).

= Trichoderma lacteum (Fr. : Fr.) Bissett, Canad. J. Bot. 69: 2367 (1992 [“1991”]).

NEoTYPE of Trichoderma citrinum: W. Gams 4031 (CBS); ex-type culture: CBS 894.85.

Holotype of Trichoderma lacteum: DAOM 167644; ex-type culture CBS 853.70. EPITYPE

of Trichoderma lacteum, here designated: W. Gams 4031 (CBS); ex-epitype culture: CBS

894.85.

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. REPREs.

SEQUENCES: tefl: FJ860631, DQ835441; rpb2: FJ179603, AF545561. REFERENCES:

Overton et al. (2006a), Jaklitsch (2011).

Note: Hypocrea citrina is a well-known name for a well-known species, particularly in

Europe, and has priority over H. lactea. Bissett (1992) erected the name Trichoderma

lacteum for the anamorph of H. lactea. The tef1 intron 5 of its ex-type strain CBS 853.70

clearly demonstrates conspecificity of T: lacteum with T. citrinum. The revised citation of

T: lacteum, above, follows the working practice advocated by Hawksworth et al. (2013).

Application of the ‘Kew rule’ would favour T: lacteum as the appropriate Trichoderma

name for a synonymised H. citrina/H. lactea. However, this would be undesirable,

because the inconsistent application of the name H. lactea to a number of different

taxa could be misleading; for example, “Hypocrea lactea“ sensu Doi (1972) was recently

described as T. pseudolacteum by Kim et al. (2013).

Trichoderma corneum (Pat.) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807424

= Hypocrea cornea Pat., J. Bot. (Morot): 64 (1890).

Ho .oryPe: K; ex-type culture: n.a.

The cultures G.J.S. 97-82 (tefl: AY937435), GJ.S. 97-75 (tefl: AY937431) and G,J.S.

97-90 (tefl: AY937426) are isolates from specimens collected in Thailand. As the

GenBank tefl accessions for these isolates vary considerably, fresh material from the

original collecting region Tonkin, Vietnam, is necessary to judge which of them match

the species. PHYLOG.: unclear; species with green ascospores. REFERENCE: Chaverri &

Samuels (2003).

Trichoderma costaricense (P. Chaverri & Samuels) P. Chaverri, Jaklitsch &

Voglmayr, comb. nov.

MycoBank MB807425

= Hypocrea costaricensis P. Chaverri & Samuels, Stud. Mycol. 48: 58 (2003).

Ho.otype: INB 0003527695; ex-type culture: P.C. 21 (lost after determination of DNA

data).

Puy oa.: Chlorospora clade, species with green ascospores. REPRES. SEQUENCES: tefl]:

AY737741 + AY391980; rpb2: AY391921. REFERENCE: Chaverri & Samuels (2003).

148 ... Jaklitsch & Voglmayr

Trichoderma danicum (Jaklitsch) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807426

= Hypocrea danica Jaklitsch, Stud. Mycol. 63: 41 (2009).

Ho.otyPe: WU 29046; ex-type culture: CBS 121273 = C.P.K. 2448 = Hypo 402.

PHyLoG.: Spinulosa clade, species with green ascospores; anamorph not formed.

REPRES. SEQUENCES: tef1: FJ860634; rpb2: FJ860534. REFERENCE: Jaklitsch (2009).

Trichoderma decipiens (Jaklitsch, K. Poldmaa & Samuels) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807427

= Hypocrea decipiens Jaklitsch, K. Poldmaa & Samuels, Mycologia 100: 981 (2008).

Ho.ortyPe: BPI 747356; ex-type culture: CBS 121307 = G.J.S. 97-207; repres. G.J.S. 91-

101.

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. REpREs.

SEQUENCES: tefl: FJ860635, EF550995; rpb2: DQ835520. REFERENCES: Jaklitsch et al.

(2008), Overton et al. (2006b; as Hypocrea farinosa). Some further accessions can be

retrieved under the name “Hypocrea farinosa’, because the name has not been updated,

pending consent by B. Overton.

Trichoderma eucorticioides (Overton) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807428

= Hypocrea eucorticioides Overton, in Overton, Stewart

& Geiser, Stud. Mycol. 56: 55 (2006).

Ho.oryPe: LPS 1719; ex-type culture: n.a.; repres. G.J.S. 99-61 (lost).

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. REpREs.

SEQUENCES: tefl: DQ835502 + DQ835474; rpb2: DQ835518. REFERENCE: Overton et

al. (2006b).

Trichoderma flaviconidium (P. Chaverri, Druzhin. & Samuels) Jaklitsch &

Voglmayr, comb. nov.

MycoBank MB807429

= Hypocrea flaviconidia P. Chaverri, Druzhin. & Samuels, in Druzhinina,

Chaverri, Fallah, Kubicek & Samuels, Stud. Mycol. 50: 404 (2004).

Ho.otype: INB 3862698; isotype BPI 746538; ex-type culture: G.J.S. 99-51 (dead);

repres. G.J.S. 99-49, CBS 116238 = G.J.S. 99-57.

PHYLOG:.: sect. Trichoderma, Hamatum subclade. REPRES. SEQUENCES: tef1: DQ020001,

AY665711; rpb2: EU883557. REFERENCE: Druzhinina et al. (2004).

Trichoderma flavipes (Peck) Seifert, Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807430

= Stilbum flavipes Peck , Ann. Rep. N.Y. St. Mus. nat. Hist. 31: 45 (1879 [“1877”]).

= Stilbella flavipes (Peck) Seifert, Stud. Mycol. 27: 68 (1985).

= Hypocrea cinereoflava Samuels & Seifert, in Seifert & Samuels, Mycologia 89: 515 (1997).

Ho torypE of Stilbum flavipes: NYS; holotype of Hypocrea cinereoflava: BPI 802847; ex-

teleotype culture: G.J.S. 92-102 = DAOM 222357.

New Trichoderma combinations ... 149

PHYLOG.: unnamed clade, ‘basal’; anamorph synnematous, stilbella-like in nature,

conidia green. REPRES. SEQUENCES: tefl: DQ834454; rpb2: DQ834461. REFERENCE:

Seifert & Samuels (1997); additional synonyms given in this publication.

Trichoderma foliicola (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807431

= Hypocrea foliicola Jaklitsch & Voglmayr, Mycologia 104: 1218 (2012).

Ho.otype: WU 31611; ex-type culture: CBS 130008 = Hypo 645.

PHyLoG.: Pachybasium Core Group, conidia hyaline. REPRES. SEQUENCES: tef1:

JQ685862; rpb2: JQ685876. REFERENCE: Jaklitsch & Voglmayr (2012).

Trichoderma hispanicum (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807432

= Hypocrea hispanica Jaklitsch & Voglmayr, in Jaklitsch,

Stadler & Voglmayr, Mycologia 104: 935 (2012).

Ho otyPe: WU 31606; ex-type culture: CBS 130540 = S453.

PHYLOG.: sect. Trichoderma. REPRES. SEQUENCES: tefl: JN715659; rpb2: JN715600.

REFERENCE: Jaklitsch et al. (2012).

Trichoderma hunua (Dingley) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807433

= Hypocrea hunua Dingley, Trans. & Proc. Roy. Soc. New Zealand 79(3-4): 327 (1952).

Ho.otyPe: PDD 10455; ex-type culture: n.a.; repres. CBS 238.63 (isolated: Dingley, No. 5).

PHyLoa.: Semiorbis clade. REPRES. SEQUENCES: tef1: AF401011; rpb2: n.a. REFERENCE:

Kullnig-Gradinger et al. (2002).

Trichoderma lacuwombatense (B.S. Lu, Druzhin. & Samuels) Jaklitsch & Voglmayr,

comb. nov.

MycoBANnkK MB807434

= Hypocrea lacuwombatensis B.S. Lu, Druzhin. & Samuels, in Lu, Druzhinina,

Fallah, Chaverri, Gradinger, Kubicek & Samuels, Mycologia 96: 338 (2004).

Ho.otyPe: PDD 77489; isotype BPI 746621; ex-type culture: CBS 122668 = G,J.S. 99-

198.

PuyLoc.: Pachybasium Core Group. REPRES. SEQUENCES: tefl: AY937452; rpb2: n.a.

REFERENCE: Lu et al. (2004).

Trichoderma megalocitrinum (Yoshim. Doi) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807435

= Hypocrea megalocitrina Yoshim. Doi, Bull. natn. Sci. Mus., Tokyo 15: 669 (1972).

Ho.otyPe: TNS-F-223220; isotype in NY; ex-type culture: n.a.; repres. BEO 00-09.

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. REPREs.

SEQUENCES: tefl: AY225855 (exon); rpb2: AF545563. REFERENCE: Overton et al. (2006a).

Trichoderma microcitrinum (Yoshim. Doi) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB MB807436

= Hypocrea microcitrina Yoshim. Doi, Bull. natn. Sci. Mus., Tokyo 15: 667 (1972).

150 ... Jaklitsch & Voglmayr

Ho.otype: TNS-F-223325; ex-type culture: n.a.; repres. BPI 744660; repres. cultures

G,J.S. 97-248, G.J.S. 91-61.

PHYLOG.: sect. Hypocreanum; anamorph acremonium- to verticillium-like, conidia

hyaline. REPRES. SEQUENCES: tefl: DQ835449 + DQ835479; rpb2: DQ835462.

REFERENCE: Overton et al. (2006a).

Trichoderma neorufum (Samuels, Dodd & Lieckf.) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807437

= Hypocrea neorufa Samuels, Dodd & Lieckf., in Dodd, Lieckfeldt,

Chaverri, Overton & Samuels, Mycol. Prog. 1: 421 (2002).

Ho otype: BPI 744493; ex-type culture: CBS 111144 = GJ.S. 96-135; repres. CBS

119498.

PHYLOG.: sect. Trichoderma. REPRES. SEQUENCES: tefl: FJ860653; rpb2: FJ860550.

REFERENCES: Dodd et al. (2002), Jaklitsch (2011).

Trichoderma novae-zelandiae (Samuels & Petrini) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807438

= Hypocrea novae-zelandiae Samuels & Petrini, in Samuels, Petrini,

Kuhls, Lieckfeldt & Kubicek, Stud. Mycol. 41: 25 (1998).

Ho.otyPe: PDD 46792; ex-type cultures: CBS 496.97 and CBS 639.92 = G.J.S. 81-265

= ATCC 208856.

PHYLOG:.: sect. Longibrachiatum. REPRES. SEQUENCES: tefl: AY865639, AY937448; rpb2:

JN133563. REFERENCES: Samuels et al. (1998, 2012), Druzhinina et al. (2012).

Trichoderma nybergianum (T. Ulvinen & H.L. Chamb.) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807439

= Hypocrea nybergiana T. Ulvinen & H.L. Chamb., in Chamberlain,

Rossman, Stewart, Ulvinen & Samuels, Karstenia 44: 21 (2004).

Ho.otyPe: OULU F 49597; isotype OULU F 49596; ex-type culture: n.a.; repres. CBS

122500 = C.P.K. 3159 = Hypo 572, CBS 122496 = C.P.K. 3163 = Hypo 577.

PHyLoG.: Pachybasium Core Group, conidia hyaline. REPRES. SEQUENCES: tefl]:

FJ179575; rpb2: FJ179611. REFERENCES: Chamberlain et al. (2004), Jaklitsch (2011).

Trichoderma ochroleucum (Berk. & Ravenel) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807440

= Hypocrea ochroleuca Berk. & Ravenel, in Berkeley, Grevillea 4: 14 (1875).

HOo.otyPe: K(M) 56075; ex-type culture: n.a.; repres. CBS 119502 = C.P.K. 1895 = Hypo

274.

PHYLOG.: sect. Trichoderma. REPRES. SEQUENCES: tefl: FJ860659; rpb2: FJ860556.

REFERENCE: Jaklitsch (2011). NoTE: GenBank accessions for G.J.S. 01-234 and G.J.S.

01-265 may represent Trichoderma albofulvum (see above).

New Trichoderma combinations ... 151

Trichoderma orientale (Samuels & Petrini) Jaklitsch & Samuels, comb. nov.

MycoBank MB807441

= Hypocrea orientalis Samuels & Petrini, in Samuels, Petrini, Kuhls,

Lieckfeldt & Kubicek, Stud. Mycol. 41: 30 (1998).

Ho otype: BPI 1109853; ex-type culture: CBS 130428 = G/J.S. 88-81, repres. CBS

131488, GJ.S. 09-784.

PHYLOG.: sect. Longibrachiatum. REPRES. SEQUENCES: tefl: EU401581, JN175573,

JQ685868; rpb2: JN175522, JQ685884. REFERENCES: Samuels et al. (1998, 2012).

Trichoderma parapiluliferum (B.S. Lu, Druzhin. & Samuels) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807442

= Hypocrea parapilulifera B.S. Lu, Druzhin. & Samuels, in Lu, Druzhinina, Fallah,

Chaverri, Gradinger, Kubicek & Samuels, Mycologia 96: 331 (2004).

HooryPe: BPI 112832; ex-type culture: CBS 112771 = G.J.S. 91-60; repres. CBS 120921

= C.PK. 1908.

PHyLoG.: Pachybasium Core Group, conidia hyaline. REPRES. SEQUENCES: tefl1:

FJ179578, AY937444; rpb2: FJ179614. REFERENCES: Lu et al. (2004), Jaklitsch (2011).

Trichoderma parmastoi (Overton) Jaklitsch & Voglmayr, comb. nov.

MycoBANK MB807443

= Hypocrea parmastoi Overton, in Overton, Stewart & Geiser, Stud. Mycol. 56: 62 (2006).

Ho.ortyPe: BPI 843639; isotype TAA(M) 169055; ex-type culture: TFC 97-143; repres.

CBS 121139.

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. RepREs.

SEQUENCES: fef1: FJ860668, DQ834456 (exon); rpb2: FJ860567, DQ834463. REFERENCES:

Overton et al. (2006b), Jaklitsch (2011).

Trichoderma patella (Cooke & Peck) Jaklitsch & Voglmayr, comb. nov.

MycoBANK MB807444

= Hypocrea patella Cooke & Peck, in Peck, Ann. Rep. N.Y.

St. Mus. nat. Hist. 29: 57 (1878 [“1875”]).

HototyPe: NYS; isotype: BPI 631626; ex-type culture: n.a.; repres. CBS 110082 = G.J.S.

95-173, CBS 110084 = G,J.S. 96-198 = ATCC 208855.

PHYLOG.: unknown, remotely associated with sect. Longibrachiatum. REPRES.

SEQUENCES: tef1: AY937428; rpb2: n.a.. REFERENCE: Dodd et al. (2002).

Trichoderma peltatum (Berk.) Samuels, Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807511

= Sphaeria peltata Jungh., Verh. Batav. Genootsch. Kunst. Wet.

17(3): 20 (1838), nom. illegit., non DC. 1805.

= Hypocrea peltata Berk., Hooker’s J. Bot. Kew Gard. Misc. 3: 205 (1851).

Ho tortyPe: L 00532089 (Junghuhn 103); ex-type culture: n.a.; repres. CBS 127107 =

GJ.S. 09-1550, CBS 127115 = GJ.S. 08-207, G.J.S. 10-103, G.J.S. 10-104, G.J.S. 10-105,

CBS 120951.

152 ... Jaklitsch & Voglmayr

PHYLOG.: unnamed clade, close to T. sambuci and T. tremelloides; anamorph not formed.

REPRES. SEQUENCES: fef1: n.a.; rpb2: HQ260609, HQ260610. REFERENCE: Samuels &

Ismaiel (2011).

Note: The original name of this taxon, Sphaeria peltata, is an illegitimate later homonym

and cannot act as basionym for new combinations (ICN Art. 6.10). The basionym is

the earliest legitimate name, Hypocrea peltata Berk. 1851, which is to be treated as a

replacement name for the illegitimate S. peltata (ICN 58.1).

Trichoderma pezizoides (Berk. & Broome) Samuels, Jaklitsch & Voglmayr,

comb. nov.

MycoBANnkK MB807445

= Hypocrea pezizoides Berk. & Broome, J. Linn. Soc., Bot. 14(74): 112 (1873).

HOoLotyPeE: n.d.; ex-type culture: n.a.; ¢repres.: CBS 101131 = G.J.S. 97-83 = C.P.K. 775,

G,J.S. 01-231 (Thailand). Originally described from Sri Lanka.

PHYLOG.: sect. Trichoderma. REPRES. SEQUENCES: tefl: AY225859; rpb2: JN715610,

AF545564. REFERENCE: Atanasova et al. (2013).

Trichoderma protopulvinatum (Yoshim. Doi) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807446

= Hypocrea protopulvinata Yoshim. Doi, Bull. natn. Sci. Mus., Tokyo 15: 695 (1972).

Ho.otype: TNS-F-223421; isotype NY 00965650; ex-type culture: CBS 739.83; repres.

C.P.K. 2434, CBS 121274 = C.P.K. 2430.

PHYLOG.: sect. Hypocreanum; anamorph acremonium-like, conidia hyaline. REPREs.

SEQUENCES: tefl: FJ860676; rpb2: FJ860574. REFERENCES: Overton et al. (2006a),

Jaklitsch (2011).

Trichoderma pulvinatum (Fuckel) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807447

= Hypocrea pulvinata Fuckel, Jb. nassau. Ver. Naturk. 23-24: 185 (1870).

LECTOTYPE: FH 00284789 (Fuckel 876); ex-type culture: n.a.; repres. CBS 121279.

PHYLOG.: sect. Hypocreanum; anamorph acremonium- to verticillium-like, conidia

hyaline. REPRES. SEQUENCES: tefl: FJ860683; rpb2: FJ860577. REFERENCES: Overton et

al. (2006a), Jaklitsch (2011).

Trichoderma rhododendri (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807448

= Hypocrea rhododendri Jaklitsch & Voglmayr, in Jaklitsch,

Fungal Diversity 48: 199 (2011).

Ho.otyPe: WU 29442 (Hypo 209); ex-type culture: CBS 119288 = C.P.K. 2015.

Puy_oa.: Psychrophila clade, anamorph unknown. REPRES. SEQUENCES: fef1: FJ860685;

rpb2: FJ860578. REFERENCE: Jaklitsch (2011).

Trichoderma rodmanii (Samuels & P. Chaverri) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807449

= Hypocrea rodmanii Samuels & P. Chaverri, Mycol. Prog. 7: 213 (2008).

Ho otype: BPI 1112859; ex-type culture: CBS 120895 = GJ.S. 91-88; repres. CBS

121553.

New Trichoderma combinations ... 153

PHYLOG.: Brevicompactum clade. REPRES. SEQUENCES: tefl: FJ860687, EU338286; rpb2:

FJ860580, EU338324. REFERENCES: Degenkolb et al. (2008), Jaklitsch (2011).

Trichoderma sambuci (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807450

= Hypocrea sambuci Jaklitsch & Voglmayr, in Jaklitsch, Fungal Diversity 48: 213 (2011).

Ho.otype: WU 29463 (Hypo 198); ex-type culture: n.a.; repres. CBS 126958 = S 94.

PHYLOG.: unnamed clade, close to T: tremelloides; conidia hyaline. REPRES. SEQUENCES

(based on DNA from the specimen WU 29467 = Hypo 426): tefl: FJ860693; rpb2:

FJ860585. REFERENCE: Jaklitsch (2011).

Trichoderma semiorbis (Berk.) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807451

= Sphaeria semiorbis Berk., J. Bot. (Hooker) 2: 146 (1840).

= Hypocrea semiorbis (Berk.) Berk., in Hooker, Fl. Tasman. 2: 278. (1859 [“1860"]).

HOo.oryPe: K; ex-type culture: n.a.; repres. CBS 244.63 = DAOM 167636, G.J.S. 99-108,

G.J.S. 99-109.

PHyLoG.: Semiorbis clade; conidia hyaline. REPRES. SEQUENCES: tefl: AF545568; rpb2:

JN133567, AF545522. REFERENCE: Chaverri et al. (2003).

Trichoderma spinulosum (Fuckel) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807452

= Hypocrea spinulosa Fuckel, Jb. nassau. Ver. Naturk. 23-24: 184 (1870).

Ho .oryPe: Fuckel (G); ex-type culture: n.a.; repres. CBS 310.50, CBS 311.50 = C.P.K.

1510, CBS 121272 = C.P.K. 2464.

PHyLoG.: Spinulosa clade, species with green ascospores; anamorph not formed.

REPRES. SEQUENCES: tef1: FJ860701; rpb2: FJ860591. REFERENCES: Chaverri & Samuels

(2003), Jaklitsch (2009).

Trichoderma stellatum (B.S. Lu, Druzhin. & Samuels) Jaklitsch & Voglmayr,

comb. nov.

MycoBank MB807453

= Hypocrea stellata B.S. Lu, Druzhin. & Samuels, in Lu, Druzhinina, Fallah,

Chaverri, Gradinger, Kubicek & Samuels, Mycologia 96: 333 (2004).

Ho.otyPe: PDD 77488; ex-type culture: CBS 112265 = GJ.S. 99-122 (as ‘G.J.S. 99-222’

in GenBank).

PHyLoG.: Pachybasium Core Group, conidia hyaline. REPRES. SEQUENCES: tef1:

AY937445, AY240855 (intron5 only); rpb2: n.a. REFERENCE: Lu et al. (2004).

Trichoderma subsulphureum (Syd.) Jaklitsch & Voglmayr, comb. nov.

MycoBANK MB807454

= Hypocrea subsulphurea Syd., in De Wildeman, Etudes

Fl. Bas- Moyen-Congo 3: 15 (1909).

Ho .orype: ?lost; ex-type culture: n.a.; ¢repres. M141 (?BPI).

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. RepREs.

SEQUENCES (from teleomorph of M141): tefl: DQ835492; rpb2: DQ835522. REFERENCE:

Overton et al. (2006b).

154 ... Jaklitsch & Voglmayr

Trichoderma sulawesense (Yoshim. Doi) Jaklitsch & Voglmayr, comb. nov.

MycoBAnk MB807455

= Hypocrea sulawesensis Yoshim. Doi, in Samuels, Doi &

Rogerson, Mem. N. Y. bot. Gdn 59: 23 (1990).

HoLortyPe: BO; isotypes NY 01169121, NY 00967625, TNS-F-243769, ex-type culture:

G,J.S. 2006 (lost), repres. G.J.S. 85-228.

PHYLOG.: unnamed clade; species with green ascospores; anamorph forming elongate

hyaline phragmoconidia in addition to green unicellular conidia. REPRES. SEQUENCES:

tefl: AY737730 + AY392002; rpb2: AY391954. REFERENCE: Chaverri & Samuels (2003).

Trichoderma sulphureum (Schwein.) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807456

= Sphaeria sulphurea Schwein., Trans. Amer. philos. Soc., Ser. 2, 4 (2): 193 (1832).

= Hypocrea sulphurea (Schwein.) Sacc., Syll. Fung. (Abellini) 2: 535 (1883).

Ho.ortyPe: n.d.; isotype: K, ex herb Schweinitz; not epitypified; ex-type culture: n.a.;

repres. C.P.K. 1593,-CBS 119929. =C.P.K..1598.

PHYLOG.: sect. Hypocreanum; anamorph acremonium- to verticillium-like, conidia

hyaline. REPRES. SEQUENCES: tefl: FJ860709; rpb2: FJ860599. REFERENCES: Overton

(2006b), Jaklitsch (2011).

Trichoderma victoriense (Overton) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807457

= Hypocrea victoriensis Overton, in Overton, Stewart

& Geiser, Stud. Mycol. 56: 55 (2006).

Ho .ortyPe: BPI 747361; ex-type culture: G.J.S. 99-200; repres. C.P.K. 3565.

PHYLOG.: sect. Hypocreanum; anamorph verticillium-like, conidia hyaline. RepREs.

SEQUENCES: tef1: FJ860718; rpb2: DQ835517. REFERENCE: Overton et al. (2006b).

Trichoderma virescentiflavum (Speg.) Jaklitsch & Voglmayr, comb. nov.

MycoBank MB807458

= Hypocrea virescentiflava Speg., Boln Acad. nac. Cienc. Cordoba 11: 529 (1889).

Ho .ortyPe: LPS 1726 (J. Puiggari 2353), ex-type culture: n.a.; repres. P.C. 278 (lost after

determination of DNA data).

PuHyLoa.: Virescentiflava clade, species with green ascospores. REPRES. SEQUENCES: tef1:

AY737749 + AY392007; rpb2: AY391959. REFERENCE: Chaverri & Samuels (2003).

Acknowledgments

We sincerely thank the pre-submission reviewers Walter Gams and Gary Samuels,

and Shaun Pennycook for his considerable nomenclatural contribution. The financial

support by the Austrian Science Fund (FWF; project P22081-B17) is gratefully

acknowledged.

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MYCOTAXON

http://dx.doi.org/10.5248/126.157

Volume 126, pp. 157-166 October-December 2013

Lactarius taxa by Zdenék Schaefer -

types and other collections in PRM

JAN HOLEc *, KAMILA PESICOVA & LENKA EDROVA

National Museum, Mycological Department,

Cirkusova 1740, CZ-193 00 Praha 9, Czech Republic

* CORRESPONDENCE TO: jan_holec@nm.cz

ABSTRACT— Extant PRM herbarium vouchers of Lactarius taxa described, validated, or

neotypified by Zdenék Schaefer are enumerated and discussed. For each taxon, the holotype,

paratypes, other original materials, and later collected specimens identified or revised by

Schaefer are cited. The nomenclatural status of each taxon is evaluated and information from

published revisions/discussions is summarized. These data provide a basis both for thorough

revisions by classical and molecular methods and for future lecto- or neotypifications.

Such revisions are encouraged because some of Schaefer’s taxa (e.g., L. albocremeus) may

represent good species while others should clearly be synonymized with earlier published

taxa. Currently accepted taxa are L. pilatii and L. bertillonii (= Lactifluus bertillonii).

Key worps—Basidiomycota, Russulales, taxonomy, nomenclature, Europe

Introduction

Zdenék Schaefer (1906-1974), a chemical engineer in the glass industry,

was a Czech amateur mycologist who became interested in fungi in 1936. In

1945 he decided to focus on Lactarius (Herink 1977), eventually publishing 29

works on the genus, mostly in the leading Czech journal, CesKA MYKOLOGIE

(now CzEcH Myco .ocy). For his biographies and bibliographies, see Pouzar

(1966) and Herink (1977). Schaefer was known for his detailed approach

and his narrow species concept. Material studied by him is now stored in the

mycological herbaria of the National Museum in Prague (PRM), the Moravian

Museum in Brno (BRNM), and (rarely) in the Slovak National Museum

in Bratislava (BRA). After his death, his family threw away his personal

herbarium (including some types; Z. Pouzar, pers. comm. 2013) because of

insect infestation; consequently, only a small part of his collection is conserved

in PRM (mostly collections from earlier periods of Schaefer’s work).

158 ... Holec, Pesicova & Edrova

In the late 1960s, Schaefer prepared a Czech-language monograph of

Czechoslovakian Lactarius species. However, the work remained unpublished

because of political restrictions following the 1968 “Prague spring” and Soviet

occupation. The original typewritten manuscript, accompanied by Schaefer's

coloured pencil drawings of Lactarii, is deposited in the Museum of Eastern

Bohemia in Hradec Kralové (HR), and photocopies are available in PRM and

BRNM. He had planned to publish about 70 new taxa (species, varieties, forms)

in his monograph.

Schaefer published more than 20 new taxa of Lactarius (species and

varieties) in various papers (1948, 1956, 1957, 1958, 1960, 1966, 1968,

1970, 1972, 1979), the last of which (Schaefer 1979) was prepared from his

unpublished paper by the CeskA Myxotoarte Editorial Board (Z. Pouzar, pers.

comm. 2013) and published five years after his death. In other papers Schaefer

also validated names published by Smotlacha (1947) and Neuhoff (1956) and

designated a neotype for a 1920 Velenovsky name. Schaefer's new taxa, which

might represent real species or varieties, need a thorough revision. The only

broadly focused revision was done by Basso (1999), who used only a part of the

available vouchers.

Our intention here is to clarify which types, other original materials, and

later collections identified by Schaefer are stored in PRM. Taxa supported by

these vouchers can be studied both by classical and DNA methods, which is

important for further development of Lactarius taxonomy. A paper devoted to

collections in BRNM should be prepared by local curators.

Materials & methods

Only published Lactarius taxa (both valid and invalid names) by Schaefer are

included. They are arranged alphabetically under Lactarius, including two recent

synonyms in Lactifluus. New taxa proposed in Schaefer’s unpublished monograph (see

Introduction) are omitted, as they are not published effectively. Each entry starts with

bibliographic data on the Latin, Czech, and/or German descriptions. As Schaefer's

publications are poorly arranged and tend to confine detailed information to the Czech

text, we compiled and translated the important voucher data to English. The following

materials from PRM are cited for each taxon: holotype, paratypes, other original materials

(not cited with all details in the protologue but certainly used for its preparation), and

other collected specimens subsequently identified or revised by Schaefer. The localities

are given not in their original form but are amended based on data accumulated from

herbarium labels, Latin descriptions, and Czech or German descriptions/discussions

and supplemented with current geographic data. Taxonomic and nomenclatural terms

are in accordance with the current International Code of Nomenclature for algae, fungi,

and plants (McNeill et al. 2012), abbreviated as ICN. Herbarium abbreviations follow

Thiers (2013). Taxonomic remarks are based on synonymies, discussions, and revisions

published in European Lactarius monographs (Korhonen 1984, Heilmann-Clausen et

al. 1998, Basso 1999, Kalamees 2011).

Lactarius taxa by Zdenék Schaefer in PRM (Czech Republic) ... 159

Results & discussions

Lactarius albocremeus Z. Schaef., Ceska Mykologie 12: 211, 1958.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 12: 211, 1958.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 12: 208-209, 1958.

Ho.LotyPeE — designated. Slovakia, Western Carpathians, NNE of Poprad, between the

villages of SpiSska Bela and Lendak, site called ,,Blomwiesen’, in fen meadow, close to

Betula and Salix trees, alt. about 800 m, 11 Aug. 1957 leg. K. Kriz (BRNM; see Antonin

& Vagner 1999).

IsoTyPE — a duplicate of the holotype collection (PRM 584029).

OTHER COLLECTIONS IDENTIFIED BY SCHAEFER — none. There are three later collected

specimens identified (respectively) by J. Kubicka, M. Svréek and Z. Pouzar, however,

they represent neither original materials nor collections revised by Schaefer.

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS —Basso (1999: 385, 389-390) regarded L. albocremeus

as identical with L. scoticus Berk. & Broome. However, the Czech Lactarius

specialist Pouzar feels that the current concept of L. scoticus differs from

Berkeley & Broome’s original concept (Z. Pouzar, pers. comm., May 2013). This

discrepancy should be clarified and the name L. albocremeus considered as a

possible name for L. scoticus auct. non Berk. & Broome.

Lactarius chrysophyllus Z. Schaef., Ceska Mykologie 11: 52, 1957.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 11: 52-53, 1957.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 11: 50-52, 1957.

HOoLotyPeE — not designated.

OTHER TYPE COLLECTIONS — the protologue mentions six localities from the Czech

Republic (Téchlovice, Cebinka, Roviny u Heroltic, Radikovice, Kuiim, Sdzava nad

Sazavou), indicating potential syntypes, but there are no corresponding vouchers in

PRM.

OTHER COLLECTIONS IDENTIFIED BY SCHAEFER — CZECH REPUBLIC, BOHEMia,

Rokycany, 22 Aug. 1942, collector unknown (PRM 195218). The collection was revised

by Basso (1999: 144).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — Basso (1999: 138, 143-144) considered L. chrysophyllus

a synonym of L. hysginus (Fr.) Fr.

Lactarius cupricolor Z. Schaef., Ceska Mykologie 20: 158, 1966.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 20: 158, 1966.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 20: 154-156, 1966.

DETAILED GERMAN DESCRIPTION: Ceska Mykologie 20: 156-157, 1966.

Ho.LotypPE — designated. Czech Republic, Turnov district, Sychrov: castle park, under

Quercus, Picea, Abies and distant Fagus, 27 Aug. 1961 leg. Z. Schaefer (PRM 584026).

160 ... Holec, PeSicova & Edrova

PARATYPES — CZECH REPUBLIC, CENTRAL BOHEMIA, OleSovice: Struharov, 15 Aug.

1957 leg. K. Kult (PRM 193972); Praha, Divoka Sarka, Quercus forest, 7 Sep. 1937 leg.

J. Herink (PRM 193865). (Five other records mentioned in the protologue are potential

paratypes, but there are no corresponding vouchers in PRM.)

NOMENCLATURAL STATUS — Valid, legitimate.

TAXONOMIC REMARKS — species of unclear identity. For type study and

discussion see Basso (1999: 131-132), who placed it under L. vietus but also

compared it with L. pallidus.

Lactarius eburneus Z. Schaef., Ceska Mykologie 33: 4, 1979, nom. illegit.,

non Thiers 1957.

LATIN DESCRIPTION: Ceska Mykologie 33: 4, 1979.

GERMAN Discussion: Ceska Mykologie 33: 4-5, 1979.

Ho.otyPE — designated. Czech Republic, central Bohemia, near Gol¢tv Jenikov, mixed

forest, 22 Jul. 1949 leg. Z. Schaefer (herb. Z. Schaefer no. 112). Location unknown but

probably discarded by Schaefer's family.

NOMENCLATURAL STATUS — valid, illegitimate (a later homonym of L. eburneus

Thiers).

TAXONOMIC REMARKS — Basso (1999: 723, 735) considered L. eburneus Z.

Schaef. identical with L. glaucescens Crossl. (= Lactifluus glaucescens (Crossl.)

Verbeken).

“Lactarius echinosporus” Z. Schaef., Ceska Mykologie 14: 237, 1960, nom. inval.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 14: 237, 1960.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 14: 233-235, 1960.

Ho.LotyPeE — not designated.

SYNTYPES — Czech Republic, Central Bohemia, SW of Prague: Barrandian region, also

called Cesky kras Karst, e.g. near Karl&8tejn castle, on calcareous bedrock, 1949 leg. K.

Kult; 9 Sep. 1958 leg. Z. Schaefer (vouchers nonexistent).

OTHER COLLECTION IDENTIFIED BY SCHAEFER (not mentioned in the protologue) —

CZECH REPUBLIC, CENTRAL BOHEMIA, between KarlStejn castle and the village of

Morina, 27 Aug. 1951 leg. A. Pilat (PRM 561154).

NOMENCLATURAL STATUS — invalid: no holotype designated.

TAXONOMIC REMARKS — a fungus belonging to Russula delica group (Basso

1999: 330, 454-455).

Lactarius hortensis Velen., Ceské houby [Czech fungi], part 1: 163, 1920, var.

hortensis

ORIGINAL CZECH DESCRIPTION: Ceské houby [Czech fungi], part 1: 163-164, 1920.

LATIN DESCRIPTION BY SCHAEFER: Ceska Mykologie 26: 143, 1972.

Lactarius taxa by Zdenék Schaefer in PRM (Czech Republic) ... 161

HoLotyPe — nonexistent (Holec & Sukova 2007). Czech Republic, central Bohemia,

SE of Prague, Mnichovice, Velenovsky’s garden, under Corylus (Velenovsky 1920: 163).

NEOTYPE — Slovakia, Banska Bystrica district, Harmanec, stand of Corylus avellana, 23

Aug. 1966 leg. Z. Schaefer (PRM 621095, designated by Schaefer 1972: 143).

NOMENCLATURAL STATUS — Valid, legitimate. The neotype designation seems

inappropriate as it originates from a different country than the original,

particularly as the species is common in the holotype area.

TAXONOMIC REMARKS —Korhonen (1984: 130) regards L. hortensis as a good

species, while Heilmann-Clausen et al. (1998: 52), Basso (1999: 117), Kalamees

(2011: 122) regard it as identical with L. pyrogalus (Bull.) Fr.

Lactarius hradecensis Z. Schaef., Ceska Mykologie 2: 85, 1948.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 2: 85, 1948.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 2: 83-85, 1948.

HOLOTYPE — nonexistent. Czech Republic, eastern Bohemia, near Hradec Kralové,

forest called Dehetnik, under Picea and admixed Pinus, Betula, Quercus, 26 Aug. 1945

leg. Z. Schaefer (Schaefer 1948: 85).

NEOTYPE — Czech Republic, eastern Bohemia, near Hradec Kralové, forest called

Dehetnik, under Picea, Quercus, Betula, 22 Aug. 1961 leg. Z. Schaefer (PRM 584032;

designated by Schaefer 1966: 152, 157, as “typus”).

OTHER COLLECTIONS MENTIONED IN NEOTYPE PUBLICATION (Schaefer 1966) —

CZECH REPUBLIC, Bonemia, Turnov, Sychrov: castle park, 24 Aug. 1961 leg. Z.

Schaefer (PRM 584025); Turnov, Valdstyn (= Vald&stejn), 19 Aug. 1945 leg. J. Herink

(PRM 193660).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — species of uncertain identity. Basso discussed it

under L. quietus (Fr.) Fr. (Basso 1999: 504-506) and mentioned it briefly under

L. subdulcis (Pers.) Gray (Basso 1999: 516).

Lactarius moravicus Z. Schaef., Ceska Mykologie 33: 10, 1979.

LATIN DESCRIPTION + GERMAN DISCUSSION: Ceska Mykologie 33: 10, 1979.

Ho.LotyPE — designated. Czech Republic, southwestern Moravia, Tel¢, near fish-pond

Parezity rybnik, Picea forest, 4 Aug. 1958 leg. Z. Schaefer (herb. Z. Schaefer no. 58: 21).

Location unknown but probably discarded by Schaefer's family.

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — species of unclear identity. Mentioned under

L. vellereus var. hometii (Gillet) Boud. by Basso (1999: 722).

Lactarius pearsonii Z. Schaef., Ceska Mykologie 22: 19, 1968.

= Lactarius mairei var. zonatus A. Pearson, Naturalist, London: 100, 1950.

LATIN DESCRIPTION: Ceska Mykologie 22: 19, 1968.

162 ... Holec, Pesicova & Edrova

CzECH piscusston: Ceska Mykologie 22: 17, 1968. Detailed Czech description cited by

Schaefer (1968: 17) was published by Pouzar (1954: 40-41, as “L. aurantiacoochraceus”).

Ho.LotyPeE — designated. Czech Republic, central Bohemia, Beroun district, Karlstejn

nature reserve, valley of the Bubovicky potok stream, site above its waterfalls, under

Fraxinus, Acer, Salix, 23 Aug. 1953 leg. Z. Pouzar, F. Kotlaba & FE Hiebik (PRM 193531).

OTHER TYPE COLLECTIONS — Schaefer (1968: 17) wrote that the species is not rare

in former Czechoslovakia. Although he listed localities in central Bohemia (vouchers

in PRM), southern Moravia (vouchers in BRNM), and southern Slovakia (voucher in

PRM), he did not cite any collections as paratypes. The existing vouchers represent

original materials on which the protologue is based.

OTHER ORIGINAL MATERIALS IN PRM (identified as L. mairei var. zonatus by their

collectors) — CZECH REPUBLIC, Bouemtia, Praha: Kinského sady park, 9 Aug. 1968

leg. E. Wichansky (PRM 682631); 31 Aug. 1964 leg. V. Brambora (PRM 602362). —

SLOVAKIA, SOUTHEASTERN SLOVAKIA, Slovensky kras Karst, the village of Vidova

near PleSivec, 1 Jul. 1963 leg. F. Kotlaba et Z. Pouzar (PRM 583432).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — Schaefer (1968: 17) rejected the synonymy of

L. mairei var. zonatus with L. aurantiacoochraceus Lj.N. Vassiljeva, which had

been proposed by Pouzar (1954). Heilmann-Clausen et al. (1998: 164), Basso

(1999: 374, 378-379), and Kalamees (2011: 106) regard L. pearsonii as identical

with L. mairei Malencon.

Lactarius pilatii Z. Schaef., Ceska Mykologie 22: 18, 1968.

LATIN DESCRIPTION: Ceska Mykologie 22: 18-19, 1968.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 22: 17-18, 1968.

Hototyre — designated. Czech Republic, southwestern Bohemia, Sumava Mts., near

the village of Horska Kvilda, Zhurska slat bog, alt. about 1100 m, under Betula pubescens,

22 Aug. 1966 leg. A. Pildt (PRM 624664).

ParaTyPEs — Czech Republic, southwestern Bohemia, Sumava Mts., near the village

of Horska Kvilda, Zhurska slat bog, alt. about 1100 m, under Betula pubescens, 16 Aug.

1965 leg. A. Pilat (PRM 624669, 624658); 16 Aug. 1967 leg. A. Pilat (PRM 628888); 18

Aug. 1967 leg. A. Pilat (PRM 628889); 26 Aug. 1967 leg. A. Pilat (PRM 628886, 628885).

OTHER COLLECTION BY A. PILAT FROM THE TYPE LOCALITY (not mentioned by Schaefer

1968 but most probably studied by him) — 19 Aug. 1967 (PRM 628887).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — good species (Heilmann-Clausen et al. 1998: 62-63,

Basso 1999: 111-117, Kalamees 2011: 174). Kalamees (2011: 174) considers

L. moseri Harmaja synonymous with L. pilatii. Kalamees cites photographs by

Korhonen (1984: 134, labelled L. moseri).

Lactarius pinicola Smotl. ex Z. Schaef., Schweizerische Zeitschrift fiir Pilzkunde 48:

141, 1970.

“Lactarius deliciosus var. pinicola” Smotlacha, Atlas hub jedlych a nejedlych [Atlas

of edible and inedible fungi]: 219, 1947, nom. inval. [no Latin description].

Lactarius taxa by Zdenék Schaefer in PRM (Czech Republic) ... 163

LATIN DIAGNOSIS: Schaefer (1970: 141-142).

GERMAN DISCUSSION: Schaefer (1970: 139-142).

HoLotyPE — designated. Czech Republic, northern Bohemia, district Jablonec

nad Nisou, near Turnov, Frydstejn: near the village of Snéhov, mixed forest (Pinus,

Picea), under Pinus, 18 Oct. 1962 leg. Z. Schaefer (PRM 584024, originally labelled as

L. salmonicolor).

OTHER COLLECTION IDENTIFIED BY SCHAEFER — CZECH REPUBLIC, BOHEMia,

Turnov, Titi near Radimovice, 21 Sep. 1966 leg. Z. Schaefer (PRM 709711).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — identical with L. deliciosus (L.) Gray (Heilmann-

Clausen et al. 1998: 140; Basso 1999: 253, 259; Nuytinck & Verbeken 2005: 131;

Kalamees 2011: 70).

Lactarius pyrogalus var. umbrosus Z. Schaef., Ceska Mykologie 26: 145, 1972.

CZECH AND LATIN DIAGNOSES: Ceska Mykologie 26: 145, 1972.

Ho.LotyPe — designated. Czech Republic, Bohemia, NE of Pardubice, near the village

of Bélecko, Picea forest, 15 Aug. 1944 leg. Z. Schaefer (herb. Z. Schaefer). Location

unknown but probably discarded by Schaefer's family.

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — taxonomic value of this variety is unclear. Discussed

under L. pyrogalus by Basso (1999: 124).

“Lactarius russuloides” Z. Schaef., Ceska Mykologie 12: 205, 1958, nom. inval.

LATIN DIAGNOSIS/DESCRIPTION: none.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 12: 205-206, 1958.

HoLotyPE — not designated. The description is based on 3 records from the Czech

Republic (Telé, Hulistata; Trebon, Spoli; Trebon, Domanin). None of these specimens

is held in PRM.

OTHER COLLECTIONS IDENTIFIED OR REVISED BY SCHAEFER — CZECH REPUBLIC,

BouEMIA, Trhové Sviny, 17 Sep. 1964 leg. J. Kubicka (PRM 603730); Tabor, Sudomérice:

Nemyil, 9 Sep. 1949 leg. M. Svréek (PRM 615083); Poricany: forest Kersko, 11 Oct. 1955

leg. Z. Pouzar (PRM 621218); Trebon, Kozakovice, 15 Oct. 1967 leg. J. Kubicka et Z.

Pouzar (PRM 709721); Trebon, Domanin, 23 Sep. 1964 leg. J. Kubicka (PRM 603726); 7

Sep. 1964 (PRM 603728).

NOMENCLATURAL STATUS — invalid: lacking Latin description or diagnosis

and holotype designation. Schaefer erroneously treated the name as a nomen

novum (= replacement name; ICN Art. 6.11) for “L. musteus” sensu Favre

(in Konrad & Maublanc 1937: tab. 331), Kithner & Romagnesi (1953: 477),

and Neuhoff (1956: 130) and “L. pubescens” sensu Bresadola (1929: tab. 360).

Because none of these is a valid name, L. russuloides must be interpreted as a sp.

nov., rather than a nom. nov.

164 ... Holec, PeSicova & Edrova

TAXONOMIC REMARKS — Basso (1999: 171, 175) considers L. russuloides is

identical with L. musteus Fr.

Lactarius syringinus Z. Schaef., Ceska Mykologie 10: 171, 1956.

LATIN DIAGNOSIS/DESCRIPTION: Ceska Mykologie 10: 171-172, 1956.

DETAILED CZECH DESCRIPTION: Ceska Mykologie 10: 168-171, 1956.

HoLotyPE — designated. Czech Republic, central Bohemia, E of Praha, near RadoSovice

(,,RadeSovice*) close to Ri¢any, site called Vojkov (name of a gamekeeper’s lodge and

a settlement E of the town of Rigany), in deciduous forest with admixed Picea and

Pinus, Aug. 1953 leg. I. Charvat. Location unknown and possibly nonexistent; probably

not preserved by Schaefer who obtained the material by surface mail from I. Charvat

(Schaefer 1956: 169).

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — Heilmann-Clausen et al. (1998: 60-61) regard

L. syringinus as a good species. Basso (1999: 130-131) discusses it under L. vietus

(Fr.) Fr. and Kalamees (2011: 158) synonymized it with L. vietus. Based on more

collections and sequences, both from Europe and Russia, L. syringinus really

seems to fall within the variation of L. vietus (Verbeken, pers. comm. 2013).

Lactarius vellereus var. bertillonii Neuhoff ex Z. Schaef., Ceska Mykologie 33: 9,

1972.

= Lactarius bertillonii (Neuhoff ex Z. Schaef.) Bon, Doc.

Mycol. 10(37-38): 92, 1980 [“1979”].

= Lactifluus bertillonii (Neuhoff ex Z. Schaef.) Verbeken,

Mycotaxon 118: 451, 2012 [“2011”].

“Lactarius vellereus var. bertillonii” Neuhoff, Die Milchlinge:

93, 1956, nom. inval. [no Latin description].

LATIN DIAGNOSIS BY SCHAEFER: Ceska Mykologie 33: 9, 1979.

GERMAN DISCUSSION BY SCHAEFER: Ceska Mykologie 33: 8-9, 11, 1979.

HoLotypPe — designated. Czech Republic, central Bohemia, Golctiv Jenikov, mixed

forest: Picea, Pinus, Abies, Betula, Quercus, 22 Jul. 1949 leg. Z. Schaefer (herb. Z. Schaefer

no. 41). Currently nonexistent, probably thrown away by Schaefer's family.

NOMENCLATURAL STATUS — valid, legitimate.

TAXONOMIC REMARKS — this variety is now generally accepted as a good

species, L. bertillonii (= Lactifluus bertillonii) (Korhonen 1984; Heilmann-

Clausen et al. 1998: 254-255; Basso 1999: 709, 713, 718; Kalamees 2011: 58).

Conclusions

We have shown that nine Lactarius taxa described by Schaefer are

documented by vouchers in PRM. Although some holotypes are missing or

nonexistent, paratypes, other original materials, or later collections identified

by Schaefer are available for most of them. The taxa should be carefully studied

Lactarius taxa by Zdenék Schaefer in PRM (Czech Republic) ... 165

by classical and molecular methods to determine whether they represent good

species (e.g. L. albocremeus) or placed into synonymy with earlier published taxa

(their current revisions are insufficient). Currently accepted taxa are L. pilatii

and L. bertillonii (= Lactifluus bertillonii). The accumulated data on available

vouchers provide a good basis for both revisions and future typifications by

experienced Lactarius specialists.

Acknowledgements

We thank Zdenék Pouzar (Prague, Czech Republic) for information on Schaefer's

private herbarium and some unpublished biographic data. We thank the reviewers,

Vladimir Antonin (Moravian Museum, Brno, Czech Republic) and Annemieke Verbeken

(Ghent University, Gent, Belgium) for their valuable comments and Shaun Pennycook

(Landcare Research, Auckland, New Zealand) and Lorelei L. Norvell (Portland, USA)

for nomenclatural and linguistic corrections. The work was financially supported by the

Ministry of Culture of the Czech Republic (NAKI project no. DF12PO1OVV021).

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MY COTAXON

http://dx.doi.org/10.5248/126.167

Volume 126, pp. 167-176 October-December 2013

Phytophthora virginiana sp. nov., a high-temperature

tolerant species from irrigation water in Virginia

XIAO YANG* & CHUANXUE HONG

Department of Plant Pathology, Physiology, and Weed Science,

Hampton Roads Agricultural Research and Extension Center, Virginia Tech,

Virginia Beach, VA 23455

* CORRESPONDENCE TO: yxiao9@vt.edu

ABSTRACT — Isolates belonging to a previously unknown species of Phytophthora were

recovered from irrigation reservoirs at several ornamental plant nurseries in Virginia.

Morphological features of this species include abundant lateral chlamydospores in

clarified V8 juice agar and carrot agar and terminal, ovoid to obpyriform, nonpapillate and

noncaducous sporangia in soil water extract. All tested isolates are silent A’ mating type. They

did not produce any sexual structures but induced A’ mating type testers of P. cinnamomi

and P. cryptogea to produce gametangia. Sequence analysis of the internal transcribed spacer

(ITS) region confirmed that this species belongs to the high-temperature tolerant cluster

within Phytophthora clade 9. Its ITS sequence differs from those of its two closest relatives,

P. hydropathica and P. parsiana, by >26 bp. Based on these morphological, physiological, and

molecular characteristics, we recognise it as a new species, described here as Phytophthora

virginiana.

KEY worDs —aquatic habitat, oomycetes, phylogeny, self-sterile, taxonomy

Introduction

The genus Phytophthora de Bary contains a number of destructive plant

pathogens (Erwin & Ribeiro 1996). They are responsible for many historical

and current epidemics of plant diseases including late blight of potatoes caused

by P infestans, ink disease of chestnut trees caused by P cambivora, and sudden

oak death caused by P ramorum (Erwin & Ribeiro 1996, Goheen et al. 2002,

Rizzo et al. 2002). One reason that Phytophthora species are widely spread plant

pathogens is that they are well adapted to aquatic environments and dispersed

by water (Blackwell 1944, Hong & Moorman 2005). If water is directly used

for irrigation in crop production areas without proper decontamination,

Phytophthora pathogens in the water can spread rapidly from field to field

through irrigation systems, severely damaging ornamental and other crops.

168 ... Yang & Hong

Previous studies have revealed the presence of at least 20 Phytophthora

species in aquatic environments (Hong & Moorman 2005), including many

notorious plant pathogens such as P capsici (Roberts et al. 2005), P. nicotianae

(Thomson & Allen 1976), and P ramorum (Werres et al. 2007). Other

Phytophthora species regarded as opportunistic plant pathogens or lacking

recorded pathogenicity also survive in the irrigation systems, such as P irrigata

(Hong et al. 2008), PR. hydrogena (Yang et al. 2014), and P. gonapodyides (Pittis

& Colhoun 1984). Many new Phytophthora taxa also exist in irrigation water

and other aquatic environments. In Virginia alone, approximately 15 known

Phytophthora species and hundreds of isolates representing unknown taxa have

been recovered from irrigation water at ornamental plant nurseries (Hong et

al. 2008). It still is not known whether these novel Phytophthora species pose a

threat to nursery production (Hong et al. 2008, Yang et al. 2014). These species

may also be confused with major plant pathogenic species. Hence, describing

these new species is important for reducing the risk of misidentifying high-

impact Phytophthora species and minimizing the threat that Phytophthora

species pose to crops and natural plants (Gallegly & Hong 2008).

In this study we formally name a novel species, Phytophthora virginiana.

Isolates were recovered during statewide surveys in Virginia, USA. We provide

detailed morphological, physiological, and phylogenetic evidence to separate

P. virginiana from all known Phytophthora species.

Materials & methods

Cultures

Isolates of Phytophthora virginiana were recovered from runoff containment basins

and irrigation reservoirs at ornamental plant nurseries in several Virginia counties.

The five representative isolates examined in this study were obtained during 2006-7

by baiting with rhododendron or camellia leaves (Bush et al. 2003). Agar blocks with

actively growing cultures in 20% clarified V8 juice agar (CV8A) were transferred

into microtubes with sterile distilled water for long-term storage at both 15 and

-80°C. Sequences from all isolates were analyzed. Isolates 44G6 and 46A2 were also

morphologically examined and tested for cardinal temperatures.

Sequencing and phylogeny

DNA extraction followed Hong et al. (2008). PCR amplifications of the internal

transcribed spacer (ITS) region covering ITS1, 5.88 rRNA gene, and ITS2 were

performed using primer pair ITS6F4R (Cooke et al. 2000). The reaction mix recipe and

PCR program were applied according to Kong et al. (2003). Sequencing was done at

the University of Kentucky Advanced Genetic Technologies Center (Lexington, KY).

Sequences of both directions were visualized with Finch TV v. 1.4.0., aligned using

Clustal W in MEGAS (Tamura et al. 2011), and edited manually to correct obvious

errors. Sequences of P virginiana isolates were aligned with those of other clade 9

species and representative Phytophthora species from other clades using Clustal W in

Phytophthora virginiana sp. nov. from irrigation water (USA) ... 169

MEGAS (Tamura et al. 2011). Pythium aphanidermatum was used as an outgroup. The

phylogeny was reconstructed in MEGAS using the Maximum Likelihood method based

on the Tamura-Nei model (Tamura & Nei 1993) with 1000 bootstrap replicates.

Cardinal temperatures

Isolates 44G6 and 46A2 were subcultured on CV8A and carrot agar (CA) and

incubated at 5, 10, 15, 20, 25, 30, 35, and 40°C. Agar blocks (5 mm diam.) were taken

from 7-day-old cultures using a flamed cork borer and placed upside down at the

center of Petri dishes. For each isolate, triplicate dishes per medium were placed in an

incubator at a designated temperature. Two colony growth measurements per colony

were taken after 4 days when the fastest growing colony was c. 1 cm from the dish

edge. This test for cardinal temperatures was repeated. Means (with standard errors) of

daily radial growth were plotted against temperature using the gplots package v. 2.11.0

(Warnes et al. 2012) in R statistical software v. 2.15.0 (R Core Team 2012). Analysis of

variance was conducted in R to determine whether the measurements were statistically

different between two isolates and experiments.

Morphology

Colony morphology of P. virginiana was noted and photographed from 10-day-old

cultures on CV8A, CA, malt extract agar (MEA), and potato dextrose agar (PDA) at

20°C in the dark.

Sporangia were produced by incubating agar blocks with actively growing cultures (<

2 weeks) in non-sterile 1.5% soil water extract solution (SWE, 15 g of non-Phytophthora

containing nature soil/1 L distilled water) for 7-8 hours under cool-white fluorescent

lamps at room temperature (c. 23°C). Mature chlamydospores were observed directly

on 20-day-old cultures. Sporangia and chlamydospores were photographed using a

Nikon Fujix Digital Camera HC-300Zi connected to a Nikon Labophot-2 microscope at

1000x. Both length and width of 50 randomly selected mature sporangia of P. virginiana

and two perpendicular diameters of 30 chlamydospores were measured using Image-

Pro® Plus v. 5.1.2.53.

Mating type of P virginiana isolates was determined by placing each isolate with

an A' or A* mating type tester of P cinnamomi in dual cultures on hemp seed agar

(HSA). The polycarbonate membrane method (Ko 1978, Gallegly & Hong 2008) was

used to induce selfed gametangia of P. virginiana in the presence of its opposite mating

type tester at room temperature. Several heterothallic species including P. cinnamomi,

P. cambivora, P. meadii, P. nicotianae, and P. cryptogea were used as mating type testers.

When no sexual organ was found after being paired for up to three months at room

temperature in dark, additional polycarbonate membrane tests were conducted at 20

and 25°C.

Results

Sequencing and phylogeny

All P. virginiana isolates produced identical ITS sequences of 745 bp (ITS1 =

180 bp, 5.8s gene = 127 bp, ITS2 = 438 bp) that differed from those of all tested

Phytophthora species. Phytophthora virginiana differed by 26 bp (11 bp in ITS1,

170 ... Yang & Hong

15 bp in ITS2) from its closest relative, the type isolate 5D1 of P. hydropathica

(GenBank EU583793).

The ITS maximum likelihood phylogenetic trees (Fic. 1) placed P virginiana

in clade 9 of Phytophthora (Blair et al. 2008). Phytophthora virginiana isolates

were supported as a taxon unique from all other known species by a strong

bootstrap (Fic. 1). GenBank accession numbers generated in this study and

used for phylogeny construction are included in Fie. 1.

Phytophthora virginiana 43G2 KC955179

Phytophthora virginiana 44G6 KC295543

® | Phytophthora virginiana 43D6 KC295542

4g | | Phytophthora virginiana 40A9 KC955178

Phytophthora virginiana 46A2 KC295544

30 High-

ms P. parsiana 47C3 KC733446 temperature

P. hydropathica 5D1 EUS83793 tolerant

species

P. hydrogena 46A3 KC249959

P. irrigata 2337 EU334634

P. chrysanthemi GF749 AB437135

P. aquimorbida 40A6 FJ666127

P. insolita PMCS-1 GU111612

P. polonica VASWS0198 DQ396410

P. macrochlamydospora 33E1 KC733445

P. quininea CBS40648 DQ275189

P. constricta CBS125801 HQ013225

P. captiosa 310C DQ297402

100 * P. fallax 310L DQ297391

P. sojae 28F9 KC733444

P. lateralis {M1040503 AF266804

P. gonapodyides CBS55467 HQ643236 Clade 1-8

P. infestans 27A8 KC733443

72 P. meadii CBS21988 HQ643268

Py. aphanidermatum P1779 GU983641 | Pythium

Clade 9

-———— 4

0.05

FiGuRE 1. Maximum likelihood phylogenetic tree for species in Phytophthora clade 9 including

Phytophthora virginiana and reference species in other clades based on ITS sequences. Alignment

was conducted with Clustal W and the phylogenetic tree was generated in MEGA 5.

Cardinal temperatures

Measurements of radial growth were statistically identical between two

P. virginiana isolates (P = 0.21) and two experiments (P = 0.25). Temperature-

growth relation of P. virginiana isolates is shown in Fic. 2. Neither isolate grew

on tested media at 5°C. The optimum temperature was 30°C on both CV8A

and CA. Both isolates grew well at 35°C with 46A2 growing >5 mm daily on

both media and 44G6 growing ~ 5 mm daily on CV8A. However, 44G6 grew

relatively slowly on CA with an average growth rate of 1.6 mm.d™ in both

experiments. No growth of 44G6 was observed on CA at 40°C, whereas 46A2

had notable growth on both media, as did 44G6 on CV8A at 40°C.

Phytophthora virginiana sp. nov. from irrigation water (USA) ... 171

44G6 CA

44G6 CV8A

46A2 CA

46A2 CV8A

Daily radial growth rate (mm)

0 5 10 15 20 25 30 35 40 45

FIGURE 2. Phytophthora virginiana. Average daily radial growth of on carrot agar (CA) and clarified

V8 juice agar (CV8A).

Colony morphology

After growing in the dark at 20°C for 10 days, the colonies of the two

P. virginiana isolates were photographed (Fic. 3). The growth patterns of

both isolates were similar on each tested medium. On CA, they grew fast and

formed radiate colonies with relatively smooth edges, producing abundant

aerial mycelia at the center and scattered at the colony intermediate and

edge. On CV8A, they formed similar chrysanthemum colony patterns with

cotton-like aerial mycelia at the colony center. On MEA, both formed tufted

to chrysanthemum colonies with discontinuous edges. Colonies on PDA had

dense aerial mycelia and expanded less than on CV8A.

Taxonomy

Phytophthora virginiana Xiao Yang & C.X. Hong, sp. nov. FIG. 4

MycoBank MB 804533

Differs from Phytophthora hydropathica by abundant, thin-walled, lateral

chlamydospores.

Type: USA, Virginia, baited with camellia leaves from irrigation water of an irrigation

runoff reservoir in a production perennial nursery, October 2007, collected by Chuanxue

Hong. Holotype, ATCC MYA-4927; ex-type culture, 46A2 (GenBank KC295544).

ETyMoLoey: ‘virginiana’ refers to the Commonwealth of Virginia, where the new

species was initially recovered.

172 ... Yang & Hong

FIGURE 3. Colony morphology of Phytophthora virginiana isolates 44G6 (top) and 46A2 (bottom)

after 10-days growth at 20°C on (left to right) carrot agar, clarified V8 juice agar, malt extract agar

and potato dextrose agar.

SPORANGIA produced occasionally by aged cultures grown on carrot agar and

20% clarified V8 juice agar, and abundantly by culture plugs submerged in

1.5% soil water extract under light within 10 hours. Sporangial shape mostly

ovoid to obpyriform and sometimes limoniform to ellipsoid with distorted-

shaped sporangia such as peapod-shaped and peanut-shaped on culture plugs

submerged in SWE after 20 hours. Sporangia terminal, nonpapillate and

noncaducous; av. 51.7 x 32.5 um. Internal proliferation of sporangiophore

common, nested or extended. SPORANGIOPHORE erect, unbranched with

occasional swelling. HyPHAL SWELLINGS common, often peanut-shaped,

angular, or variously distorted in shapes. CHLAMyDOsPORES abundant,

thin-walled, spherical and lateral produced by mature cultures av. 43.5 um

diam, sometimes on short stalks, with tapered base or clustered. Terminal

chlamydospores rare.

Phytophthora virginiana is self-sterile, producing no sexual organ in single

cultures. In the polycarbonate membrane tests, P. virginiana isolates belong to

silent A' mating type, producing no sexual organ after up to 90-day-pairing,

but stimulating A’ mating type isolates of P cinnamomi and P. cryptogea to

produce gametangia after 30-day-pairing at 20, 25°C and room temperature

(ex23°@),

ADDITIONAL ISOLATES EXAMINED: USA, VIRGINIA, baited with rhododendron or

camellia leaves from irrigation runoff reservoirs in several perennial nurseries: June

2006, 40A9 (GenBank KC955178); November 2006, 43D6 (GenBank KC295542); May

2007, 43G2 (GenBank KC955179); May 2007, 44G6 (GenBank KC295543).

Phytophthora virginiana sp. nov. from irrigation water (USA) ... 173

FiGuRE 4. Phytophthora virginiana. Morphology of asexual structures. A-G: Sporangia in

various shapes; A; Ovoid sporangium; B: Ovoid sporangium appearing semi-papillate just before

releasing zoospores; C: Limoniform sporangium; D: Ellipsoid sporangium; E: Obpyriform

sporangium; F: Distorted, peapod-shaped sporangium; G: Distorted, peanut-shaped sporangia.

H: Extended proliferation. I-K: Nested proliferation with intercalary swelling on sporangiophore

in K. L: Sporangium releasing zoospores with an incipient proliferation. M-N: Distorted hyphal

swellings. O: Intercalary swelling. P: Lateral chlamydospore and hyphal swelling. Q-S: Lateral

chlamydospores; S: Lateral chlamydospore with a tapered base on a short stalk. T: Terminal,

aborted chlamydospore. Bars = 10 um.

Discussion

Phytophthora virginiana was recovered from nursery irrigation water. This

new species has unique morphological and physiological attributes as well as

molecular signature, so that it can be readily distinguished from all known

Phytophthora species. It is the only species that is heterothallic, grows well at

35°C, and produces abundant lateral chlamydospores. Phytophthora insolita

(Ann & Ko 1980), which also grows well at 35°C and produces abundant thin-

walled chlamydospores, is homothallic and produces terminal chlamydospores.

Phytophthora lateralis (Tucker & Milbrath 1942) and P quininea (Crandall 1947)

also produce abundant lateral chlamydospores but are homothallic and did not

sustain growth at 35°C (Tucker & Milbrath 1942, Crandall 1947). Phytophthora

quininea also produces intercalary chlamydospores (Crandall 1947) whereas

174 ... Yang & Hong

P. virginiana does not. Among the known heterothallic, high-temperature

tolerant species that grow well at 35°C, only P. virginiana produces abundant

lateral chlamydospores. Phytophthora melonis, P. hydrogena, and P. irrigata do

not produce chlamydospores in fresh agar media (Ho et al. 2007, Hong et al.

2008, Yang et al. 2014). Some isolates of P drechsleri were reported to produce

much smaller chlamydospores (diam. 7.9 um) (Erwin & Ribeiro 1996) than

those of P. virginiana (diam. 43.5 um). Phytophthora hydropathica and some

P. parsiana isolates frequently produce chlamydospores in fresh cultures (Hong

etal. 2010, Mostowfizadeh-Ghalamfarsa et al. 2008). However, these two species

produce mostly terminal chlamydospores, which are rare in P. virginiana. The

average chlamydospore sizes of P hydropathica (diam. 37 um) and P. parsiana

(diam. 27-37.5 um) are also smaller than that of P. virginiana. Additionally,

the ITS sequence of P. virginiana differs from its two closely related species

by 26 bp (BP hydropathica) and 39 bp (P. parsiana). This sequence difference is

another important criterion for separating P. virginiana from all other existing

Phytophthora species.

The phylogenetic placement of P virginiana provides further evidence

to the argument that there is a high-temperature tolerant cluster (Fic. 1) in

clade 9 of the genus Phytophthora (Yang et al. 2014). Phytophthora virginiana

and all other species within this unique cluster grow well at 35°C and some

survive at 40°C (Fic. 2). To date, species belonging to this cluster have been

found on several continents including Europe, North America, South America,

and Asia. Phytophthora polonica was first described in Poland (Belbahri et al.

2006). Irrigation reservoirs in Virginia contain diverse species belonging to this

cluster including P. aquimorbida (Hong et al. 2012), P hydropathica (Hong et

al. 2010), P. hydrogena (Yang et al. 2014), P irrigata (Hong et al. 2008), and this

new species P. virginiana. Phytophthora chrysanthemi, P. insolita and P. parsiana

were first reported from Asia (Ann & Ko 1980, Mostowfizadeh-Ghalamfarsa

et al. 2008, Naher et al. 2011). Also, a provisional species belonging to this

cluster, Phytophthora sp. “lagoariana,” recovered from the Amazonian

rainforest, also grows well at 35°C (data not shown). The unique correlation

between physiological and phylogenetic characteristics indicates that these

high-temperature tolerant species may originate from tropical or subtropical

environments, and increasing international trade may have contributed to

current global distribution of these species.

The ecological and economic impacts caused by P. virginiana are not clear at

this time. All isolates were recovered from irrigation water at several ornamental

plant nurseries in Virginia by baiting. However, plants diseased by P virginiana

have not been observed at the same nurseries. This does not necessarily

mean that P virginiana will not cause severe damage when introduced to

Phytophthora virginiana sp. nov. from irrigation water (USA) ... 175

new environments. Investigations into its host range and ecological roles in

irrigation systems are warranted.

Acknowledgments

This research was supported in part by grants from USDA/NIFA (2005-51101-

02337, 2010-51181-21140). We thank Mrs. Patricia Richardson for her assistance with

submission of PCR products for DNA sequencing during the study. We thank Drs. Zia

Banihashemi, Ross Beever, Treena Burgess, Michael Coffey, David Cooke, Kenneth

Deahl, Erhard Halmschlager, and Brett Tyler for providing isolates of reference species

used in the phylogenetic analysis in this study. We thank Dr. Steven Jeffers for helpful

discussion about the terminology in the genus Phytophthora. Our thanks extend to Mr.

Xitong Li for his help with processing and editing figures.

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MY COTAXON

http://dx.doi.org/10.5248/126.177

Volume 126, pp. 177-182 October-December 2013

New records of Puccinia on Poaceae

from Khyber Pakhtunkhwa, Pakistan

A. IsHAQ’*, N.S. AFSHAN? & A.N. KHALID’

™Department of Botany & *Centre for Undergraduate Studies, University of the Punjab,

Quaid-e-Azam Campus, Lahore, 54590, Pakistan

*CORRESPONDENCE TO: aamna_ishaq@yahoo.com

AsBstRAcT —Puccinia arthraxonis-ciliaris and P. pseudocesatii were collected from Khyber

Pakhtunkhwa and are new records for Pakistan.

Key worps — Naran, Pucciniales, rust fungi

Introduction

Khyber Pakhtunkhwa (KPK), formerly known as the Northwest Frontier

Province (NWFP), is one of the five provinces of Pakistan, located in the

northwest of the country. It is situated at approximately 34.00°N 71.32°E

(Fic. 1). Khyber Pakhtunkhwa is famous for hill coniferous forests, herbal

plants, and large biodiversity. This floristically rich area is affected by many

biotic and abiotic stresses including plant diseases such as leaf blights, red rot,

smut, mosaic, and rust (Asad et al. 2007). One hundred seventy taxa of rust

fungi have been reported from this area (Afshan & Khalid 2008, 2009; Afshan

et al. 2007, 2008a,b,c,d, 2010; Khalid & Afshan 2009; Ahmad et al. 1997; Iqbal

et al. 2008). The number of rust species recorded from this area is expected to

increase in future due to extensive exploration.

We describe below three new rusts from Khyber Pakhtunkhwa that bring

the total rust taxa recorded from the area to 173: Puccinia pseudocesatii on

Dichanthium annulatum and P. arthraxonis-ciliaris on Arthraxon sp. (both

new records for Pakistan) and P helictotrichi var. pakistanica on Helictotrichon

junghuhnii.

Materials & methods

During an exploration of the rust diversity in KPK, rust infected plants and their

inflorescences were collected from different areas of the province and compared

morphological with earlier herbarium collections. Freehand sections & scrape mounts

178 ... Ishaq, Afshan & Khalid

70° 72° 74 76°

North i

Northern Areas

36° ——e eee eae 36°

CT lady os 78°

Afghanistan od \

34° —-—- L. — Péshawa ir 5 ee Reena.

Islamabad

Provisional Boundaries ———————

District Boundaries

Sampling sites * *&¥

Federally Administrated eS

Tribal Areas

60 Kilometers

FiGuRE 1: Map of Khyber-Pakhtunkhwa showing sampling sites.

of infected plant materials were made in lactic acid. The plants were photographed

and infected portions were observed under stereomicroscope. Twenty spores of each

spore state were examined under microscope (Nikon YS 100) and paraphyses and spore

dimensions were taken by an Ocular micrometer (Zeiss Eyepiece Screw Micrometer).

Sections, paraphyses and spores were microphotographed by digiporo-Labomed.

Illustrations of spores and paraphyses were made under Lucida camera (Ernst Leitz

Wetzlar Germany).

New records

Puccinia arthraxonis-ciliaris Cummins, Uredineana 4: 16 (1953) Fig. 2

SPERMOGONIA, AECIA & UREDINIA not found. TELIA amphigenous, black,

scattered. TELIOSPORES ellipsoid to oblong-ellipsoid, light brown, 1-2 celled,

mostly 2-celled spores (19-)21-28 x 42-49(-57) um; apex 6-10 um thick; wall

2-4 um thick; pedicel hyaline, 5-9 x (23-)29-54 um. 1-celled spores 21-26

(-31) x (26-)34-38 tm; apex 7-10 um thick; wall 2-3 um thick; pedicel up to

44 um long.

MATERIAL EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA, Ghora gali, 1691 ma. s.

l., on Arthraxon sp., stage III, 30 April 2008. A. Ishaq AM69 (LAH 1166).

ComMENTs: Puccinia arthraxonis-ciliaris has previously been reported on

Arthraxon hispidus (Thunb.) Makino from China, India, Japan, Mauritius,

Puccinia taxa new to Pakistan ... 179

FIGURE 2: Puccinia arthraxonis-ciliaris. Teliospores. Scale bar = 14 um.

New Guinea, and Philippines (Cummins 1971 and represents a new species

record for Pakistan. Puccinia arthraxonis (Henn.) Syd. et al. has previously been

reported from Pakistan on different Arthraxon species (Ahmad 1976; Khalid et

al. 1993).

Puccinia pseudocesatii Cummins, Uredineana 4: 71 (1953) Fiess

SPERMOGONIA, AECIA & UREDINIA not found. TELIA amphigenous, pulvinate,

black. TEL1osporEs ellipsoid, hyaline to light brown, 2-celled, 3-celled rarely,

10-20 x (35-)44-60 um, sometimes upper cell is wider than lower one,

10-27 um wide; apex 5-15(-18) um thick; wall 1.5-2 um thick; pedicel hyaline,

fragile, 5-8 x 23-67 um.

MATERIAL EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWaA, Ghora Gali, 2009 ma.s.L.,

on Dichanthium annulatum (Forssk.) Stapf, stage III, 29 March 2009, A. Ishaq AM31

(LAH 1172)

ComMENTs: Puccinia pseudocesatiihas previously been reported on Bothriochloa

ischaemum (L.) Keng and Chrysopogon gryllus (L.) Trin. from southern Europe

180 ... Ishaq, Afshan & Khalid

FIGURE 3: Puccinia pseudocesatii. Teliospores. Scale bar = 17 um.

(Cummins 1971). From Pakistan, P. cesatii J. Schrot., P duthiei Ellis & Tracy,

P. dichanthii Afshan & Khalid, Uromyces andropogonis-annulati Syd. et al. and

U. clignyi Pat. & Har. have previously been reported on Dichanthium annulatum

(Ahmad 1956a,b; Hasnain et al. 1959; Ghaffar and Kafi 1968; Afshan et al.

2010).

This is a new record for Pakistan and Dichanthium annulatum represents a

new host for this rust fungus.

Puccinia helictotrichi var. pakistanica Afshan & Khalid, Mycotaxon 112: 487

(2010) Fic. 4

SPERMOGONIA and AECIA unknown. UREDINIA scattered on whole leaf

surface, naked, amphigenous, golden brown. UReEpiniospores globose

to subglobose, hyaline, 15-24 x 20-26 um; wall 1.5-3 um thick, hyaline,

echinulate; germ pore obscure. TELIA amphigenous, golden brown, loculate,

100-200 x 40-80 mm. TELIOsporEs elongated to cylindrical, 14-24 x 36-49

um; apex 2-3.7 um thick; wall 1.5-3 um thick; pedicel hyaline, 3-7 x 3.3-13

uum.

Puccinia taxa new to Pakistan ... 181

FiGurE 4: Puccinia helictotrichi var. pakistanica.

A. Urediniospores showing echinulate surface ornamentation. B. Teliospores.

Scale bars: A = 9 um; B = 10 um.

MATERIAL EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA, Shogran, 2362 m a.s.l.,

on Helictotrichon junghuhnii (Buse) Henrard (= H. asperum (Munro ex Thwaites) Bor),

stages II & III, 28 August 2010, A. Ishaq AM04 (LAH 1145); Naran, 2427-3200 ma.s.L.,

on leaves of Helictotrichon sp., stages II & II, 29 August 2008, A. Ishaq AM15 (LAH

1149).

ComMENts: Puccinia helictotrichi var. pakistanica has been previously reported

on Helictotrichon junghuhnii (= H. virescens (Nees ex Steud.) Henrard) from

Fairy Meadows, Northern Areas (Afshan et al. 2010). This represents a new

record for Khyber Pakhtunkhwa.

Acknowledgements

We sincerely thank Dr. Abdul Rehman Niazi (Department of Botany, University

of the Punjab, Lahore, Pakistan) and Dr. Omar Paino Perdomo (Dominican Society

of Mycology, Santo Domingo, Dominican Republic) for their valuable suggestions to

improve the manuscript and acting as presubmission reviewers.

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Afshan NS, Khalid AN. 2009. New records of Puccinia and Pucciniastrum from Pakistan.

Mycotaxon 108: 137-146. http://dx.doi.org/10.5248/108.137

Afshan NS, Khalid AN, Abbasi M, Niazi AR. 2007. New records of rust fungi from Pakistan.

Mycotaxon 101: 233-237,

Afshan NS, Khalid AN, Niazi AR. 2008a. New records and distribution of rust fungi from Pakistan.

Mycotaxon 105: 257-267.

Afshan NS, Khalid AN, Niazi AR. 2008b. New records of graminicolous rust fungi from Pakistan.

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182 ... Ishaq, Afshan & Khalid

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40(3): 1285-1289.

Afshan NS, Berndt R, Khalid AN, Niazi AR. 2008d. New graminicolous rust fungi from Pakistan.

Mycotaxon 104: 123-130.

Afshan NS, Niazi AR, Khalid AN. 2010. Three new species of rust fungi from Pakistan. Mycol.

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Ahmad S. 1956a. Uredinales of Pakistan. Biologia 2: 29-101.

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Asad S, Iftikhar S, Munir A, Ahmad I, Ayub N. 2007. Pathogenic diversity in Bipolaris sorokiniana

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Bahri B, Shah S, Hussain S, Leconte M, Enjalbert J, Pope CDV. 2011. Genetic diversity of the wheat

yellow rust population in Pakistan and its relationship with host resistance. Plant Pathology 60:

649-660. http://dx.doi.org/10.1111/j.1365-3059.2010.02420.x

Cummins GB. The rust fungi of cereals, grasses and bamboos, Springer-Verlag, New York; 1971.

http://dx.doi.org/10.1007/978-3-642-88451-1

Fazal H, Nisar A, Rashid A, Farooq S. 2010. A checklist of phanerogamic flora of Haripur Hazara,

Khyber Pakhtunkhwa, Pakistan. Pak. J. Bot. 42(3): 1511-1522.

Ghaffar A, Kafi A. 1968. Fungi of Karachi. Pak. J. Bot. 4: 195-208.

Hasnain SZ, Khan A, Zaidi AJ. 1959. Rusts and smuts of Karachi. Bot. Dept. Karachi Univ. Mont.

Iqbal SH, Khalid AN, Afshan NS, Niazi AR. 2008. Rust fungi on Saccharum species from Pakistan.

Mycotaxon 106: 219-226.

Khalid AN, Afshan NS. 2009. Addition to the graminicolous rust fungi of Pakistan. Mycotaxon

108: 175-183. http://dx.doi.org/10.5248/108.175

Khalid AN, Iqbal SH, Ahmad F. 1993. Rust flora of Pakistan. Uredinales collected in Salt Range,

Pakistan. Sci. Int. (Lahore) 5(2): 211-214.

MY COTAXON

http://dx.doi.org/10.5248/126.183

Volume 126, pp. 183-190 October-December 2013

The olive goblet: Peziza oliviae,

a new cup fungus growing underwater in Oregon

JONATHAN L. FRANK

Department of Biology, Southern Oregon University,

1250 Siskiyou Blvd., Ashland, OR 97520-5071, USA

CORRESPONDENCE TO: frankjon@sou.edu

ABSTRACT — A new species, Peziza oliviae, is described from stream habitats in the Oregon

Cascade Mountains (USA) based on morphology, habitat, and ITS and LSU nrDNA sequence

data. This new member of the Ascomycota grows underwater on sticks and decomposing

wood as well as near running water where the woody substrate is saturated.

Key worps — Pezizaceae, underwater, monitoring, survey, manage

Introduction

Surveys for rare and endangered fungi in Oregon continue to be part of

management practices, which now include stream habitats in search protocols

for rare macrofungi that fruit underwater, such as Vibrissea truncorum

(Ascomycota) and Psathyrella aquatica (Basidiomycota) (Frank et al. 2010;

Oregon Biodiversity 2010). While only one gilled mushroom, P. aquatica, from

the Rogue River in southern Oregon, is known to grow and fruit underwater,

several ascomycetes fruit when submerged. Cudoniella clavus forms tan to gray

fruiting bodies on submerged sticks, while Vibrissea truncorum and Mitrula

spp. form orange caps on submerged sticks as well as on wet woody debris

near water. Sessile macrofungi (e.g., Vibrissea filisporia, Scutellinia scutellata,

Pachyella clypeata, Adelphella babingtonii) fruit underwater in streams in

Oregon, and the new genus Aquapeziza (Pezizaceae) was recently described

for a cup fungus fruiting underwater in China (Pfister 1973; Wang et al. 2005;

Pfister et al. 2008; Hu et al. 2012). Aquatic microfungi are also classified in

the Oomycota, Chytridiomycota, and aquatic hyphomycetes as well as new

genera in the Leotiomycetes and Sordariomycetes from Florida and Costa

Rica (Raja et al. 2009; Ferrer et al. 2012). Documentation of new and unusual

fungal biodiversity provides target descriptions and DNA barcodes useful for

monitoring and conservation (Halme et al. 2012).

184 ... Frank

In the broad sense, Peziza historically encompassed a variety of ascomal

morphologies, including the archetypal sessile cup, stalked goblets, and

convoluted truffle-like sequestrate fungi. Molecular data indicate that Peziza

s.l. is polyphyletic and should be separated into several distinct genera, with

the core group (Peziza s.s.) surrounding the generic type, P. vesiculosa. Fungi

in the core group are saprotrophic and characterized by an amyloid ring at the

tips of the asci, which form a palisade with thinner paraphyses along a more or

less cup-shaped hymenium. They are represented by ellipsoid spores that may

be smooth or ornamented; a stalk may be present but is usually not prominent,

forming just a point of attachment to the substrate (Dennis 1968; Korf 1973;

Breitenbach & Kranzlin 1984; Tylutki 1979; Hansen et al. 2001, 2002).

During surveys for rare species in central Oregon, I repeatedly collected

an olive-brown stalked cup fungus growing underwater in small streams in

the Willamette National Forest. Previously, no species of Peziza s.s. have been

reported growing underwater in Oregon. Aquatic fruiting is such an unusual

phenomenon that mycologists apply special scrutiny to confirm that a fungus

has not originally fruited above water and subsequently fallen in or become

submerged in rising waters. The fungus, which does not match any previously

described species on survey lists or in scientific literature, is described here as

Peziza oliviae.

Materials & methods

Streams, riverbanks, and saturated woody substrates were surveyed for macrofungi

during May—November 2011 and May—August 2012 in the Willamette and Mount Hood

National Forests. Site locations (UTM NAD83 Zone 10) were determined and recorded

using a Garmin 450 portable GPS unit. Collections were mapped, described in the field.

Photographs of fresh material were taken with a Canon T2i digital camera (field) or

under a Leica MZ75 dissecting microscope (laboratory). Fresh hand sections mounted

in distilled water and Melzer’s reagent were viewed under a Leica DMLB compound

microscope. Thirty spores from each of eight collections were measured at 1000x. All

microscopic measurements were made from sections mounted in distilled water.

DNA was extracted, amplified, and sequenced following Frank et al. (2010): fresh

fungal tissue was ground with micropestles in CTAB buffer, extracted with chloroform,

and amplified with PCR primers ITS1f, ITS4, and TW13. Internal primers ITS1 and

ITS4r were used for sequencing reactions. After sequences were edited with Chromas

1.45 (McCarthy 1998), contiguous sequences were assembled in Sequencher v4.7 (Gene

Codes Corp. Ann Arbor, MI) and compared to other fungal ITS and LSU sequences in

GenBank with BLAST (Altschul et al. 1990). MAFFT and ClustalX were used to generate

multiple sequence alignments (Katoh et al. 2002; Thompson et al. 1997). Alignments

were edited manually using BioEdit and Mesquite (Hall 1999; Maddison & Maddison

2011). Phylogenetic trees, using maximum likelihood with 1000 bootstrap replicates,

were generated using fastDNAml (Felsenstein 1981; Olsen et al. 1994) through the

Mobyle portal of the Pasteur Institute (http:/mobyle.pasteur.fr). ML analyses using

Peziza oliviae sp. nov. (U.S.A.) ... 185

settings that correspond to the HKY85 model, and parsimony analyses with 1000

bootstrap replicates and 1000 jack-knife replicates, were generated using PAUP*

4.10b10 (Swofford 2002). Consensus trees with 50% majority-rule were generated using

the tree-bisection-reconnection branch-swapping algorithm. All characters were given

equal weight; gaps were treated as missing.

Of the 41 aligned ITS sequences, five were generated from collections of Peziza

oliviae, one was generated from Peziza sp. collected in southern Oregon, and 35

were obtained from GenBank. ITS and LSU rDNA sequences have been deposited in

GenBank; ITS alignment and trees were submitted to TreeBase.org.

Results

Ten collections of P. oliviae were made during July-October 2011 and June-

August 2012. Ascocarps were found growing from stream bottoms submerged

10-25 cm in running water or on saturated wood near running water in first

order streams in the Santiam and Breitenbush watersheds at 900-1400 m

elevations.

ITS nrDNA was amplified and sequenced for five collections of P. oliviae and

for one morphologically similar Peziza sp. collected underwater in southern

Oregon (JLF2167). LSU nrDNA was amplified and sequenced for three

collections of P. oliviae and for Peziza sp. (JLF2167). Sequence data have been

deposited in GenBank (JX415339-JX415342 and KC916727-KC916729).

Edited alignments of 41 ITS sequences generated consensus trees using

parsimony with 774 total, 256 constant, 117 variable, and 401 parsimony-

informative characters. Maximum Likelihood and Maximum Parsimony

analyses support the molecular uniqueness of this new species, with P. oliviae

nearest to the European P lohjaoensis (PLATE 1). Maximum Parsimony analysis

and pairwise comparisons of LSU sequence data confirm the placement of

P. oliviae in Peziza s.s. Alignments and trees are available at http://purl.org/

phylo/treebase/phylows/study/TB2:S14001.

Morphological and molecular analyses support the designation of this

unusual stalked cup as a new species. This cup fungus grows underwater on

sticks and wood embedded in the sediments, pebbles, and cobble of mountain

stream bottoms and above water on saturated wood near cold mountain

streams in the Oregon Cascades. It has a prominent stalk, olive-brown color,

operculate asci with amyloid tips, smooth nonguttulate ellipsoid spores, and

ITS and LSU nrDNA sequences that place it in Peziza s.s. (PLATE 1).

Taxonomy

Peziza oliviae J.L. Frank, sp. nov. PL. 2

MycoBank MB803430

Differs from the genetically related Peziza lohjaoensis by its smooth spores and North

American distribution and from other Peziza spp. by its submerged habitat.

186 ... Frank

AF491576 Peziza lohjacensis

JLF2088 Peziza oliviae

100) | sir2001 Peziza oliviae TYPE

JLF2479 Peziza oliviae

95 JLF2474 Peziza oliviae

JLF2118 Peziza oliviae

AF491500 Peziza ampliata

JF08636 Peziza ninguis

65 AF491614 Peziza domiciliana

85 JFe0ss29 Peziza perdicina

JF008849 Peziza udicola 2

AF491508 Peziza fimeti 7

AF491620 Peziza nivalis 2

AF401884 Peziza arvernensis §

60 “ AF491575 Peziza echinospora 5

00 FJ235142 Peziza cf. echinospora | &

FJ236144 Peziza cf. echinospora

94 AF491621 Peziza ammophila

JFe08558 Peziza granularis

O00 | JFe08sé6s Peziza vesiculosa

AF491623 Peziza vesiculosa

JFO08S55 Peziza bovina

JLF2167 Peziza sp.

AF491612 Peziza alcis

100 Jreosses Peziza proteana

JFeoss27 Peziza petersii

JP908534 Peziza moseri

AF491628 Peziza subcitrina

100 AY830858 Hydnotryopsis setchellii

80 pazoes43 Sarcosphaera coronaria

AY818334 Peziza ostracoderma

AY830852 Cazia flexiascus

86 100 FJ268642 Peziza stuntzii

AY830851 Peziza ellipsospora

JF808544 Peziza badia

AF276869 Tirmania pinoyi

JFQ08570 Peziza polaripapulata

100 JF90sss6 Peziza succosa

AY830853 Peziza infossa

Ay920528 Pachyphloeus sp.

100

PLATE 1. Maximum Likelihood tree using ITS nrDNA showing the placement of Peziza oliviae in

Peziza s.s. 23,336 rearrangements tried, score of best tree = 9722.72452. MP Bootstrap numbers

greater than 50% are included above branches; analyses performed in PAUP* using 1000 bootstrap

replicates.

TYPE: United States, Oregon, Marion Co., Willamette NE, Dunlap Creek, UTM E571615

N4962062, elevation 990 m, underwater in stream, 15 Jul 2011, JLF2091 (HOLOTYPE-

OSC148300; GenBank JX415340).

Erymo_oey: for Doris Olivia for help finding this cryptic cup fungus; also reflecting the

olive hue of the ascomata.

ASCOMATA Olive to golden-brown stalked cups 7-25 mm tall, 8-40 mm diam,

margin crenulate. Stipe 1-10 x 3-7 mm, external surface roughened, glabrous.

Ascomal morphology varies from stalked cup or goblet-shape with upturned

margin to a nearly sessile slightly convex disc with a short but prominent stalk.

SUBHYMENIUM 15-25 um thick, composed of small isodiametric cells 4-10

um. MEDULLARY EXCIPULUM 500-1800 um thick, composed of large inflated

Peziza oliviae sp. nov. (U.S.A.) ... 187

isodiametric cells 45-110 x 35-85 um. EcTaL ExcrpuLUM 50-150 um thick,

composed of elongate and contorted cells 20-40 x 15-35 um with olive-brown

pigmentation.

Asct 250-380 x 16-22 um, cylindrical, operculate, arranged in a regular

palisade with amyloid tips forming a distinct blue ring zone. PARAPHYSES

3-5 um diam, with enlarged clavate tips 5-8 um diam equal to or extending

slightly (3-8 um) beyond the asci, septate with septa 15-40 um apart. SPORES

nonguttulate smooth ellipsoid 19.5-24 x 9.5-12 um, 8 per ascus.

ECOLOGY & DISTRIBUTION - small streams in the Cascade Range of north

central Oregon between 800 and 1500 m elevation, associated with dead woody

debris embedded in stream bottoms or on saturated wood, at the stream surface

or on stream banks; June through October.

ADDITIONAL SPECIMENS EXAMINED —UNITED STATES. OREGON: MARION Co.,

Willamette NF: underwater in Dunlap Creek, UTM E584752 N4964113, elev. 990

m, 15 Jul 2011, JLF2087 (OSC148297); UTM E571892 N4962471, elev. 1070 m,

JLF2088 (OSC148298; GenBank JX415339); UTM E571641 N4962234, elev. 1010 m,

JLF2089 (OSC148299); UTM E577193 N4964125, elev. 1050 m, 27 Jul 2011, JLF2118

(OSC148301; GenBank JX415341); UTM E584753 N4964113, elev. 1212 m, 10 Aug

2011, JLF2140 (OSC148302); UTM E559896 N4957372, elev. 1270 m, 15 Oct 2011,

JLF2189 (OSC148303); UTM E571880 N4962471, elev. 1090 m, 27 Jun 2012, JLF2474

(OSC148304; GenBank KC916728); UTM E571935 N4962478, elev. 1120 m, 27 Jul 2012,

JLF2541 (OSC148307); below Scorpion Mtn. underwater in E. fork of Humbug Creek,

UTM E578141 N4963797, elev. 1295 m, 29 Jun 2012, JLF2479 (OSC148305; GenBank

KC916729); underwater in upper Byars Creek, UTM E568438 N4959910, elev. 1240 m,

26 Jul 2012, JLF2538 (OSC148306).

OTHER SPECIES EXAMINED: Peziza sp.: UNITED STATES. OREGON: JACKSON Co.,

Rogue River-Siskiyou NF, underwater in the Rogue River, UTM E540414 N4745695,

elev. 995 m, 7 Sep 2011, JLF2167 (OSC148308; GenBank JX415342, KC916727)

ComMENTS- Peziza oliviae is distinguished from other species in Peziza

s.s. by the combination of a stipe, smooth nonguttulate spores, olive-brown

pigmentation, its growth underwater or on saturated woody substrates, and its

western United States distribution. Among similar species, P. vesiculosa lacks a

prominent stalk and grows on dung, P domiciliana forms smaller biguttulate

spores, P. echinospora produces spiny spores, and P. ammophila grows in sand

and has a cracked margin. Its closest relative, P lohjaoensis, is separated by

a 20bp difference (>3%) in the ITS1+ITS2 region, warty spores, growth on

decaying hardwood litter, and a European distribution (Hansen et al. 2002).

Another stalked Peziza sp. (JLF2167), collected underwater in the upper Rogue

River in southern Oregon, appears to be a distinct species in Peziza s.s., but

more collections are needed to confirm its taxonomic position and status

(PLATE 1).

Peziza oliviae differs from other aquatic or semiaquatic members of the

Pezizaceae with amyloid asci as follows: Aquapeziza forms globose spores,

188 ... Frank

Pe. may 5 mae ie aa

S gene. <<

Pa EE pages F 10 um

ieee? ; LG r 1

PLATE 2. Peziza oliviae. A: ascocarp in situ underwater (JLF2189). B: ascocarp with irregular

crenulate margin (JLF2189). C: ascocarp (JLF2091). D: ascocarp cross-section (JLF2140). E: spores

in asci, amyloid in Melzer’s toward the operculate tip (JLF2091). F: spores (JLF2474). Bars: A, B = 1

cm; C = 0.5 cm; D = 1 mm; E,F = 10 um.

Peziza oliviae sp. nov. (U.S.A.) ... 189

and Pachyella and Adelphella are non-stipitate and have gelatinous tissues.

While the aquatic Cudoniella clavus is not closely related, it has many forms

including a tan stalked cup that can macroscopically resemble P oliviae, so

that surveyors might mistake this for P oliviae in the field. However, C. clavus

is easily distinguished microscopically by its smaller spores and non-amyloid

inoperculate asci.

Peziza oliviae appears to be rare and associated with undisturbed forest

habitats. The streams in which P oliviae was collected contained aquatic insect

larvae and other ascomycetes including Vibrissea truncorum, V. filisporia,

C. clavus, Scutellinia scutellata, and Adelphella babingtonii. These streams are

small and do not experience regular disturbances, such as major sediment

moving floods and scouring, as do larger stream systems. These fast-moving

small streams, found at 800-1500 m in the central Oregon Cascade Mountains,

maintain relatively consistent conditions with temperatures and moisture levels

not fluctuating nearly as much as the surrounding terrain. Ascocarps were

collected at the bottom of streams attached to woody substrates embedded

in gravel and cobble. These fungi did not fruit terrestrially and thus were not

subsequently inundated by rising waters. Some fruiting bodies were collected

from saturated old logs and log fragments from large conifers (probably

Pseudotsuga menziesii or Tsuga heterophylla) near the water. Growth on dead

wood suggests that P. oliviae, like other species in Peziza s.s. (Hansen et al.

2002), is saprotrophic.

The discovery of this unusual species documents new biodiversity in healthy

old-growth forests, underscores the importance of monitoring and maintaining

such habitats, and adds to the list of species that represent measurable units

affected by adaptive management practices. Its common name, “the olive

goblet” (based on its chalice- or grail-like appearance), should help forest

managers and surveyors find and identify this unique cup fungus.

Acknowledgements

I thank Don Pfister, Keith Egger, Nancy Smith Weber, Karen Hansen, Matthew

Smith, Scot Loring, Lorelei Norvell, Shaun Pennycook, and Darlene Southworth for

helpful comments and suggestions. DNA extraction, amplification, and sequencing

were performed (by JLF) in the Biotechnology Center at Southern Oregon University.

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MY COTAXON

http://dx.doi.org/10.5248/126.191

Volume 126, pp. 191-226 October-December 2013

One hundred fourteen years of Pluteus in Brazil:

collections studied by Hennings and Rick

NELSON MENOLLI JR.’?* & MARINA CAPELARI?

‘Instituto Federal de Educagdao, Ciéncia e Tecnologia de Sao Paulo, Campus Sao Paulo,

CCT / Biologia, Rua Pedro Vicente 625, Sado Paulo, SP 01109-010, Brazil

?Nucleo de Pesquisa em Micologia, Instituto de Botanica,

Caixa Postal 68041, Sdo Paulo, SP 04045-902, Brazil

* CORRESPONDENCE TO: menollijr@yahoo.com.br

ABSTRACT — The re-examination of Pluteus collections from Brazil studied by P.C. Hennings

and J. Rick during the early 20th century expands current Brazilian knowledge of the genus.

Searches of the bibliographical and herbarium records revealed a total of 32 Pluteus names

linked to specimens collected in Brazil and studied by Hennings and Rick. Of the ten species

represented by Brazilian types, five types could not be located in any of the consulted herbaria

and therefore must be treated as nomina dubia. None of previously published collections

listed under five other names could be located. All other collections (in BPI, FH, PACA

and SP) were studied for morphology. In all cases, European names originally attributed by

Hennings and Rick have been found to be misapplied, and re-identifications based especially

on species described from the Neotropics are suggested. Sections Pluteus and Celluloderma

are represented by most collections.

Key worps — biodiversity, Pluteaceae, taxonomy

Introduction

Pluteus Fr. (Pluteaceae, Agaricales, Basidiomycota) is a genus represented

by approximately 300 species worldwide (Kirk et al. 2008). Although Pluteus

species are easily recognizable by their free lamellae, pinkish spore print,

inamyloid basidiospores, inverse hymenophoral trama, and the absence of

annulus and volva (Singer 1986), modern molecular studies have helped

resolve taxonomic inconsistencies, establish synonymies of species described

multiple times, and enlarge the generic circumscription to include annulate

species historically classified in the allied genus, Chamaeota (W.G. Sm.) Earle

(Minnis et al. 2006, Justo et al. 201lab, Vizzini & Ercole 2011). However,

molecular tools are limited in their ability to resolve taxonomic problems in old

192 ... Menolli & Capelari

and especially poorly preserved collections. An accurate morphological study

is an alternative in these situations, although it is not always possible to retrieve

useful information from old collections in which the hymenial structures are

frequently collapsed and not revivable.

Paul Christoph Hennings was the first author to describe a Pluteus species

from Brazil, PR. scruposus (Hennings 1900) based on material collected by R.

Pilger in 1899. Hennings (1904a,b) also described P termitum Henn. from

material collected by E.H.G. Ule in 1901 and P cervinus var. griseoviridis from

material collected by A. Puttemans in 1903.

Hennings was a German mycologist who studied the fungi from Brazil

in the late 19th and early 20th centuries. He examined materials collected in

different regions of Brazil by many collaborators, including E.H.G. Ule, EA.G,J.

Moller, A.EM. Glaziou, C.A.W. Schwacke, I.D.J. Huber, C.E Baker, E.A. Goeldi,

A. Puttemans, and R. Pilger (Fidalgo 1968). Hennings was born in 1841 in

Heide (Germany) and from approximately 1885 onward devoted himself to the

study of fungi from around the world, mainly those from German colonies and

Brazil. He published more than 250 papers and brought together in Berlin one

of the best fungal collections in the world (Perkins 1909). Many collections

described by Hennings were deposited in herbarium B of the Botanical Museum

Berlin-Dahlem, which was partly destroyed by fire in a bombing raid in 1943

(Merrill 1943, Hiepko 1987). However, several of Hennings’ types were saved

because duplicates of different fungal groups (especially Fungi Imperfecti and

Uredinales) were deposited in other herbaria (Hiepko 1987, Hein 1988). Hein

(1988) cited all species described by Hennings and data about the preservation

states of these collections in B, but unfortunately no Pluteus collections survived

the fire in B.

Among Henning’s collaborators was Ernst Heinrich Georg Ule, an

important and active collector of fungi in Brazil. Ule was born in 1854 in

Halle an der Saale (Germany), and in 1883 he moved to Brazil, first living

in Santa Catarina state before moving to Rio de Janeiro, where he became a

traveling naturalist of the National Museum (Martius 1906, Fidalgo 1968). The

P. termitum specimen described by Hennings (1904a) was collected during Ule’s

expedition to the Amazonian Forest in Fortaleza, Amazonas state (19.X.1901-

18.X1.1901; Hennings 1904a, Martius 1906). Ule accumulated approximately

3361 species of fungi from the Brazilian States of Santa Catarina, Rio de Janeiro,

Minas Gerais, Goiads, and Amazonas (Martius 1906). These collections were

distributed to many mycologists, including G. Winter, FO. Pazschke, H. Rehm,

H. Sydow, P. Sydow, E. Jahn, P. Dietel, and P.C. Hennings (Fidalgo 1968).

Other collaborators in Henning’s Pluteus studies were Robert Pilger and

Arsene Puttemans. Pilger, who was born in 1876 in the Heligoland Archipelago

(Germany), in 1899 conducted expeditions to the state of Mato Grosso, where

Hennings & Rick Pluteus revisions (Brazil) ... 193

he collected the first Pluteus recorded from Brazil. Puttemans, born in Brussels

(Belgium) in 1873, moved to Brazil in 1892, where he lived in Rio de Janeiro

state for two years before moving to Sao Paulo. Puttemans, who is undoubtedly

the most important name in Brazilian phytopathology, was also an important

collector of various non-pathogenic fungi. Although many other collections sent

to Germany were destroyed in the fire in B, fortunately duplicates of almost all

of Puttemans’ collections were deposited in other herbaria, and approximately

7000 samples were preserved at UFRJ of the ‘Universidade Federal Rural do

Rio de Janeiro’ (Fidalgo 1968); additional collections of Puttemans are also

preserved in other herbaria, including the type of P. cervinus var. griseoviridis

now deposited in SP ("Instituto de Botanica").

As the study of Brazilian fungi began with foreign collectors, Johann Rick

played an extremely important role and is considered the father of Brazilian

mycology. In 1904, he started a series of mycological papers representing the

beginning of the systematic study of fungi from Brazil (Fidalgo 1962, 1968).

His collections brought together approximately 12,000-15,000 exsiccatae from

the state of Rio Grande do Sul that are now deposited in PACA of the ‘Instituto

Anchietano de Pesquisas’ (Fidalgo 1968). Rick described and recorded 21

Pluteus taxa from that state (Rick 1907, 1919, 1930, 1938, 1961), including

seven new taxa from Brazil: P cristatulus, “P. exiguus var. venosa,” P. fibrillosus,

P. leptonia, P. sensitivus, P. straminellus, and P. velatus.

Rick was born in Dornbirn (Austria) in 1869. Around 1896 he began to

study fungi and came into contact with H. Rehm, G. Bresadola, and C.G.

Lloyd. He shared many duplicates with Lloyd that formed what was most likely

the largest collection of Brazilian fungi at the time in the ‘Lloyd Herbarium

(Fidalgo 1962). These “Lloyd Herbarium’ collections are now part of the U.S.

National Fungus Collections (BPI), which also includes recent collections

and contributions from many collectors, giving BPI approximately 14,000

collections representing different groups of fungi from Brazil (Farr & Rossman

2013). After Rick arrived in Brazil in 1903, he was appointed a professor of the

Jesuit College of Sao Leopoldo, Rio Grande do Sul State, where he collected

most of his specimens (Fidalgo 1962), including what likely represents his first

Pluteus specimen (dated 1906 and later identified as P cervinus; Rick 1938,

1961). Rick continued studying Brazilian fungi until 1946, the year he died in

Brazil.

Considering that foreign mycologists made the first studies of Pluteus from

Brazil, many names proposed for European specimens were used for Brazilian

materials. The current state of knowledge of Pluteus in South America is

somewhat problematic, mainly due to the abundance of species described for

this region (Singer 1959, Horak 1964) and to misapplied identifications in

the past. Nevertheless, there has not yet been any publication correcting past

194 ... Menolli & Capelari

misapplications. With the aim of complementing the study of Pluteus from

Brazil, we present our re-examination of historical Pluteus collections studied

by Hennings and Rick and comment on the identifications. Undoubtedly,

studies involving a combination of multiple methods and different types

of evidence are relevant for insights into the evolution of the genus and to

determine whether morphologically similar taxa from different continents are

conspecific or distinct. However, where we have only old or poorly preserved

collections, historical studies are equally important for correcting past mistakes

involving European names misapplied to South American species and for

halting dissemination of errors, such as those involving the geographical

distribution of certain species.

Materials & methods

The materials examined herein include types and other Pluteus collections studied

by Hennings and Rick and cited in their publications (Hennings 1900, 1904ab; Rick

1907, 1919, 1930, 1938, 1961). We also re-examined unpublished collections that they

studied. To locate as many collections studied by them as possible, we consulted B, BPI,

FH, K, KIEL, L, PACA, S, SP, and W; herbarium acronyms follow Thiers (2013). When

necessary, additional collections were also studied to help identify specimens.

The taxa are arranged alphabetically according to the names used by Hennings and

Rick. Collections studied by them under the name presented in the taxonomic header

are listed under SPECIMEN(S) EXAMINED with previously published names followed by

an asterisk (*). Under ADDITIONAL SPECIMEN(S) EXAMINED are listed the collections that

share the same characters as the materials listed in SPECIMEN EXAMINED but preserved in

herbaria under other names. Specimens that do not represent the species suggested for

the other materials involved under this name are also listed in ADDITIONAL SPECIMEN(S)

EXAMINED, but the correct identification and name are discussed in the text. For each

taxon, we comment upon the name attributed by Hennings or Rick and about what

these collections actually represent based on our morphological examination. Original

descriptions are presented for the taxa described from Brazilian types as well as for

published names for which we could not locate specimens. A summary is provided in

TABLE 1. Generic and infrageneric concepts follow Singer (1959, 1986) as complemented

by Justo et al. (2011a,b).

For microscopic analyses, the dried material was wetted with 70% ethanol and

then rehydrated in 5% KOH or stained with Melzer’s reagent to determine the amyloid

reaction of the spores. The notation [a/b/c] preceding basidiospore measurements =

(a) basidiospores were measured from (b) basidiomata taken from (c) collections; Q =

range of the length/width ratio for all spores measured; Qm = average of all calculated

Q values for all spores measured; and Lm (Wm) = the average of all the lengths (widths)

of the measured basidiospores. At least 20 basidiospores from each basidioma were

measured in lateral view, and the terms denoting basidiospore shape follow Bas (1969).

For the pileipellis having a hymeniderm, the length of the pedicel was included in the

measurement of the pileipellis elements, and its length is also reported separately.

Hennings & Rick Pluteus revisions (Brazil) ... 195

Results

Of the 32 names of taxa associated with collections recovered, Hennings

studied three and Rick the other 29 (TABLE 1). In addition to the 21 taxa

published by Rick (1907, 1919, 1930, 1938, 1961), eight additional names

were recovered from his unpublished materials. Of the ten taxa represented by

Brazilian types, five were not found in any herbarium consulted, and we now

regard these as nomina dubia. We were also unable to locate collections bearing

five other names listed here. Our suggested identifications are based on species

actually described from the Neotropics, making the European names originally

applied by Hennings and Rick inappropriate in all cases.

TABLE 1: Pluteus species reported from Brazil by Hennings and Rick.

SPECIES

“P. brunneopictus”*

“P. cervinus”

P. cervinus var. griseoviridis®

“P. cervinus vat. patricius”

P. cristatulus®

“P. dissimilis”

“P. egregius”

“P. exiguus”

“P. exiguus var. venosa”®

“P. eximius”

P. fibrillosus®

“P. fuscidulus”

“P. granulatus”

“P. hispidulus”

“P. leoninus”

P. leptonia®

“P. longistriatus”

“PR. melanodon”

“P. murinus”

“P. nanus”

“P. nanus var. podospileus”

“P. pellitus”*

“P. princeps”

“P. phlebophorus”*

P. scruposus®

P. sensitivus®

“P. sensitivus var. macrospora”

P. straminellus®

P. termitum®

“P. umbrosus”

P. velatus®

“P. wehlianus”

REFERENCES

Rick 1938, 1961

Hennings 1904b

Rick 1907

Rick 1938, 1961

Unpublished

Rick 1938, 1961

Rick 1961

Rick 1919, 1938, 1961

Rick 1938, 1961

Unpublished

Rick 1930, 1938, 1961

Rick 1919, 1938, 1961

Rick 1938, 1961

Unpublished

Rick 1938, 1961

Unpublished

Rick 1919, 1938, 1961

Rick 1938, 1961

Rick 1930, 1938, 1961

Unpublished

Rick 1938, 1961

Hennings 1900

Rick 1930, 1938, 1961

Unpublished

Rick 1961

Hennings 1904a

Rick 1919, 1938, 1961

Rick 1961

Unpublished

COMMENTS

Probably P. tucumanus and P. cubensis

P. xylophilus

Probably P. angustisporus

Not found

Not found (nomen dubium)

Probably P. angustisporus

P. xylophilus

Not found

Probably P jamaicensis

Not found

Not found (nomen dubium)

Insufficient for study

Probably P. glaucotinctus

Probably P. yungensis

Probably P. conizatus

Entoloma s.1.

Probably a true P. longistriatus

Not found

P. albostipitatus

Probably P. sapiicola

Not found

Probably P petasatus

or another sp. in sect. Hispidoderma

Probably P. angustisporus

Probably P. sapiicola and P. tucumanus

Not found (nomen dubium)

P. albostipitatus

Nomen dubium

Not found (nomen dubium)

Probably P. angustisporus

Nomen dubium

Bolbitiaceae or Strophariaceae

® = type locality in Brazil; * = species represented by collections of two different species;

Bold font = references that provide collection/herbarium information.

196 ... Menolli & Capelari

Sections Pluteus and Celluloderma are represented by the greatest number

of collections; the number of sampled collections cited by Justo et al. (2012b)

suggest that these two sections are probably the most representative of the

genus. A current study (Menolli, Meijer & Capelari, unpublished data)

including a morphological revision of Pluteus specimens collected during

more than 30 years of random field trips throughout South Brazil also reveals

a preponderance of species in these sections, further confirming observations

on sections Celluloderma and Pluteus in the Neotropics by Singer (1959) and

Horak (1964).

Taxonomy

“Pluteus brunneopictus Berk.’ sensu Rick, Iheringia, Sér. Bot. 8: 420.1961. Fic.1

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Santa Maria, 1936, J. Rick s.n.

(PACA14522*).

Basipiospores [20/1/1] 5.0-6.2(-7.5) x 3.7-5.0 um [Q = 1.24-1.35

(-1.50); Qm = 1.31; Lm = 5.7 um; Wm = 4.4 um], broadly ellipsoid to ellipsoid,

inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIs a hymeniderm,

composed of sphaeropedunculate or clavate to clavate-pedicellate cells, 27-40

(-50) x 11.2-18.7(-22) um, with a medium to long pedicel (5.0-17.5 um long),

thin-walled, with brown internal content, frequently condensed or sometimes

dissolved. PLEURO- AND CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS

absent.

ADDITIONAL SPECIMEN EXAMINED — Brazil Rio Grande do Sul: Sao Leopoldo, 1943, J.

Rick s.n. (PACA14531).

Figure 1. “Pluteus brunneopictus” (PACA14522).

a. Basidiospores; b. Pileipellis cells. Scale bars = 10 um.

Hennings & Rick Pluteus revisions (Brazil) ... 197

FIGuRE 2. “Pluteus brunneopictus” (PACA14531).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

ComMENTS — Pluteus brunneopictus (Berk. & Broome) Sacc. isan obscure species

described from Sri Lanka based on one sample collected in 1868 (Berkeley &

Broome 1871, Saccardo 1887). Although there are no recent collections, Singer

(1956) and Pegler (1986) re-examined the type and confirmed the occurrence

of broadly ellipsoid to ellipsoid basidiospores measuring 5.3-7.3 x 4.5-4.8 um

(Singer) and 7.0-9.0 x 5.0-6.0 um (Pegler) and a cellular pileipellis composed

of ellipso-pedicellate elements with brownish pigment. Although these data are

similar to what we observed in the material studied by Rick (1938, 1961), they

are not enough (mainly because pleuro- and cheilocystidia were not found) to

correlate it to P brunneopictus.

Moreover, considering the geographical distance, it is unlikely that

the specimen collected by Rick in South America actually represents

P. brunneopictus. Singer (1959) suggested that “P brunneopictus” sensu

Rick might be P. umbrinoalbidus Singer, P. rimosoaffinis Singer, or any other

species in stirps Pulverulentus, Jamaicensis, or Tucumanus. Based on the

insufficient information available because the hymenial structures were

not recovered, we consider that this collection most likely represents P

tucumanus Singer, originally described from Argentina. According to Singer

(1959), PB tucumanus has ellipsoidal basidiospores (3.8-5.5 x 2.7-3.8 um)

and subglobose-pedicellate pileipellis cells with dissolved or condensed

intracellular pigment, and it represents one of the few South American species

198 ... Menolli & Capelari

in sect. Celluloderma characterized by a predominantly brown pileus color and

ellipsoid basidiospores. More collections of Pluteus from Brazil and Argentina

with these characteristics would be required to establish the actual relationship

between Rick’s collection and Singer's species and to confirm its occurrence in

Brazil.

It is also important to note that Rick (1961) mentioned under P. brunneopictus

two collections (PACA14522 and PACA14532) made in Santa Maria County

in 1936. However, the exsiccata deposited at PACA mistakenly labeled as

(PACA)14532 actually contains PACA14531 (Brazil. Rio Grande do Sul: Sao

Salvador, 1943). PACA14531 (Fic. 2) represents an unpublished sample, and our

re-examination showed it has broadly ellipsoid to rarely ellipsoid basidiospores

measuring [20/1/1] 6.2-7.5(-8.7) x 5.0-6.2(-7.5) um [Q = (1.16—-)1.21-1.24

(-1.50); Qm = 1.24; Lm = 7.1 um; Wm = 5.7 um], clavate or elongate-ventricose

pleurocystidia (47-68 x 15.0-22 um), a pileipellis that seems to be a cutis

with ascendant elements, and no clamp connections. Cheilocystidia were not

found. Based on these characteristics and on a current study (Menolli, Meijer &

Capelari, unpublished data) that includes a broad examination of collections of

P. cubensis (Murrill) Dennis, this seems to be the better name for PACA14531.

Pluteus cubensis is widely distributed in Central and South America, and its

representatives display wide morphological variation, especially in basidiospore

shape and size.

“Pluteus cervinus Schaeff. sensu Rick, Iheringia, Sér. Bot. 8: 418. 1961. FIG. 3

SPECIMENS EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1906, J. Rick

s.n. (PACA14519*); 1934, J. Rick sn. (PACA14516*); 1934, leg. Steffen, det. J. Rick

(PACA14526).

BASIDIOSPORES [62/4/4] (5.0-)6.2-7.5(-8.7) x (3.7-)5.0-6.2 um [Q =

(1.21-)1.24-1.68; Qm = 1.45; Lm = 6.7 um; Wm = 4.8 um], broadly ellipsoid,

ellipsoid to elongate, inamyloid, hyaline, smooth, thick-walled, guttulate.

PLEuUROCYSTIDIA of three types: (1) the normal Cervinus-type, 58-76 x 15.0-

17.5 um, fusoid-ventricose, colorless, hyaline, with four to six apical to lateral

prongs at the apices, thick-walled, the wall sometimes tapering toward the base,

abundant to very abundant; (II) the modified Cervinus-type as the majority

of those found in P. harrisii Murrill, 56-87 x 12.5-20 um, fusoid-ventricose,

colorless, hyaline, usually without prongs and a subcapitate apex or with two

short lateral prongs and slightly strangulated at the apex, moderately thick-

walled, common; and (III) the Magnus-type, 56-78 x 12.5-21 um, lageniform

to fusoid, with an acute apex, colorless, hyaline, moderately thick-walled, rare.

CHEILOCYSTIDIA not recovered. PILEIPELLIS a cutis. CLAMP CONNECTIONS

absent.

ADDITIONAL SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Pareci Novo, 1918, J.

Rick s.n. (SP33928 as “P. egregius” Rick).

Hennings & Rick Pluteus revisions (Brazil) ... 199

Ficure 3. “Pluteus cervinus” (PACA14519).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

COMMENTS — Our re-examination of Rick’s collections identified as P. cervinus

shows that they actually represent P xylophilus (Speg.) Singer. Although we

were unable to find any cheilocystidia, the characteristic pleurocystidia

combined with the basidiospore size and shape and the macroscopic aspect

of the pileus are distinctive enough to identify Rick's collections. Pluteus

xylophilus can be easily recognized by its large pileus, the presence of three types

of pleurocystidia, and the broadly ellipsoid to elongate basidiospores (Singer

1959, Menolli et al. 2010). There is much discussion about the identity and

circumscriptions of P cervinus (Schaeff.) P. Kumm., and it is likely restricted to

the northern hemisphere, as shown in the phylogenetic analyses published by

Justo et al. (2011b). As suggested by Singer (1959), almost all collections from

Brazil referred to P cervinus are probably P. xylophilus, a species well known

in South America and which forms a distinct clade in sect. Pluteus (Justo et al.

2011b).

Another collection (PACA14529) of P. cervinus recorded by Rick (1961) was

also not found; however, “PACA14529” may be a typographical error for the

specimen PACA14526 that we studied.

200 ... Menolli & Capelari

Pluteus cervinus var. griseoviridis Henn., Hedwigia 43: 204. 1904. FIGS. 4, 5

SPECIMEN EXAMINED — Brazil. Sao Paulo: Sao Paulo, Serra da Cantareira, March 1903,

A. Puttemans 871 (Holotype, SP141797).

ORIGINAL DESCRIPTION — “Pileo carnoso, convexo explanato, centro obtuso, laevi glabro,

subglutinosulo, margine striatulo, griseo-viridi, 5-7 cm diam.; stipite solido, cylindraceo,

pallido, laevi, glabro, 4-7 x 3-5 y [sic; = cm], basi incrassato; lamellis liberis, confertis,

ventricosis, ca. 3-4 mm latis, flavido-incarnatis; sporis ellipsoideis, 1-guttulatis, carneis,

4-5 x 4 u; cystidiis lageniformibus, apice 2-vel 3 dentato-hamatis, 40-60 x 15-20 wy.”

Basiprospores [100/5/5] 5.0-7.5(-8.7) x (3.1-)3.7-5.0 um [Q = 1.24-

1.70(-2.03); Qm = 1.57; Lm = 6.3 um; Wm = 4.1 um], broadly ellipsoid,

ellipsoid to elongate, rarely cylindrical, inamyloid, hyaline, smooth, thick-

walled, guttulate. PLEURocysTIpIA 50-95 x 10.0-24 um, fusoid-ventricose,

colorless, hyaline, mostly of the Cervinus-type with four to six apical or lateral

prongs, or like those found in P harrisii that are usually without or with two

short lateral prongs, sometimes with lateral secondary spinules such as those of

P spinulosus Murrill, thick-walled, abundant to very abundant. CHEILOCYSTIDIA

not recovered in most collections but a clavate form (39 x 15.0 um) was found

in BPI770905. PILEIPELLIS a cutis. CLAMP CONNECTIONS absent.

FiGurE 4. Pluteus cervinus var. griseoviridis (holotype, SP141797).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

Hennings & Rick Pluteus revisions (Brazil) ... 201

FIGuRE 5. “Pluteus cervinus” (BPI770858).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

ADDITIONAL SPECIMENS EXAMINED — Brazil. Rio Grande do Sul: Arroio do Meio, 1920,

J. Rick s.n. (SP33926 as P. cervinus); 1920, J. Rick s.n. (BPI770858 as P. cervinus); 1920, J.

Rick s.n. (BPI770893 as “P. princeps” Rick); 1920, J. Rick s.n. (BP1770905 as P. umbrosus);

Sao Leopoldo, 1929, leg. Steffen, det. J. Rick (PACA14517 as “P. dissimilis” Rick).

COMMENTS — Our re-examination of the holotype of P cervinus var.

griseoviridis, along with some unpublished collections identified by Rick as

P. cervinus and P. umbrosus and others named as “P. dissimilis” and “P. princeps,”

showed that they share some morphological features with P angustisporus

Singer described from Bolivia (Singer 1959).

Pluteus angustisporus is morphologically close to P. xylophilus and P. harrisii,

from which it differs mainly by the narrow basidiospores and the absence of

Magnus-type pleurocystidia as found in P. xylophilus. Although the collections

we examined have large pilei, such as those observed in P. xylophilus (<110 mm;

Menolli et al. 2010) and much bigger than those reported for P. angustisporus

(<25 mm; Singer 1959), Magnus-type pleurocystidia were not found in any

collection studied. In addition, Singer (1959) also reported pleurocystidia

with polymorphic apices for P angustisporus like those we observed (Fie. 5).

However, due to the condition of the exsiccatae, it was not possible to confirm

the presence of polymorphic cheilocystidia and pileipellis elements as reported

by Singer (1959).

202 ... Menolli & Capelari

Although we cannot confirm the actual identity of these collections, the

recovered data indicate that these collections are clearly close to or identical

with either P. angustisporus or P. xylophilus, and we tentatively conclude that

P. angustisporus is the most appropriate name.

It is important to note that although P cervinus var. griseoviridis was

described before P. angustisporus, if they are the same taxon, the earlier varietal

name does not have priority at species rank.

“Pluteus cervinus var. patricius Schulz.; sensu Rick, Brotéria, Sér. Bot. 6: 71. 1907.

ORIGINAL DESCRIPTION — “Pileo 1 dm. lato, crasso, firmo; stipite solido, usque 4 cm.

lato, albido, griseo-squamoso. Differt a typo pileo albo, griseo-squamoso, primitus cinereo.

Sporis 7-8 yu longis, 5-6 u latis. Est varietas prorsus identica cum illa a cl. Schulzer in

Hungaria lecta. Legi quoque typum.”

ComMENTs — ‘The infraspecific taxon P cervinus var. patricius (Schulzer)

Massee (= P. cervinus subsp. patricius (Schulzer) Sacc.) was described from

Europe. Although Rick (1907) recorded P. cervinus var. patricius, he did not

cite the collection studied. No sample collected by Rick under this name was

found in the herbaria, but it most likely also represents P. xylophilus.

FiGure 6. Pleurocystidia of “Pluteus dissimilis” (PACA14517). Scale bar = 10 um.

Hennings & Rick Pluteus revisions (Brazil) ... 203

Pluteus cristatulus Rick, Lilloa 3: 445. 1938.

ORIGINAL DESCRIPTION — “Pileo membranaceo, albo-brunneo, innato-squamuloso,

1 cm. lato; stipites vitreo, albo-consperso, curto, 2 mm. lato, laterali; lamellis liberis, albis,

ventricosis, subconfertis acie serrates, versus stipitem rotundatis; sporis 4.5 x 3 uw rubris. In

trunco. Differt a Pl. exiguo lamellis albis serratis, pileo non striato et statura. ”

COMMENTS — Rick (1938, 1961) did not mention the collection studied, and

no sample collected by Rick under this name was found in the herbaria. We

therefore regard this name as a nomen dubium.

“Pluteus dissimilis Rick? nom. in herb. FIG. 6

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1929, leg. Steffen, det.

J. Rick (PACA14517).

ComMMENTS — See the description under P. cervinus var. griseoviridis, with

which it shares the same features. It probably represents P. angustisporus.

“Pluteus egregius Rick; nom. in herb. FIG. 7

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Pareci Novo, 1918, J. Rick s.n.

(SP33928).

Figure 7. Pleurocystidia of “Pluteus egregius” (SP33928). Scale bar = 10 um.

204 ... Menolli & Capelari

CoMMENTS — See the description under P. cervinus, with which it shares the

same features. We feel it represents P. xylophilus.

“Pluteus exiguus Pat.’ sensu Rick, Lilloa 3: 446. 1938.

ORIGINAL DESCRIPTION — “Pileo convexo, dein plano, tenui, 1-2 cm. lato, initio laevi,

dein margine striatulo, pilis brevissimis, erectis, brunneis, flocculosis, velutino, brunneo-

rufo, centro obscuriore, lamellis rotundatis inaequalibus, a stipites remotis rufis; stipites

fareto, 10-15 mm. longo, basi incrassato, albido, pruinose, non striato; sporis sphaeroides,

1-guttulatis, roseis 6-7 = |x] 3 u. Ad terram. PI. nano affinis.”

ComMENts — Although Rick (1938) did not mention any collection studied,

he (Rick 1961) later referred this name to PACA14518, but the package of this

collection is labeled “P exiguus var. venosa” (see below). The occurrence of

P. exiguus (Pat.) Sacc. or any variety of this species in Brazil seems unlikely. It

is important to consider that the description of P exiguus presented in Rick

(1938, 1961) is almost identical to that provided by Saccardo (1887) and that no

other material under this name was located in the herbaria, making it difficult

to confirm its occurrence in Brazil. In addition, as far as we know, P. exiguus

appears to be restricted to Europe, where it has been rarely documented

(Kithner & Romagnesi 1956, Orton 1986, Justo & Castro 2007a, Ripkova 2009).

“Pluteus exiguus var. venosa” Rick, Theringia, Sér. Bot. 8: 417. 1961, nom. inval. Fic. 8

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Marcelino Ramos, 1936, J. Rick s.n.

(PACA14518, as P. exiguus var. venosa).

ORIGINAL DESCRIPTION — “Pileo brunneo, venoso, lamellis mediis, densis, stipite vitreo.

Sporis 4-5 my [um]. Ad ligna.”

BasiprosPores [20/1/1] 5.6-7.5 x (4.6-)5.0-6.2 um [Q = (1.09-)1.12-1.24);

Qm = 1.20; Lm = 6.3 um; Wm = 5.3 um], subglobose to broadly ellipsoid,

inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIs a hymeniderm,

composed of subglobose to spheropedunculate cells, 26-37 x 18.7-26.5 um,

almost non-pedicellate or with a short pedicel (1.2-3.7 um long), thin-walled,

with brown dissolved content. PLEURO- AND CHEILOCYSTIDIA not recovered.

CLAMP CONNECTIONS absent.

ComMENts — Although PACA14518 is labeled “P. exiguus var. venosa, in the

protologue Rick (1961) cited this collection under P exiguus, not under the

variety “venosa.” As no holotype was designated for the variety, the name is not

validly published.

The proposal of a new variety by Rick (1961) seems inappropriate because

PACA14518 has a hymenidermal pileipellis more typical of sect. Celluloderma

than sect. Hispidoderma under which P. exiguus is classified. The scarce

morphological data obtained from Rick’s collection are insufficient to designate

a correct name for it, but we agree with Singer (1959) that Rick most likely had

Hennings & Rick Pluteus revisions (Brazil) ... 205

FiGure 8. “Pluteus exiguus var. venosa” (PACA14518).

a. Basidiospores; b. Pileipellis cells. Scale bars = 10 um.

P. jamaicensis Murrill, which is characterized by non-globose basidiospores

[5.0-6.8(-7.5) x 4.5-6.2 um] and pileipellis cells (21-60 x 20-45 um) with

brown dissolved content (Singer 1956, 1959; Smith & Stuntz 1958; Horak 1964;

Pegler 1983; Banerjee & Sundberg 1993; Menolli et al. 2010).

“Pluteus eximius Saund. et Sm.’ sensu Rick, Lilloa 3: 445. 1938.

ORIGINAL DESCRIPTION — “Pileo laevi, pellicula separabili viscida tecto, rufescenti-

umbrino; stipites fibrilloso, demun nigricante. In serragine. Maximus speciosus. Est forma

luxurians PI. cervini.”

CoMMENTS — ‘The English infraspecific taxon P cervinus var. eximius (W.

Saunders & W.G. Sm.) Massee (= P. cervinus subsp. eximius (W. Saunders &

W.G. Sm.) Sacc.) has never been raised to species rank. Rick (1919, 1938, 1961)

did not mention the collection studied, and no sample collected by Rick was

found in the herbaria under this name.

Pluteus fibrillosus Rick, Lilloa 3: 444. 1938.

ORIGINAL DESCRIPTION — “Pileo conico, carnoso, fibrilloso, atro-brunneo, fibris

glandulosis ab umbone usque ad margine picto et albo-rimoso usque 5 cm. lato; stipites 1

dm. longo virgineo, laevi, lamellis remotis, confertis, vix ventricosis, versus stipitem liberis

et rotundatis; sporis sphaerics 5 u. Ad terram. Videtur Pl. psichiophoro, albo-lineato et

apilopu affinis. Est varietas Pl. longestriati Peck. Hi omnes exceptis 2 primis videntur unam

sister speciem proximan Pl. nano Europaeo.”

COMMENTS — Rick (1938, 1961) did not mention the collection studied, and

no sample collected by Rick was found in the herbaria under this name. We

therefore regard this name as a nomen dubium.

206 ... Menolli & Capelari

“Pluteus fuscidulus Fr.? nom. in herb.

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Salvador, 22 Jul 1944, J. Rick s.n.

(PACA20926).

Comments — This collection is in too poor condition for study of any

microstructures or to gain insights about its identification. Moreover, there is

no Pluteus (nor Agaricus) named by Fries as ‘fuscidulus’ Because the collection

is unfit for study and the name not attributable to Fries, we have nothing to add

to this record.

oS)

Figure 9. Basidiospores. a. “Pluteus granulatus” (PACA14523);

b-c. “Pluteus hispidulus” (b. PACA14528; c. PACA14532). Scale bars = 10 um.

“Pluteus granulatus Bres.’ sensu Rick, Iheringia, Sér. Bot. 8: 418. 1961. FIG. 9A

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1929, J. Rick s.n.

(PACA14523*).

BASIDIOSPORES [20/1/1] 6.2-7.5 x 5.0-6.2 um (Q = 1.21-1.50; Qm = 1.28;

Lm = 7.2 um; Wm = 5.7 um), preponderantly broadly ellipsoid or sometimes

ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIs

composed of hyphal elements typical of a cutis but it is too difficult to be

recovered. PLEURO- AND CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS

not observed.

ComMENTs — Pluteus granulatus Bres. is a rarely recorded mushroom that

seems to be restricted to Europe (Saccardo 1887, Kihner & Romagnesi 1956,

Orton 1986). It is considered a synonym of P. plautus (Weinm.) Gillet, in a

broad sense (Vellinga & Schreurs 1985, Legon & Henrici 2011).

Singer & Digilio (1952) recorded P plautus from Argentina, but Singer

(1959) later re-identified the collection as P. argentinensis Singer. According

to Singer (1959), Rick’s collection was misidentified, and our re-examination

suggests that it most likely represents some species with a cutis-like pileipellis

characterized by an umber fibrillose pileus and broadly ellipsoid to ellipsoid

basidiospores. It is possible that Rick had P. argentinensis, P. fastigiatus Singer,

Hennings & Rick Pluteus revisions (Brazil) ... 207

P. fuliginosus Murrill, or (most likely) P. glaucotinctus E. Horak that was recently

recorded from Brazil (Justo et al. 201 1a,b).

“Pluteus hispidulus Fr’ sensu Rick, Theringia, Sér. Bot. 8: 419. 1961. FIG. 9B, C

SPECIMENS EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, Rick s.n.

(PACA14528*); 1932, Rick s.n. (FH00301668); Sao Salvador, 1943, Rick s.n.

(PACA14532*).

Basip1ospores [60/3/3] 6.2-10.0 x 5.0-7.5 um (Q = 1.16-1.50; Qm = 1.30;

Lm = 7.8 um; Wm = 6.0 um), broadly ellipsoid to ellipsoid, inamyloid, hyaline,

smooth, thick-walled, guttulate. PILEIPELLIS and HYMENIAL STRUCTURES not

recovered. CLAMP CONNECTIONS not observed.

ADDITIONAL SPECIMEN EXAMINED — Brazil. J. Rick s.n. (BPI839783).

COMMENTS — Originally described from Europe, P. hispidulus (Fr.) Gillet has

not yet been definitively recorded from South America. Because our recovered

data are insufficient to allow our renaming of Rick's collections, it is not possible

to determine what species Rick had collected.

Singer (1959) suggested that this might be a species of sect. Hispidoderma

(stirps Fuliginosus). The most probable species might be P. yungensis Singer,

which has the biggest basidiospores comparable in size to those observed in

Rick’s collections. Modern collections with these characteristics are needed

to clarify whether P yungensis really occurs in Brazil. We revised another

unpublished collection (BPI839783), labeled by Rick as P. hispidulus, that

actually represents Entoloma s.1.

“Pluteus leoninus Schaeff.’ sensu Rick, Iheringia, Sér. Bot. 8: 417. 1961. FIG. 10

SPECIMEN EXAMINED — Brazil. Rio Grande Sul: Sao Leopoldo, 1932, J. Rick s.n.

(FH00301669).

Basipiospores [20/1/1] 5.0-6.2(-7.5) x 3.7-4.6(-5.0) um [Q = (1.09-)

1.22-1.51(-1.68); Qm = 1.37; Lm = 5.7 um; Wm = 4.2 um), broadly ellipsoid

to ellipsoid, rarely subglobose or elongate, inamyloid, hyaline, smooth, thick-

walled, guttulate. PLEUROCYSTIDIA 52-87 x 12.5-26 um, fusoid-ventricose

to lageniform, usually with a rounded apex, metuloidal with thickened wall

and lacking any apical or lateral horns, frequently with pale straw or colorless

internal condensed or dissolved content. CHEILOCYSTIDIA not recovered.

PILEIPELLIs difficult to recover but apparently a cutis composed of hyphae with

pale brown pigment. CLAMP CONNECTIONS absent.

ComMMENTS — Pluteus leoninus (Schaeff.) P. Kumm. belongs to sect.

Hispidoderma and is apparently restricted to the northern hemisphere.

The species is characterized by golden yellow basidiomata and thin-walled

pleurocystidia with little excrescences at the apex (Singer 1956, Vellinga &

Schreurs 1985, Orton 1986, Takehashi & Kasuya 2007). However, “P. leoninus”

208 ... Menolli & Capelari

Ficure 10. “Pluteus leoninus” (FH00301669).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

sensu Rick has thick-walled pleurocystidia. Considering other species with

yellowish basidiomata and non-horned metuloidal pleurocystidia, it is likely

that Rick had P. conizatus (Berk. & Broome) Sacc. or P. amphicystis Singer,

which differ from each other in pileipellis structure and the absence of metuloid

caulocystidia in P conizatus (Singer 1956, 1959; Pegler 1983, 1986; Pradeep et

al. 2002). Because the pileipellis in Rick’s collection was not well characterized,

we verified that the caulocystidia are absent in this collection. We believe that

Rick probably had P. conizatus instead of P leoninus.

Pluteus leptonia Rick, Lilloa 3: 445. 1938.

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1936, J. Rick s.n.

(Holotype, PACA14525*).

ORIGINAL DESCRIPTION — “Pileo conico, demum expanso-fisso, carnoso, 3-4 cm. lato,

subumbonato, laevi, minute ruguloso, exstrio; lamellis aperte liberis, densis, subventricosis,

evidenter erosis carneis; stipite usque 1 dm. Longo 5 mm lato, firmo, demum cavo, nudo,

striato, cartilagineo; sporis angulatis 7-10 micr. apiculatis. Omnibus partibus pulcherrime

Hennings & Rick Pluteus revisions (Brazil) ... 209

lazulinus aut coeruleus exceptis lamellis carneis e carne carnea. Odore et sapore nullo. In

terra inter Bambusas. Pluteus phaeus Sacc. XVI et sine dubio proximus nisi identicus, sed

mea species non crescit in ligno, sed in terra.”

ComMENnts — ‘The type of P leptonia is not a Pluteus; it is actually a member

of Entolomataceae. Its basidiospores are predominantly isodiametric, with 5-7

angles. Singer (1956, 1959) mentioned the presence of typical Rhodophyllus

basidiospores, and he suggested that Rick’s collection might represent Nolanea

howellii Peck or another allied species.

“Pluteus longistriatus (Peck)? nom. in herb.

SPECIMEN EXAMINED — Brazil. Rio Grande Sul: Sao Leopoldo, 1932, J. Rick s.n.

(FH00301670).

BASIDIOSPORES [20/1/1] 6.2-7.5 x 5.0-6.8 um (Q = 1.07-1.24; Qm = 1.19; Lm

= 7.0 um; Wm = 5.9 um), subglobose to broadly ellipsoid, inamyloid, hyaline,

smooth, thick-walled, guttulate. PILEIPELLIS and HYMENIAL STRUCTURES not

recovered. CLAMP CONNECTIONS not observed.

ComMENTS — Pluteus longistriatus (Peck) Peck, which is common in subtropical

and temperate areas of North and South America, is usually recognized by

a pileus with a deeply sulcate-striate margin and a hymenidermal pileipellis

with heteromorphic elements (Singer 1959, Menolli & Capelari 2010). The

macroscopic appearance of FH00301670 suggests that it had a sulcate-striate

pileus when fresh, and the basidiospore size and shape agree with those usually

reported for P. longistriatus (Singer 1959, Menolli & Capelari 2010). Based

on these data, we suspect that Rick’s collection is actually P longistriatus, but

we cannot confirm the identification and the occurrence of this species from

the state of Rio Grande do Sul with certainty because the pileipellis and the

hymenial structures were not recovered from this collection.

“Pluteus melanodon Secr: sensu Rick, Theringia, Sér. Bot. 8: 418. 1961.

ORIGINAL DESCRIPTION — “Stipite cavo; lamellis acie atris, incises; an P. umbrosus? Ad

lignum.”

ComMMENTs — The European species invalidly described by Secretan was

subsequently named as P melanodon (Fr.) Sacc. Rick (1938, 1961) did not

mention the collection studied, and no sample collected by Rick under this

name was found in the herbaria.

“Pluteus murinus Bres.} nom. in herb. FIG. 11

SPECIMEN EXAMINED — Brazil. J. Rick s.n. (BPI770882).

BASIDIOSPORES [40/2/2] (6.2—)7.5-8.7 x (5.0—)6.2-7.5 um [Q = (1.00-)1.16-

1.24(-1.60); Qm = 1.19; Lm = 7.6 um; Wm = 6.4 um), preponderantly broadly

ellipsoid or sometimes globose or ellipsoid, inamyloid, hyaline, smooth, thick-

210 ... Menolli & Capelari

a. (

ivi

FiGuRE 11. “Pluteus murinus” (BPI770882).

a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 um.

walled, guttulate. PLEUROCYSTIDIA 51-69(-86) x 10.0-22.5 um, elongate-

clavate or slightly ventricose, thin- to relatively thick-walled, sometimes with

a colorless internal condensed content. CHEILOCYSTIDIA 50-66 x 12.5-15 um,

clavate with a well-developed pedicel at the base, thin-walled, colorless and

hyaline. PILEIPELLIS a cutis composed of hyphal elements with a rounded apex

and brownish dissolved pigment. CLAMP CONNECTIONS not observed.

ADDITIONAL SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Salvador, 1944, J.

Rick s.n. (PACA20770 as P. sensitivus var. macrospora).

ComMENTs — Pluteus murinus Bres., apparently restricted to Europe (Singer

1956, Orton 1986, Banerjee & Sundberg 1993), is considered a synonym of

P. ephebeus (Fr.) Gillet (Vellinga & Schreus 1985). The data recovered from

Rick’s collections indicate that it shares the main characteristics found in

P. albostipitatus (Dennis) Singer, a widely distributed and commonly cited

species with some synonyms and wide morphological variation: mainly the

presence of fibrils on the pileus, pleurocystidial wall thickness, and basidiospore

size and shape (Menolli et al. 2010; Justo et al. 2011b).

“Pluteus nanus Pers.’ sensu Rick, [heringia, Sér. Bot. 8: 418. 1961. Fic. 12

SPECIMENS EXAMINED — Brazil. Rio Grande do Sul: Marcelino Ramos, 1936, Rick s.n.

(FH00301671); Sao Salvador, 29 Feb. 1944, Rick s.n. (PACA22618*).

Hennings & Rick Pluteus revisions (Brazil) ... 211

FiGuRE 12. “Pluteus nanus”. (a-b. FH00301671) a. Basidiospores; b. Pileipellis cells.

(c-d. PACA22618) c. Basidiospores; d. Pileipellis cells. Scale bars = 10 um.

BasIpiosPorEs [40/2/2] 5.0-7.5 x 5.0-6.2(-7.5) um [Q = 1.00-1.12(-1.24);

Qm = 1.08; Lm = 6.3 um; Wm = 5.9 um], globose or occasionally broadly

ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS a

hymeniderm composed of subglobose cells, 20-59 x 15.0-34 um, with a short

to moderately long pedicel (1.2-10.0 um long), thin-walled, with brownish

condensed or dissolved content. PLEURO- AND CHEILOCYSTIDIA not recovered.

CLAMP CONNECTIONS not observed.

ComMMENTS — Pluteus nanus (Pers.) P. Kumm. is a species with many

interpretations and seems to be restricted to Europe (Vellinga & Schreurs 1985,

Orton 1986, Justo et al. 2007, Justo et al. 2011b). Although there is a suspect

record from North America (Homola 1972), it remains unclear if P nanus is

really found in the USA according to any of these interpretations (Minnis &

Sundberg 2010). Singer (1959) considered Rick’s identification to be dubious.

Considering the rugose-venose pileus, globose basidiospores, and pileipellis

cells with brownish condensed or dissolved content, it is possible that Rick had

212 ... Menolli & Capelari

P. pulverulentus Murrill or (most likely) P sapiicola Singer, a species known

to occur in Brazil (Menolli, Meijer & Capelari, unpublished data). However,

other species with pigmented cystidia, such as P. beniensis Singer, P. eliae Singer,

P. rimosoaffinis, and P. riograndensis Singer, should also be considered.

“Pluteus nanus var. podospileus (Sacc. et Cub.)” sensu Rick, Lilloa 3: 444. 1938.

ORIGINAL DESCRIPTION — “Differt stipite brunneo.”

ComMENTS — Pluteus podospileus Sacc. & Cub. is a widespread European

species, distinct from P nanus; it is unlikely that Rick would have collected

authentic material in Brazil. Rick (1938, 1961) did not mention the collection

studied, and no sample collected by Rick under this name was found in the

herbaria.

“Pluteus pellitus Pers.” sensu Rick, Iheringia, Sér. Bot. 8: 418. 1961. FIG. 13

SPECIMENS EXAMINED — Brazil. Rio Grande do Sul, Sao Leopoldo, Rick s.n.

(PACA14524*); 1931, Rick s.n. (FH00301673); 1932, Rick s.n. (PACA14530*); Sep.

1933, Rick s.n. (FH00301672).

Basipiospores [80/4/4] (5.6-)6.2-7.5(-8.7) x (3.7-)5.0-6.2 um [Q =

1.21-1.51(-1.68); Qm = 1.28; Lm = 6.7 um; Wm = 5.3 um], preponderantly

broadly ellipsoid or sometimes ellipsoid, rarely elongate, inamyloid, hyaline,

smooth, thick-walled, guttulate. PLEUROCYsTIDIA 61-78 x 16.2-18.7 um,

slightly ventricose to lageniform, with an amorphous pale straw content,

pigmentation usually concentrated at apex and base, thin-walled (recovered

only in FH00301673). CHEILOCYSTIDIA and PILEIPELLIS not recovered. CLAMP

CONNECTIONS not observed.

ComMENTts — Although there are many concepts of P. pellitus (Pers.) P. Kumm.,

Justo & Castro (2007b) and Justo et al. (2011b) accepted it as a species apparently

restricted to Europe and characterized mainly by the white basidiomata, a

habitat on angiosperm wood, basidiospores of 5.0-7.5 x 3.5-5.0 um, and the

presence of clamp connections. Singer (1959) mentioned that “P pellitus”

sensu Rick “is obviously P. viscidulus, although he did not cite Rick’s collection

in the ‘examined material’ for this species. Pluteus viscidulus Singer, another

species with white basidiomata but that lacks clamp connections was described

from Argentina (Singer 1959) and recorded also from Brazil (from the same

State from which Rick's material was collected). Singer (1959) also recorded

P. viscidulus from the U.S.A. (Florida), although he mentioned that the North

American collection had bigger basidiospores and that this character can be

used to separate it from P. viscidulus.

Our re-examination of the Brazilian collection determined by Singer as

P. viscidulus (Fic. 14) [Brazil. Rio Grande do Sul: Sao Salvador, 9 Nov. 1951, Singer

B111 (LIL)] showed that it has broadly ellipsoidal to elongate basidiospores,

[20/1/1] (5.6-)6.2(-7.5) x 3.7-5.0 um [Q = 1.24-1.68; Qm = 1.35; Lm = 6.3 um;

Hennings & Rick Pluteus revisions (Brazil) ... 213

FiGurE 13. “Pluteus pellitus”. a-c. Basidiospores (a. PACA14524; b. PACA14530;

c. FH00301673); d. Pleurocystidia. (FH00301673). Scale bars = 10 um.

Wm = 4.7 um], fusoid-ventricose pleurocystidia (69-83 x 11.2-25 um) with

thick walls and apices with 2-4 lateral prongs, clavate cheilocystidia (37-61

x 12-15 um), cutis-like pileipellis, and no clamp connections. Justo & Castro

(2007b) found similar features in the isotype of P. viscidulus. Until very recently,

P. viscidulus was thought to represent an authentic South American species

with white basidiomata and without clamp-connections; however, molecular

analyses by Justo et al. (2011b) place P. viscidulus within the molecular variation

of PB. petasatus (Fr.) Gillet, and it is now treated as a synonym of P. petasatus.

Although P. petasatus (= P. viscidulus) can be confirmed from Brazil based on

Singer's collection, its morphological diagnostic characters were not seen in

Rick’s collections; the unpublished FH00301673 has thin-walled pigmented

pleurocystidia, not the horned pleurocystidia typical of sect. Pluteus found

214 ... Menolli & Capelari

FiGurE 14. Pluteus viscidulus (Singer B111, LIL).

a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 um.

in P. pellitus, P. petasatus, and allied species. In addition, according to notes

for FH00301673, Rick denoted it as P pellitus ‘videtur, which indicates that

although it resembles P pellitus, a different allied species can also be considered.

Because there are no pileus color notes for FH00301673, it is difficult to

suggest a name for it. There are no previously published species that combine

Hennings & Rick Pluteus revisions (Brazil) ... 215

broadly ellipsoid basidiospores and pigmented pleurocystidia with a whitish

pileus. However, because there is no mention whether the pileus is actually

whitish, if we consider only the species of sect. Hispidoderma with pigmented

pleurocystidia, FH00301673 might represent any species from stirps Aethalus,

Circumscissus, or Plautus, according to Singer’s classification (Singer 1959).

Although no reliable conclusion can be made about the identity of Rick’s

collections, we suggest that they do not represent P pellitus. We should not

disregard the possibility that the collections (PACA14524, PACA14530)

published in Rick (1961) could be P. petasatus, and FH00301673 is most likely

a species of sect. Hispidoderma.

“Pluteus princeps Rick; nom. in herb. FIG. 15

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Arroio do Meio, 1920, J. Rick s.n.

(BPI770893).

ComMMENTS — See the description under P. cervinus var. griseoviridis, with

which it shares the same features. The above collection probably represents

P. angustisporus.

FiGurE 15. “Pluteus princeps” (BP1770893).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

216 ... Menolli & Capelari

FiGuRE 16. “Pluteus phlebophorus”. (a-b. PACA14520) a. Basidiospores; b. Pileipellis cells.

(c-d. PACA20895) c. Basidiospores; d. Pileipellis cells. Scale bars = 10 um.

“Pluteus phlebophorus Dittm.” sensu Rick, Theringia, Sér. Bot. 8: 419.1961. Fic. 16

SPECIMENS EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1934, leg. Steffen,

det. J. Rick (PACA 14520*); Sao Salvador, 28 Jan 1944, J. Rick s.n. (PACA20895).

(PACA14520) — Basipiospores [20/1/1] 5.0(-6.2) x 3.7(-5.0) um [Q =

(1.24—)1.35(-1.68); Qm = 1.39; Lm = 5.2 um; Wm = 3.8 um], preponderantly

ellipsoid or sometimes broadly ellipsoid or elongate, inamyloid, hyaline,

smooth, thick-walled, guttulate. PILEIPELLIs a hymeniderm composed of

subglobose cells, 24-36 x 18.7-22 um, with a short pedicel (2.5-3.7 um long),

thin-walled, with brownish condensed content. PLEURO- AND CHEILOCYSTIDIA

not recovered. CLAMP CONNECTIONS not observed.

(PACA20895) — Basip1ospores [20/1/1] 5.0-6.2 x 5.0-6.2 um (Qm =

1.00; Lm = 5.7 um; Wm = 5.7 um), globose, inamyloid, hyaline, smooth, thick-

walled, guttulate. PILEIPELLIS a hymeniderm composed of subglobose cells,

31-36 x 16.2-25 um, with a short to moderately long pedicel (2.5-12.5 um

long), thin-walled, with brownish condensed or dissolved content. PLEURO-

AND CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS not observed.

Hennings & Rick Pluteus revisions (Brazil) ... 217

ComMENnts — These two collections studied by Rick have different anatomical

characters and represent two species, although they were both listed as

P. phlebophorus (Ditmar) P. Kumm. Singer (1959) suggested that

“P. phlebophorus” sensu Rick is likely P. jamaicensis, which is characterized

by non-globose basidiospores of 5.0-6.8(-7.5) x 4.5-6.2 um (Singer 1956,

1959; Smith & Stuntz 1958; Horak 1964; Pegler 1983; Menolli et al. 2010)

differing in size and shape from those observed in both of Rick's collections.

PACA14520, which shares characteristics with PACA14522 named by Rick as

P. brunneopictus, most likely represents P. tucumanus (see discussion under P

brunneopictus). PACA20895, which most likely represents P sapiicola, is similar

to PACA22618, referred by Rick to P nanus (see discussion under P nanus).

Pluteus scruposus Henn., Beibl. Hedwigia 39: 136. 1900.

ORIGINAL DESCRIPTION — “Pileo carnoso, campanulate, obtuso, scruposo, verrucis crassis

atrobrunneis tecto, 2%-3 cm diametro, brunneo; stipites farcto, tereti, albido, laevi, 3% cm

longo, 5 mm crasso, basi bulboso tomentosulo ca. 1 cm incrassato; lamellis liberis, confertis

lanceolatis pallidis, dein subincarnatis, cystidiis lanceolatis vel clavate-lageniformibus 55-

90 x 25-35 yu; basidiis clavatis 35-45 x 15-20; sporis globosis intus flavido-subincarnatis

15-17 pu.”

ComMENTsS — As we could locate no specimens, we regard this name as a

nomen dubium.

Pluteus sensitivus Rick, Brotéria, Sér. Bot. 24: 104. 1930.

ORIGINAL DESCRIPTION — “Pileo 1 cm lato, campanulato, striato usque ad centrum,

membranaceo, viscido, centro brunneo-squamuloso, ceterum fibrilloso, margine hyalino,

lamellis confertis, linearibus, acie serratis, rubescentibus, liberis; stipites filiformi 1 mm.

lato, 3 cm. alto, vitreo, consperso; spora 4-5 4, subangulata, grosse guttulata.”

ComMEnts — Although Rick (1930, 1938) described P. sensitivus without

indicating a holotype, he later (Rick 1961) cited PACA20770 as the material

studied for this species. However, according to the exsiccata notes, PACA20770

represents an unpublished variety, “P. sensitivus var. macrospora” (see below);

moreover, as this specimen was collected in 1944, it cannot be the material on

which Rick (1930) based his original description. As no other collection has

been found for the epithet sensitivus, we regard this name as a nomen dubium.

“Pluteus sensitivus var. macrospora Rick; nom. in herb. FIG. 17

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Salvador, 1944, J. Rick s.n.

(PACA20770).

ComMENTS — See the description under P. murinus, with which it shares the

same features. Although PACA20770 was published under P sensitivus (Rick

1961), the specimen notes cite it as “P sensitivus var. macrospora.” Based on

218 ... Menolli & Capelari

FiGureE 17. “Pluteus sensitivus var. macrospora”. (PACA20770).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

the data recovered from the specimen, we suggest that it may actually represent

P. albostipitatus (see discussion under P. murinus).

Pluteus straminellus Rick, Iheringia, Sér. Bot. 8: 417. 1961.

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1930, J. Rick s.n.

(Holotype, PACA15531*).

ORIGINAL DESCRIPTION — “Pileo stramineo, innato-squamuloso, centro lurido, stipite

brevi, aquoso; lamellis densis, arcuatis, carneolis, liberis. Sporis 4 x 3 my [um]. Ad lignum.”

BASIDIOSPORES [20/1/1] 3.7(-4.4) x 2.5(-3.1) um [Q = (1.42-)1.48; Qm

= 1.48; Lm = 3.7 um; Wm = 2.5 um), ellipsoid, inamyloid, hyaline, smooth,

thick-walled, guttulate. PILEIPELLIS and HYMENIAL STRUCTURES not recovered.

CLAMP CONNECTIONS not observed.

ComMENTS — Based only on basidiospore size and shape, it is not possible to

establish P. straminellus as sufficiently well known, leading us to treat it as a

nomen dubium. If there were data about the pileipellis and hymenial structures

and pileus color, it might be possible to suggest a morphological relationship

between P straminellus and other small-spored species, such as P microsporus

(Dennis) Singer, P ugandensis Pegler, or P. unakensis Murrill.

Hennings & Rick Pluteus revisions (Brazil) ... 219

Pluteus termitum Henn., Hedwigia 43: 183. 1904, as “termitarum.”

ORIGINAL DESCRIPTION — “Pileo carnosulo, campanulato, vertice umbonato-

obtuso, brunneolo, radiatim substriato, albido, 3-4 cm diam., stipites fistuloso,

tereti, substriato, laevi basi interdum curvulo incrassato, pallido-brunneolo, 4-5 cm

longo, 2-3 mm crasso; lamellis liberis, confertis, ventrocsis, ca. 3 mm latis, pallidis

dein flavidis; sporis ellipsoideo-fusoideis utrinque acutiusculis, 1-2 guttulatis,

episporio pallide incarnate, laevi, 7-10 x 4-5 ys.”

COMMENTS — No specimens can be located. We therefore regard this name as

a nomen dubium.

“Pluteus umbrosus Pers.’ sensu Rick, Iheringia, Sér. Bot. 8: 418. 1961. Fic. 18

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Arroio do Meio, 1920, J. Rick s.n.

(BPI770905).

COMMENTS — See the description under P. cervinus var. griseoviridis, with which

it shares the same features. The name attributed by Rick (1919, 1938, 1961)

FIGURE 18. “Pluteus umbrosus” (BPI770905).

a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 um.

220 ... Menolli & Capelari

FIGURE 19. Pluteus velatus (holotype, PACA14527).

a. Basidiospores; b. Pleurocystidia. Scale bars = 10 um.

is obviously inaccurate because P umbrosus (Pers.) P. Kumm. represents sect.

Hispidoderma (Singer 1956, Vellinga & Schreurs 1985, Orton 1986) whereas

the above collection has the horned pleurocystidia typical of sect. Pluteus. Our

morphological examination indicates that Rick’s collection probably represents

P. angustisporus (see discussion under P cervinus var. griseoviridis).

Pluteus velatus Rick, Iheringia, Sér. Bot. 8: 417. 1961. FIG. 19

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1939, J. Rick s.n.

(Holotype, PACA14527* as P. velutinus).

ORIGINAL DESCRIPTION — “Campanulatus, 3 cm latus, griseo-brunneus, membranaceus,

squarrosulus ad marginem, secus nudus, fere usque ad centrum leviter striatus; lamellis

liberis, ventricosis, densis, albis, margine rubro. Basidiis brevibus, cystidiis paucis,

latis; stipite 6 cm alto, 3 cm lato, subtus crassiore, albo-lanoso, ceterum nudo, striatulo,

pallido, velo arachnoideo. Sporis 6-10 x 5-10 my [um], polygonalibus, rubris. Ad lignum

putridum.”

BASIDIOSPORES [20/1/1] (6.2-—)7.5-8.7(-10.0) x 5.0-6.2(-7.5) um [Q = (1.16-)

1.21-1.50(-1.61); Qm = 1.32; Lm = 7.9 um; Wm = 6.0 um), broadly ellipsoid to

Hennings & Rick Pluteus revisions (Brazil) ... 221

FiGuRE 20. Pluteus fibulatus (isotype, Singer T141, MICH).

a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 um.

ellipsoid or rarely almost elongate, inamyloid, hyaline, smooth, thick-walled,

guttulate. PILEIPELLIS composed of hyphal elements typical of a cutis but it is

too difficult to be recovered. PLEUROCYSTIDIA 56-69(-87) x (13.7—)18.7-24

(-30) um, clavate or slightly ventricose, thin-walled. CHEILOCYSTIDIA not

recovered. CLAMP CONNECTIONS not observed.

COMMENTS — Singer (1953, 1959) recognized Rick’s collection as representing

P fibulatus Singer (described from Argentina; Singer & Digilio 1952). Although

it is not clear whether Singer actually examined this collection, he did mention

that, “the indication of a veil in the unpublished notes must be due to an error.”

Pluteus velatus was published in Rick’s posthumous paper (Rick 1961), and

according to Singer (1953, 1959), P. velatus should be considered a synonym of

P. fibulatus. However, our re-examination of the two types confirms that they

represent different species.

Pluteus fibulatus is typical of sect. Pluteus, characterized by a fibrillose

deep-colored pileus, clamp-connections, and horned metuloid pleurocystidia

222 ... Menolli & Capelari

FIGuRE 21. Pluteus fibulatus (Pegler et al. 3899, K(M)41525).

a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 um.

(Singer 1959). Our re-examination of the isotype of P fibulatus [Argentina.

Prov. Tucuman: Capital, 22 Feb. 1949, Singer T141 (MICH)] (Fic. 20) showed

globose to broadly ellipsoidal (rarely ellipsoidal) basidiospores, [20/1/1] (5.6-)

6.2-7.5(-8.7) x (5.0—)5.6-6.8(-7.5) um [Q = 1.00-1.24(-1.40); Qm = 1.16; Lm

= 7.1 um; Wm = 6.1 um]; fusoid-ventricose pleurocystidia (65-88 x 17.5-28.7

um) with thickened to moderately thickened walls and apices with 2-4 apical

prongs (although sometimes without prongs and with an acute to rounded

apex), clavate cheilocystidia (44 x 13.7 um), a cutis-like pileipellis that was

difficult to recover, and difficult to detect clamp connections. Similar features

were also found in the collection of P. fibulatus (Fic. 21) recorded by Pegler

(1997) from the state of Sao Paulo, Brazil [Brazil. Sao Paulo: Mogi Guagu, 30

Jan. 1987, Pegler et al. 3899 (K(M)41525)].

However, globose basidiospores and metuloid or horned cystidia were not

found in our re-examination of the holotype of P. velatus. All pleurocystidia

observed in P velatus had thin walls, suggesting that the specimen represents

sect. Hispidoderma rather than P fibulatus or another species in sect. Pluteus.

Thus, although we can confirm the occurrence of P fibulatus in Brazil, the

Hennings & Rick Pluteus revisions (Brazil) ... 223

data recovered from Rick’s collection are not sufficient to confirm the identity

of P. velatus, which remains a nomen dubium. Taking into consideration the

macroscopic characteristics described by Rick (1961), such as pileus color and

lamellae with colored edges (‘margine rubro’), as well as the size and shape of the

basidiospores and pleurocystidia described herein, it is likely that the material

collected by Rick represents P fernandezianus Singer, originally described from

Chile (Singer 1959).

“Pluteus wehlianus Miiller? nom. in herb.

SPECIMEN EXAMINED — Brazil. Rio Grande do Sul: Sao Leopoldo, 1939, J. Rick s.n.

(PACA14529).

Basipiospores [4/1/1] 16.2-20 x 11.2-12.5 um (Q = 1.4-1.6; Qm = 1.51;

Lm = 18.4 um; Wm = 12.2 um), ellipsoid with germ pore. PILEIPELLIS &

HYMENIAL STRUCTURES not recovered. CLAMP CONNECTIONS not observed.

ComMENts — Although referred to P wehlianus (F. Muell.) Sacc., this

unpublished collection obviously does not represent a Pluteus species due to

the large basidiospores with germ pore. It is likely a species of Bolbitiaceae or

Strophariaceae.

Conclusion

Among the names published by Hennings and Rick, only three collections

referred to P. cervinus could be assigned to the correct name, P xylophilus.

Three other unpublished collections could be correctly identified: “P egregius”

as P xylophilus and “P murinus” and “P. sensitivus var. macrospora” as

P. albostipitatus. Materials were not found for ten taxa, of which we consider

five to be nomina dubia. We also regard P. straminellus and P. velatus as nomina

dubia because the morphological data recovered from Rick’s collections are

insufficient to confirm their identity.

Of the remainder, we only suggested another name for the collections of

Hennings and Rick when it was not possible to recover complete morphological

data for a correct identification, although the data available were sufficient to

reject the originally applied names. Finally, there is insufficient material of

“P. fuscidulus” for study, P leptonia belongs to Entolomataceae, and

“P. wehlianus” belongs to Bolbitiaceae or Strophariaceae.

Acknowledgments

The authors thank Andrew M. Minnis and Else C. Vellinga for critical review

and for providing very insightful comments in the manuscript; Shaun R. Pennycook

for his review and helpful suggestions regarding the nomenclature, formatting, and

presentation; Lorelei L. Norvell for her final review and editorial suggestions to improve

the text; all curators who helped with this work (especially those from BPI, FH, K,

224 ... Menolli & Capelari

LIL, MICH, PACA and SP) for the loan of collections; Klei R. Sousa for inking the

illustrations; Fernanda Karstedt for her assistance regarding the collections considered

members of Entolomataceae; the Graduate Program in “Biodiversidade Vegetal e Meio

Ambiente” for the financial support toward the costs involving this manuscript; and

the “Conselho Nacional de Desenvolvimento Cientifico e Tecnoldgico” (CNPq) for the

support and grant to the authors.

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Rick J. 1961. Basidiomycetes eubasidii in Rio Grande do Sul - Brasilia. 5. Agaricaceae. Iheringia,

Série Botanica 8: 301-449.

Ripkova S. 2009. Pluteus exiguus - a threatened or overlooked species? Catathelasma 11: 49-52.

Saccardo PA. 1887. Sylloge fungorum omnium hucusque cognitorum, vol. 5. Patavii.

Singer R, Digilio APL. 1952 [“1951”]. Prédomo de la flora Agaricina Argentina. Lilloa 25: 5-461.

Singer R. 1953. Type studies on Basidiomycetes VI. Lilloa 26: 57-159.

Singer R. 1956. Contributions towards a monograph of the genus Pluteus. Trans. Br. Mycol. Soc. 39:

145-232. http://dx.doi.org/10.1016/S0007-1536(56)80001-6

Singer R. 1959 [“1958”]. Monographs of South American Basidiomycetes, especially those of the

east slope of the east slope of the Andes and Brazil. 1. The genus Pluteus in South America.

Lloydia 21: 195-299.

Singer R. 1986. The Agaricales in modern taxonomy, 4th edn. Koeltz Scietific Books, Koenigstein.

Smith AH, Stuntz DE. 1958. Studies on the genus Pluteus I. Redescriptions of American species

based on a study of type specimens. Lloydia 21: 115-136.

226 ... Menolli & Capelari

Takehashi S, Kasuya T. 2007. First record of Pluteus chrysophaeus and reexamination of Pluteus

leoninus from Japan. Mycoscience 48: 321-325.

Thiers B. 2013 (continuously updated). Index Herbariorum: a global directory of public herbaria

and associated staff. New York Botanical Garden’ Virtual Herbarium. Accessed February 2013.

http://sweetgum. nybg.org/ih/.

Vellinga EC, Schreurs J. 1985. Notulae ad floram agaricinam neerlandicam - VIII Pluteus Fr. in

West-Europe. Persoonia 12: 337-373.

Vizzini A, Ercole E. 2011. A new annulate Pluteus variety from Italy. Mycologia 1034: 904-911.

http://dx.doi.org/10.3852/10-382

Wartchow F, Cortez VG, Coelho G. 2006. New records of Pluteus (Pluteaceae, Agaricales) from

Brazil. Mycotaxon 96: 241-252.

MY COTAXON

http://dx.doi.org/10.5248/126.227

Volume 126, pp. 227-230 October-December 2013

A new species of Bahusutrabeeja from Guangxi, China

XIAO-XIA LI”?’, JI-WEN X1A’, L1-Guo Ma’,

RAFAEL FE. CASTANEDA- RUIZ? & XIU-GUO ZHANG”

" Life Sciences Institute, Zunyi Normal College, Zunyi, Guizhou, 563002, China

* Department of Plant Pathology, Shandong Agricultural University,

Taian, Shandong, 271018, China

3 Instituto de Investigaciones Fundamentales en Agricultura Tropical “Alejandro de Humboldt”

(INIFAT), calle 1, esq. 2, Santiago de Las Vegas, Ciudad de La Habana, Cuba C. P. 17200

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

ABSTRACT —Bahusutrabeeja exappendiculata sp. nov. was discovered on rotten branches

from subtropical forest of Guangxi Province, China. It differs from previously described

Bahusutrabeeja species in having colourless globose conidia without appendages. The fungus

is described, illustrated and compared with similar taxa.

KEY worpDs — anamorphic fungi, taxonomy

Introduction

During ongoing surveys of saprobic microfungi from subtropical forests of

Guangxi Province, China, an interesting anamorphic fungus was collected on

rotten branches. Its conidiogenesis and conidia suggest that the fungus belongs

in Bahusutrabeeja Subram. & Bhat (Subramanian & Bhat 1977).

The type of this genus, B. dwaya Subram. & Bhat, has distinct, mononematous

conidiophores with integrated, terminal, cylindrical conidiogenous cells.

Conidiogenous cells produce conidia in succession by percurrent proliferation

from a single fertile locus. The conidia are colourless, smooth, thick walled,

aseptate, spherical or rounded-cubical or obpyriform to obclavate, with several

to many slender appendages distributed over the surface. They accumulate

in a slimy mass at the apex of the conidiogenous cell. The genera Nawawia

Marvanova (Marvanova 1980) and Chalarodes McKenzie (McKenzie 1991),

which are similar, were also considered, as they both have macronematous

conidiophores, conidiogenous cells each with a single fertile locus producing

appendaged conidia in succession by percurrent proliferation. The conidia are,

228 ... Li & al.

Fic. 1. Bahusutrabeeja exappendiculata. A. Colonies on natural substratum.

B. Globose conidia. C, D. Conidiophores with developing conidia.

however, pyramidal or tetrahedric in Nawawia, while in Chalarodes they are

obconical, with each bearing two distal setulae.

Worldwide, five species are currently included in Bahusutrabeeja. Those

species are separated by conidial shape, size, and appendages (Ma et al. 2011,

2012). All of those species occur on rotten twigs and branches. The present

fungus is morphologically distinct from all five previously known species and

is, therefore, described here as a new species. A key to all six species is provided.

Bahusutrabeeja exappendiculata Xiao X. Li & X.G. Zhang, sp. nov. FIG. 1

MycoBank MB 804166

Differs from all other Bahusutrabeeja species in having colourless, globose conidia

without appendages.

Bahusutrabeeja exappendiculata sp. nov. (China) ... 229

Type: China, Guangxi Province: Dayaoshan Nature Reserve, on rotten branches of an

unidentified broad-leaved tree, 10 Nov. 2012, Xiao X. Li (Holotype HSAUP H 9070;

isotype, HMAS 243429).

ETyMo_oey: exappendiculata, in reference to conidia without appendages.

Co.LoniEs on the natural substratum effuse, dark brown to blackish, velvety,

with scattered conidiophores visible under the stereomicroscope, each with

a colourless, globose conidial mass at the tip. Mycelium partly superficial,

partly immersed in the substratum, composed of almost colourless to brown,

septate, branched hyphae. CoNIDIOPHORES mononematous, erect, straight or

slightly flexuous, occasionally branched, cylindrical, smooth, thick-walled, 6-

to 14-septate, 161-275 x 5-6.5 um, dark reddish brown to dark brown at the

base which is up to 11.5 um wide, pale brown to brown at the apex, with a

single conidiogenous cell at the apex, often proliferating percurrently through

the collarette of that conidiogenous cell, and produce further growth with

additional septa, and eventually a new conidiogenous cell. CONIDIOGENOUS

CELLS terminal, integrated, cylindrical, slightly swollen toward the belly, up to

7.5 um wide, producing conidia by percurrent non-progressive proliferation

from a single fertile locus at the apex where there is a prominent, slightly

narrower collarette, 3-4 um wide. Conip1A colourless, globose, thick-walled,

aseptate, 10.5-16 um diam., without appendages, aggregating in a slimy

colourless mass at the apex of the conidiogenous cell.

ComMENTS — Bahusutrabeeja exappendiculata differs from all previously

described species in the genus in producing conidia with no appendages. The

new species resembles B. dwaya (Subramanian & Bhat 1977), B. globosa Bhat

& W.B. Kendr. (Bhat & Kendrick 1993), and B. bunyensis McKenzie (McKenzie

1997) in conidial shape and pigmentation. The conidia of B. exappendiculata

are, however, clearly larger than those of B. bunyensis (7.5-10.5 x 7.5-9.5 um),

and smaller than those of B. globosa (18-22 um). In B. dwaya, the first-formed

conidium is pear-shaped while subsequent conidia are spherical.

Key to described Bahusutrabeeja species

Lem, Conidivnotspheticale s2).58 ve. she Asa seeks ee AS aah aoe eee gates as 2

Conidiaspherical or subsphericaly po So sess be bytes. wh a,c te Re 3

2.— k@onidiaanowlar, ‘7S! Uni iaiiis ee plein pete cde Aa lan de Send B. angularis

Conidia obpyriform to obclavate, 15-20 x 5-8 um ............... B. dubhashii

3. Conidia without appendages, 10.5-16 um diam. ........... B. exappendiculata

Monidia Withrappendaces®, ot. .h labs at tat ohh hee ha Rie al ge at 8 +

4. Conidia with 3 appendages, 7.5-10.5 x 7.5-9.5 um ............00. B. bunyensis

Cotidra with: 8 12-dppendages Fi se. na ee ia tase tien aceathis nutaty ae eases toes 5

bee MEG Onidia 12.514 rm, AAT oo. og dking dihnieg diated disatercd dineerg diners Libeehe B. dwaya

Conidia US-2 9s evel alii cs. kee dale tet ale apelin eect aia d on slaaa dat ladle dl B. globosa

230 ... Li & al.

Acknowledgments

The authors thank Dr De-Wei Li and Dr David Minter 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).

Literature cited

Bhat DJ, Kendrick B. 1993. Twenty-five new conidial fungi from the Western Ghats and the

Andaman Islands (India). Mycotaxon 49(1): 19-90.

Ma J, Wang Y, Ma LG, Zhang YD, Castafieda RF, Zhang XG. 2011. Three new species of

Neosporidesmium from Hainan, China. Mycol. Prog. 10: 157-162.

http://dx.doi.org/10.1007/s11557-010-0685-2

Ma LG, Ma J, Zhang YD, Zhang XG. 2012. A new Corynesporella species and two first records from

China. Mycotaxon 119: 83-88. http://dx.doi.org/10.5248/119.83

Marvanova L. 1980. New or noteworthy aquatic hyphomycetes. Clavatospora, Heliscella, Nawawia

and Heliscina. Trans. Br. Mycol. Soc. 75(2): 221-231.

http://dx.doi.org/10.1016/S0007-1536(80)80083-0

McKenzie EHC. 1991. Dematiaceous hyphomycetes on Freycinetia (Pandanaceae). 3. Chalarodes

gen. nov. Mycotaxon 42: 89-93.

McKenzie EHC. 1997. Bahusutrabeeja bunyensis sp. nov. (hyphomycetes) from Queensland,

Australia, and a new name for Chalara australis McKenzie. Mycotaxon 61: 303-306.

Subramanian CV, Bhat DJ. 1977. Bahusutrabeeja, a new genus of the hyphomycetes. Can. J. Bot.

55(16): 2202-2206. http://dx.doi.org/10.1139/b77-249

MYCOTAXON

http://dx.doi.org/10.5248/126.231

Volume 126, pp. 231-233 October-December 2013

Flavophlebia sphaerospora,

a new corticoid species from India

MANINDER Kaur, AVNEET P. SINGH * & G.S. DHINGRA

Department of Botany, Punjabi University, Patiala 147002, India

*CORRESPONDENCE TO: avneetbot@gmail.com

ABSTRACT — Flavophlebia sphaerospora sp. nov. is described from Himachal Pradesh, India.

Key worps — Basidiomycota, Northwestern Himalaya, Narkanda

During the fungal forays conducted during August 2012 in the Hattu Peak area,

Narkanda, district Shimla of Himachal Pradesh, India, Maninder and Avneet

collected a fungus on decaying wood of Cedrus deodara in a mixed forest.

After comparison of macroscopic and microscopic characters (Parmasto 1968,

Hjortstam & Larsson 1977, Bernicchia & Gorjén 2010, Mycobank 2013) we

concluded that the material represented the monotypic genus Flavophlebia but

differed from E sulfureoisabellina (Litsch.) K.H. Larss. & Hjortstam.

Flavophlebia sphaerospora Man. Kaur, Avneet P. Singh & Dhingra, sp. nov. PLATE 1

MycoBank 804821

Differs from Flavophlebia sulfureoisabellina by its denser subiculum and its spherical to

subspherical basidiospores.

Tye: India, Himachal Pradesh: Shimla, Narkanda, on way to Hattu Peak, on the bark of

decaying wood of Cedrus deodara (Roxb. ex D. Don) G. Don, 19 August 2012, Maninder

Kaur & Avneet 5166 (PUN, holotype).

Erymo toy: The epithet refers to the shape of the basidiospores.

Basidiocarp resupinate, adnate, effused, ceraceous, <180 um thick in section;

hymenial surface smooth, grayish green to grayish yellow to grayish orange;

margins thinning, paler concolorous, to indeterminate. Hyphal system

monomitic; generative hyphae branched, septate, clamped; basal hyphae

<3.3 um wide, more or less parallel to the substrate, thick-walled, forming a

dense subiculum; subhymenial hyphae <2.7 um wide, vertical, densely united.

232 ... Kaur, Singh & Dhingra

PLATE 1. Flavophlebia sphaerospora (holotype). 1. Basidiocarp showing hymenial surface. 2.

Vertical section through basidiocarp showing hyphae, basidia, cystidia and basidiospores.

Flavophlebia sphaerospora sp. nov. (India) ... 233

Cystidia 45-60 x 8-8.5 um, cylindrical, flexuous, thin-walled, with basal clamp.

Basidia 28-36 x 8.5-10 um, clavate, somewhat stalked, 4-sterigmate, with basal

clamp, filled with oily contents; sterigmata <6.0 um long. Basidiospores 6.6-8.0

um in diameter, spherical to subspherical with subapical apiculus reminding of

Radulomyces confluens (Fr.) M.P. Christ., with numerous oil-droplets, thin- to

somewhat thick-walled, inamyloid, somewhat cyanophilous.

REMARKS— Both Flavophlebia sphaerospora and F. sulfureoisabellina occur on

the bark of coniferous trees and have a similar greenish-yellowish colour, but F.

sulfureoisabellina differs in having a comparatively loose subiculum, ellipsoid

to subglobose basidiospores, and slightly narrower basidia.

Acknowledgements

The authors thank Head (Department of Botany, Punjabi University, Patiala)

for providing research facilities, Dr. Nils Hallenberg (Professor Emeritus in Botany,

Goteborg, Sweden) for peer review and expert comments, and Prof. B.M. Sharma

(Department of Plant Pathology, COA, CSKHPAU, Palampur, H.P., India) for peer

review.

Literature cited

Bernicchia A. & Gorjon SP. 2010. Corticiaceae s.1. Fungi Europaei 12. Edizioni Candusso. Alassio.

Italia. 1008 p.

Hjortstam K. & Larsson, KH. 1977. Notes on Corticiaceae (Basidiomycetes). Mycotaxon 5(2):

475-480.

MycoBank. 2013. Fungal databases. Nomenclature and species banks. [Accessed: 11/07/2013]

http://www.mycobank.org/

Parmasto E. 1968. Conspectus Systematis Corticiacearum. Tartu. 262 p.

MYCOTAXON

http://dx.doi.org/10.5248/126.235

Volume 126, pp. 235-237 October-December 2013

Peniophora hallenbergii sp. nov. from India

SAMITA & G.S. DHINGRA*

Department of Botany, Punjabi University, Patiala 147 002, India

*CORRESPONDENCE TO: dhingragurpaul@gmail.com

ABSTRACT — A new corticioid species, Peniophora hallenbergii, is described on a stick of Rosa

indica from Uttarakhand state in India.

Key worps - Basidiomycota, Agaricomycetes, Chaurangi Khal, Uttarkashi

While conducting fungal forays in Chaurangi Khal area of district Uttarkashi,

Uttarakhand (India), Samita collected an unknown corticioid fungus on a stick

of Rosa indica. The presence of gloeocystidia, metuloids, and smooth inamyloid

basidiospores indicates that the material belongs to genus Peniophora (Rattan

1977, Eriksson et al. 1978, Boidin et al. 1991, Dhingra 1993, Boidin 1994,

Wu 2002, Bernicchia & Gorjén 2010). The material, which keys out near

P. boidinii but from which it differs in basidiospore shape, is described here as

a new species. A portion of the basidiocarp was sent to Prof. Nils Hallenberg

(Sweden), who confirmed the findings.

Peniophora hallenbergii Samita & Dhingra sp. nov. Fries 1-9

MycoBAank 804960

Differs from Peniophora boidinii by its broadly ellipsoid basidiospores.

Type: India, Uttarakhand: Uttarkashi, Chaurangi Khal, on a stick of Rosa indica L., 29

September 2011, Samita 5167 (PUN, holotype).

Erymo oey: In honor of Nils Hallenberg, Professor Emeritus, University of Gothenburg,

Sweden.

Basidiocarps resupinate, adnate, effused, <180 um thick in section, hymenial

surface smooth, grayish orange; margins thinning, fibrillose, paler concolorous

to whitish. Hyphal system monomitic; generative hyphae <4.5 um wide,

branched, septate, clamped; basal hyphae parallel to the substrate, thin- to

somewhat thick-walled, subhyaline to pale brown; subhymenial hyphae

vertical, subhyaline, compactly arranged. Gloeocystidia 25-46 x 5-8.5 um, few,

236 ... Samita & Dhingra

OAV

terre ten CH

sas

ret

SOYA

ifs

9 o

ai — ‘>

OTE}

PLATE 1. Peniophora hallenbergii (holotype). 1. Basidiocarp showing hymenial surface.

2. Basidiospores. 3. Basidia. 4. Hyphae. 5. Gloeocystidium. 6. Metuloid 7. Vertical

section through basidiocarp. 8. Cystidia. 9. Basidia.

subfusiform, slightly thick-walled, negative to sulphovanillin. Metuloids 30-58

x 8-15 um, abundant, conical to subfusiform, heavily encrusted, thick-walled,

with basal clamp. Basidia 28-37 x 3.5-5 um, clavate, 4-sterigmate, with basal

clamp; sterigmata <4.5 um long. Basidiospores 6.3-8 x 4.5-5.6 um, broadly

ellipsoid, inamyloid, acyanophilous.

Peniophora hallenbergii sp. nov. (India) ... 237

ADDITIONAL MATERIAL STUDIED: India, Uttarakhand: Dehradun, Chakrata, on angio-

spermous wood, 18 September 2012, Samita 5168 (PUN).

REMARKS— Peniophora hallenbergii resembles P. boidinii D.A. Reid (described

from southwestern Europe) in having a similar basidiocarp morphology and

two kinds of cystidia. However, P. boidinii differs in having narrowly ellipsoid

to reniform or suballantoid basidiospores (6-9 x 3-3.8 tm).

Acknowledgements

The authors thank Head of the Department of Botany (Punjabi University, Patiala)

for providing research facilities, Prof. Nils Hallenberg (Gothenburg, Sweden) for expert

comments and peer review, and Prof. B.M. Sharma (Department of Plant Pathology,

COA, CSKHPAU, Palampur, H.P., India) for peer review.

Literature cited

Boidin J. 1994. Les Peniophoraceae des parties tempérées et froides de lhémisphére nord

(Basidiomycotina). Bull. Mens. Soc. Linn. Lyon 63(9): 317-334.

Boidin J, Lanquetin P, Gilles G. 1991. Les Peniophoraceae de la zone intertropicale (Basidiomycetes,

Aphyllophorales). A. Espéces paleotropicales. Bull. Soc. Mycol. France 107: 91-147.

Bernicchia A, Gorjon SP. 2010. Corticiaceae s.1. Fungi Europaei 12. Edizioni Candusso. Alassio.

Italia. 1008 p.

Dhingra GS. 1993. Three new records of genus Peniophora Cooke from Eastern Himalayas.

Geobios New Reports 12: 101-104.

Eriksson J, Hjortstam K, Ryvarden L. 1978. The Corticiaceae of North Europe-V. Oslo. pp.

890-1046.

Rattan SS. 1977. The resupinate Aphyllophorales of the North Western Himalayas. Bibliotheca

Mycologica 60: 1-427.

Wu SH. 2002. A study of Peniophora species in Taiwan with clamped hyphae. Bot. Bull. Acad. Sin.

43: 241-250.

MYCOTAXON

http://dx.doi.org/10.5248/126.239

Volume 126, pp. 239-245 October-December 2013

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

It is a pleasure to compile this present set of reviews of recently published

books. Two important European contributions to our knowledge of genera in

the Agaricales came out, a thorough monograph of Rosellinia saw the light,

and the book announcements show the results of many more systematic and

taxonomic activities concerning various groups of fungi in different places of

the world. The book that catches the eye and most attention is THE KINGDOM

OF FUNGI by J.H. Petersen, as it is par excellence the way to bring fungi to the

attention of everyone.

ASCOMYCOTA

Atlas of soil ascomycetes. By J. Guarro, J. Gene, A.M. Stchigel and M.J. Figueras.

2012. CBS Biodiversity Series 10. CBS-KNAW Fungal Biodiversity Centre, P.O. Box

85176, 3508 AD Utrecht, The Netherlands. <info@cbs.knaw.nl>. Pp. 486, 322 figs.

Price 70 €.

Soil fungi are ubiquitous and are found in great numbers in every environmental

study focusing on fungi, but their identities often remain obscure.

This atlas gives an overview of the ascomycetes found in soil that make

ascomata in culture, and it provides identification tools for this specific group

of fungi.

Keys to genera and species, descriptions, and illustrations (photographs and

beautifully executed line drawings) will help with the identification process.

‘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.

240 ... Vellinga, BOOK REVIEW EDITOR

Different culturing techniques and observation methods, and identification

criteria are elaborated on.

The genera are presented in alphabetical order, as are the species within each

genus.

The authors realize both that the classifications of the treated groups are

very much in flux and that these identification methods may soon be replaced

by new generation sequencing. Nevertheless, the morphological information

presented here is valuable and might give clues to the diverse life styles of

these fungi and their environments. It might also stimulate future students and

studies in the field. This non-ascomycete-specialist, who spent a pleasurable

hour browsing through this volume, certainly hopes that the work will find

its place in the hands of many soil biologists and other environment-oriented

mycologists.

Rosellinia -a world monograph. By L.E. Petrini, 2013. BrsLioTHECA MYCOLOGICA

205. Schweizerbart science publishers, Johannesstr. 3A, D-70176 Stuttgart, Germany,

< www.schweizerbart.de>. ISBN 978-3-443-59107-6. Pp 410, figs 72. Price 119 €

Rosellinia necatrix and its close relatives are important root pathogens of

many woody food plants, but the rest of the genus Rosellinia has sparsely been

investigated. Liliane Petrini’s monograph has changed that dramatically.

The work presented here is an old-school morphological study of

herbarium specimens from all over the world. Rosellinia fruitbodies are

small, inconspicuous, not abundant, and hard to find. The fruitbodies form in

stromata on a subiculum; the spores often have appendages or are embedded

in a mucous sheath.

A short introduction to the genus and its history, generic and specific

delimitation criteria, and a short methodology overview precedes the real part

of the book, comprising keys and species descriptions. Species descriptions

follow a fixed format, so the descriptions are easily compared; line drawings and

photos are included, and keys to species groups and species are dichotomous

and arranged by group. A total of 142 species (including 37 new taxa) is treated.

The species concept is narrow, and subtle characters in stromatal and spore sizes

are used. Some of the new taxa are based on only one collection. Very valuable

are the lists of names that are not covered in this work as well as the chapters on

geography and ecology of the species. The briefness of the chapter on ecology is

an indication of the lack of knowledge on this group. Also included are country

and host species lists (each organized by both main categories). The numerous

indices make this a very useful book.

So far Rosellinia has not undergone phylogenetic studies, but with this

monograph and the morphological basis laid, it should be much more feasible.

MycotTaxon 126 Book Reviews ... 241

Sequence data of endophytes could then also be linked to named species. I also

hope that more attention will be paid to the ecology and pathogenicity of the

species.

Dr. Petrini must be congratulated on the completion of a work certain to

become a standard, and I am looking forward to seeing other monographs on

equally intriguing and understudied genera in the future.

BASIDIOMYCOTA

Agaricus L. - Allopsalliota (Parte II). By L.A. Parra Sanchez. 2013. Funer

Europazl 1A, 2° Ed. Candusso Edizioni, Via Ottone Primo 90, I-17021 Alassio

(SV), Italy. ISBN 978-88-905310-2-6. Pp 1168, 679 coloured Pl. 89 €

The second part of the volume in the series FuNGr EUROPAE! on the genus

Agaricus came out in 2013, 5 years after part 1. Luis Alberto Parra Sanchez

and colleagues have produced a tome of 1168 pages, in which the European

species of sections Xanthodermatei, Arvenses, Minores, and Lanosi (in subg.

Lanagaricus) plus the genus Allopsalliota are treated, a total of 56 species.

The format follows that of Part I. Text is in both Spanish and English, and

the keys have also been translated into Italian. The descriptions precede 150

pages of illustrations, mainly colour photos of fruitbodies and microscopical

characters; multiple entries per species illustrate variability according to age

and circumstance. There is a section with published and unpublished older

plates, two spore size scatter diagrams, and a number of ITS sequence-based

phylogenetic trees showing species relationships and justification for describing

some of the new species. Bibliographic references and addenda to the first

volume complete this work.

Twelve new species are proposed. Several were only discovered through

ITS sequence differences while others are also morphologically supported.

Taxonomic confusion from the past is cleared up (as in the case of Agaricus

semotus and its various interpretations throughout time), and four species are

now lectotypified.

It is a pleasure to see Rick Kerrigan, the North American Agaricus specialist,

honoured with a species named for him, along with Jozsef Geml (now in the

Netherlands) and Jacques Guinberteau (France). The author also named a

species in honour of his wife, and the names of his two children are given to yet

another species.

Many more taxa and names than the ones described in full are discussed

in the text, and this work is truly a must for every one interested in the group.

It also serves as an example of thorough work on a very difficult and daunting

242 ... Vellinga, BOOK REVIEW EDITOR

group. The combination of morphological and phylogenetic work is great, and

I now await even more eagerly Kerrigan's North American Agaricus book that

will also combine both sets of characters.

Parra Sanchez is not a professional mycologist in the sense of someone who

earns a living by doing mycological research. But the book reflects collaboration

with many others in the field (many of the new species are group efforts), and

the translations and editing have been done by again another set of people.

It amazes me to see some western North American species present in Europe

and vice versa, and I would love to see a biogeographic study of the genus and

speculations on migration routes and centres of diversity.

I have one minor bone to pick, which has more to do with the formatting

and layout than with the contents. I really miss page headings indicating which

species is treated on that particular page; the species entries are long, and one

gets easily lost in the sea of words and information.

In short: highly recommended, but because of its weight and bulk definitely

not a book for the field.

The genus Tricholoma. By M. Christensen & J. Heilmann-Clausen. 2013.

FUNGI OF NoRTHERN Europe VOL. 4. The Danish Mycological Society. Svampetryk,

<www.svampe.dk; svampetryk@svampe.dk>. ISBN 978-87-983581-8-3. 228 pp,

numerous colour pl. Price 300 DKK. Also available in Danish.

A new book in the series FUNGI OF NORTHERN EuropPE is always a happy event,

and this installment does not disappoint.

Tricholoma is the subject, and the authors give a thorough introduction

to the genus, including a preliminary tree based on ITS sequences with

the promise of a more in-depth article, information on the ecology of the

species and where to find species assemblages, and some notes on edibility.

The main part presents keys, descriptions, and illustrations of all the treated

species. Following are lists of collections on which the descriptions are based,

geographical distribution data, interpretation of the colour illustrations in the

literature, colour references, and literature references. For a number of species,

neo- or epitypes have been selected, which are also elaborated on at the end.

Though the title may suggest that only northern Europe is taken into account,

the authors have traveled all throughout Europe to gain insight into species

diversity, variability, and distribution.

Tricholoma species lack interesting microscopical characters, and

identification is for the most part based on the macromorphology: colour and

shape of the fruitbodies, the characters of the veil, and the structure of the

pileipellis. The keys handle these characters well. The descriptions are good,

and the colour illustrations are of high quality; for most species, only one

MycotTaxon 126 Book Reviews ... 243

photo is given, but for some highly variable taxa (e.g., Tricholoma saponaceum),

the range of variation is illustrated. For each species, spore drawings and a

distribution map are given. Short discussions point to comparisons with similar

looking species, problems in the literature etc. It is clear that some names have

been misapplied in North America.

This is a beautiful and very useful book, not only within Europe, but also

outside the continent. I hope that this, together with the North American

Tricholoma book (Bessette et al. 2013), will stimulate the study of this important

ectomycorrhizal genus, which includes one of the most highly valued edible

taxa in the world.

Bessette AE, Bessette AR, Roody WC, Trudell SA. 2013. Tricholoma of North

America. A mushroom field guide. Austin, University of Texas Press. 208 pp.

GENERAL MYCOLOGY

Texas mushrooms. A field guide. By S. Metzler & V. Metzler, 1992; 2013.

University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, U.S.A.

<www.utexaspress.com>. ISBN: 978-0-292-75126-2. Pp. 358, numerous colour pl.

Price US$ 23.42 (web)

This is a reprint of the 1992 edition with a new preface, in which the revolutions

in mushroom classification of the last 20 years are mentioned. The new

classifications resulting from in-depth phylogenetic research show that species

first considered closely related because of similarities in morphology (think

Coprinus comatus and C. lagopus, both of which autodigest their lamellae) are

now placed in different genera, families, or orders (for the ink cap example

above, the genera are now Coprinus in the Agaricaceae and Coprinopsis in

the Psathyrellaceae). However, as the nomenclature in the book has not been

updated according to the newly gained insights, the reader is encouraged to

search the Internet and other resources to do so herself.

The quality of the photos, a very important component of a field guide,

varies considerably: some pictures are too dark and too blue; others—such as

the close-up of Phyllotopsis nidulans—are beautiful. Some species are clearly

misidentified (e.g., the photo of Laccaria amethystina shows Mycena pura).

And as usual in books like this, the source of descriptions and photos is not

revealed.

So far, this is still the only popular book dealing with Texas mushroom

species, but I sincerely hope that someone will post updates on the names and a

critical evaluation of the species online for all to use, and that future books will

reflect our current knowledge and insights.

244 ... Vellinga, BOOK REVIEW EDITOR

The Kingdom of Fungi. By J.H. Petersen. 2013. Princeton University Press, 41

William Street, Princeton, New Jersey 08540 U.S.A. <orders@cpfsinc.com>. ISBN

978-0-691-1574-2. Pp 272, 800+ colour pl. Price US $ 29.95

Also available in other languages (Dutch as ‘Het leven van paddenstoelen &

schimmels, Danish as ‘I svampenes rige, and Estonian as ‘Seeneriigi illustreeritud

entsiiklopeedia’), and as an e-book (eISBN: 978-1-4008-4687-0).

One of the most exciting events for mycology in 2013 was the publication of

Jens Petersen’s book, THE KINGDOM OF FUNGI. It is a book full of beautiful,

stunning, and insightful pictures of fungi and mushrooms, interspersed with

short pieces of text. But all are so arranged and crafted that in a nutshell a

thorough introduction to the kingdom is given and various evolutionary trends

are made clear. The book also introduces fungal ecology and the different

habitats where these organisms can be found. Mushroom-forming fungi in

the Basidiomycota and Ascomycota (including lichenized species) get the most

attention, but other groups feature as well.

It is not a guidebook to identify mushrooms in a certain area; there are

photos from many different places, in Europe, Asia, Africa, and South America.

The microphotos of asci and basidia, the close-ups of pores, lamellae, spines,

and many other details make you look at the fungal world with different eyes.

The book is above all destined to live on your living room or office table, laid

open at one of the detailed photos to convey the message that mushrooms are

varied, intriguing, diverse, and still full of mystery for the human viewer.

The low price makes it affordable for many, and if you have not yet bought

it, please do so soon, and give it to your students, prospective students, and

anybody with the slightest interest in the natural world.

BOOK ANNOUNCEMENTS

Cultivation and diseases of Proteaceae: Leucadendron, Leucospermum and Protea.

By P.W. Crous, S. Denman, J.E. Taylor, L. Swart, C.M. Bezuidenhout, L. Hoffman,

M.E. Palm & J.Z. Groenwald. 2013 CBS BIODIVERSITY SERIES 13. CBS-KNAW

Fungal Biodiversity Centre, P.O. Box 85176, 3508 AD Utrecht, The Netherlands.

<info@cbs.knaw.nl>. Pp. 360. ISBN: 978-90-70351-95-3 Print ISSN: 1571-8859.

Pricew5-4i

Flora of the Guianas Series E: (fungi and lichens) Cladoniaceae. By S. Mota de Oliveira.

2013. Kew Publishing <www.kewbooks.com>. ISBN 9781842464793. 150 pp, 60

B&W pl. Price US$ 99.00.

Operculate Discomycetes (Pezizales, Ascomycota) of Israel. By G.S. Barseghyan &

S.P. Wasser. 2013. BIODIVERSITY OF CYANOPROKARYOTES, ALGAE AND FUNGI OF

MycoTaxon 126 Book Reviews ... 245

IsRAEL. Koeltz Scientific Books, P.O.Box 1360, D-61453 Koenigstein, Germany.

<www.koeltz.com>. ISBN 978-3-87429-442-3.Pp 240, 11 figs. Price 118 €

Ophiostomatoid fungi. By K.A. Seifert, Z.W. de Beer & M.J. Wingfield. 2013 CBS

BIODIVERSITY SERIES 12. CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85176,

3508 AD Utrecht, The Netherlands. <info@cbs.knaw.nl>. Pp. 337. ISBN: 978-90-

70351-94-6. Price 75 €.

Phytopathogenic Dothideomycetes. By P.W. Crous, G.J.M. Verkley & J.Z. Groenewald

(eds). 2013. SrupiEs In Mycotoey no. 75. CBS-KNAW Fungal Biodiversity Centre,

P.O. Box 85176, 3508 AD Utrecht, The Netherlands. <info@cbs.knaw.nl>. Pp. 406,

illustr. Price 70 € (paper copy, download free)

Plant pathogenic and endophytic Botryosphaeriales known from culture. By A.J.L.

Phillips, B. Slippers, J.Z. Groenewald & P.W. Crous (eds). 2013. STUDIES IN

Myco.oey no. 76. CBS-KNAW Fungal Biodiversity Centre, PO. Box 85176, 3508

AD Utrecht, The Netherlands. <info@cbs.knaw.nl>. Pp. 167, illustr. Price 65 € (paper

copy, download free)

The lichens and allied fungi of Great Smoky Mountains National Park. An annotated

checklist with comprehensive keys. By J.C. Lendemer, R.C. Harris & E.A. Tripp.

2013. MEMoIRS OF THE NEW YORK BOTANICAL GARDEN 104. The New York

Botanical Garden Press, 2900 Southern Boulevard, Bronx, NY 10458-5126, U.S.A.

<nybgpress@nybg.org>. 164 pp. ISBN 978-0-89327-521-1. Price US$55.00.

MYCOTAXON

http://dx.doi.org/10.5248/126.247

Volume 126, pp. 247-248 October-December 2013

Regional annotated mycobiotas

new to the Mycotaxon website

ABSTRACT — MycotTaxon is pleased to announce three new species distribution lists to

our ‘web-list’ page covering Cortinariaceae in Argentina (by Romano & Lechner), lichens

in Iran (by Valadbeigi), and lichenized and lichenicolous fungi in Spain’s Balearic Islands

(by Atienza & al.). The 6" edition of the myxomycetes and larger asco/basidiomycetes in

Turkey by Sesli & Denchev has also been uploaded. This brings to 110 the number of free

access mycobiotas now available on the MycoTaxon website:

http://www.mycotaxon.com/resources/weblists.html

SOUTH AMERICA

Argentina

G.M. ROMANO & B.E. LECHNER. The Cortinariaceae of Argentina's Nothofagus

forests. 35 p.

ABSTRACT — We present a checklist of species in the Cortinariaceae (Agaricales)

described from the Nothofagus forests of Argentina up to 2013. Names of the 247

species in Cortinarius, Dermocybe, Descolea, and Stephanopus have been revised

and are accompanied by references and updated distributions.

EUROPE

Spain

VIOLETA ATIENZA, ELENA ARAUJO, MARIA DEL ROSARIO ARROYO, ANA

Rosa BURGAZ, GEMMA FIGUERAS, ANTONIO GOMEZ-BOLEA, NESTOR L.

HLADUN, XAVIER LLIMONA, ESTEVE LLop, MARiA EUGENIA LOPEZ DE

SILANES, BERNARDA MARCOS, ISRAEL PEREZ-VARGAS, RAQUEL PINO-

Bopas, EsTELA SERINA & VicToR J. Rico. Lichenized and lichenicolous

fungi from the Pititises Archipelago (Eivissa and Formentera Islands and

Islets), Balearic Islands, Spain. 36 p.

AsstTract — A preliminary checklist for the lichenized and lichenicolous fungi

from the Pititises Archipelago in the Balearic Islands is presented, based on original

records complemented with literature references. The catalogue of our collections

in the Pitiiises Archipelago contains 302 lichens and lichenicolous fungi. To date,

the checklist, together with records from the literature, include 360 lichens and

248 ... New regional mycobiotas online

lichenicolous fungi, representing 113 genera. Muellerella lecanactidis is a new

record for Europe, 6 lichens and lichenicolous fungi are new for Spain and the

Iberian Peninsula, 57 for the Balearic Islands and 116 for the Pititises Archipelago.

Mip-EASstT

Iran

TAHEREH VALADBEIGI. Lichen flora of the Ilam Province, South West Iran. 7 p.

ABSTRACT — 121 lichen species are recorded as new to the Ilam province, and four

species, Candelariella rosulans, Lecanora sulphurata, Lecidella scabra, and Lecanora

klauskalbii, are new to Iran. Two newly reported species, Verrucaria macrostoma

and Xanthoria candelaria, have not been seen since 1957.

MY COTAXON

http://dx.doi.org/10.5248/126.249

Volume 126, pp. 249-251 October-December 2013

NOMENCLATURAL NOVELTIES AND TYPIFICATIONS

PROPOSED IN MYCOTAXON 126

Ambomucor R.Y. Zheng & X.Y. Liu, p. 99

Ambomucor clavatus R.Y. Zheng & X.Y. Liu, p. 105

Ambomucor seratioinflatus R.Y. Zheng & X.Y. Liu, p. 99

Ambomucor seratioinflatus var. brevior R.Y. Zheng & X.Y. Liu, p. 102

Annulohypoxylon orientale Lar.N. Vassiljeva & S.L. Stephenson, p. 5

Bahusutrabeeja exappendiculata Xiao X. Li & X.G. Zhang, p. 228

Craterium corniculatum B. Zhang & Yu Li, p. 72

Embryonispora G.Z. Zhao, p. 78

Embryonispora bambusicola G.Z. Zhao, p. 78

Entocybe haastii (G. Stev.) Largent, p. 64

Flavophlebia sphaerospora Man. Kaur, Avneet P. Singh & Dhingra, p. 231

Hypoxylon cyanescens H.X. Ma, Lar.N. Vassiljeva & Yu Li, p. 7

Linodochium sinense D.W. Li, Jing Y. Chen & Yi X. Wang, p. 16

Lophodermium circinatum Li Chen bis & Y.R. Lin, p. 110

Nectria araliae Lar.N. Vassiljeva & S.L. Stephenson, p. 8

Nipponoparmelia perplicata S.Y. Kondr., Tschab., Elix & Hur, p. 38

Passalora lepistemonis L. Xia, Y.L. Guo & Y. Li, p. 51

Peniophora hallenbergii Samita & Dhingra, p. 235

Peziza oliviae J.L. Frank, p. 185

Phyllobaeis crustacea S.N. Cao & J.C. Wei, p. 35

Phytophthora virginiana Xiao Yang & C.X. Hong, p. 171

Spadicoides bawanglingensis J.W. Xia & X.G. Zhang, p. 55

Spadicoides longchiensis J.W. Xia & X.G. Zhang, p. 57

Terriera nitens Y.R. Lin, p. 114

Trichoderma albocorneum (Yoshim. Doi) Jaklitsch & Voglmayr, p. 145

Trichoderma albofulvum (Berk. & Broome) Jaklitsch & Voglmayr, p. 145

Trichoderma alcalifuscescens (Overton) Jaklitsch & Voglmayr, p. 145

Trichoderma americanum (Canham) Jaklitsch & Voglmayr, p. 145

250 ... MYCOTAXON 126

Trichoderma andinense (Samuels & Petrini) Samuels, Jaklitsch & Voglmayr, p. 146

Trichoderma atrogelatinosum (Dingley) Jaklitsch & Voglmayr, p. 146

Trichoderma avellaneum (Rogerson & S.T. Carey) Jaklitsch & Voglmayr, p. 146

Trichoderma britdaniae (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, p. 146

Trichoderma caerulescens (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, p. 146

Trichoderma citrinum (Pers. : Fr.) Jaklitsch, W. Gams & Voglmayr (epitypified),

p. 147

Trichoderma corneum (Pat.) Jaklitsch & Voglmayr, p. 147

Trichoderma costaricense (P. Chaverri & Samuels) P. Chaverri, Jaklitsch &

Voglmayr, p. 147

Trichoderma danicum (Jaklitsch) Jaklitsch & Voglmayr, p. 148

Trichoderma decipiens (Jaklitsch, K. Poldmaa & Samuels) Jaklitsch &

Voglmayr, p. 148

Trichoderma eucorticioides (Overton) Jaklitsch & Voglmayr, p. 148

Trichoderma flaviconidium (P. Chaverri, Druzhin. & Samuels) Jaklitsch &

Voglmayr, p. 148

Trichoderma flavipes (Peck) Seifert, Jaklitsch & Voglmayr, p. 148

Trichoderma foliicola (Jaklitsch & Voglmayr) Jaklitsch & Voglmayyr, p. 149

Trichoderma hispanicum (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, p. 149

Trichoderma hunua (Dingley) Jaklitsch & Voglmayr, p. 149

Trichoderma lacuwombatense (B.S. Lu, Druzhin. & Samuels) Jaklitsch &

Voglmayr, p. 149

Trichoderma megalocitrinum (Yoshim. Doi) Jaklitsch & Voglmayr, p. 149

Trichoderma microcitrinum (Yoshim. Doi) Jaklitsch & Voglmayyr, p. 149

Trichoderma neorufum (Samuels, Dodd & Lieckf.) Jaklitsch & Voglmayr, p. 150

Trichoderma novae-zelandiae (Samuels & Petrini) Jaklitsch & Voglmayyr, p. 150

Trichoderma nybergianum (T. Ulvinen & H.L. Chamb.) Jaklitsch & Voglmayr, p. 150

Trichoderma ochroleucum (Berk. & Ravenel) Jaklitsch & Voglmayr, p. 150

Trichoderma orientale (Samuels & Petrini) Jaklitsch & Samuels, p. 151

Trichoderma parapiluliferum (B.S. Lu, Druzhin. & Samuels) Jaklitsch &

Voglmayr, p. 151

Trichoderma parmastoi (Overton) Jaklitsch & Voglmayyr, p. 151

Trichoderma patella (Cooke & Peck) Jaklitsch & Voglmayr, p. 151

Trichoderma peltatum (Berk.) Samuels, Jaklitsch & Voglmayyr, p. 151

Trichoderma pezizoides (Berk. & Broome) Samuels, Jaklitsch & Voglmayyr, p. 152

Trichoderma protopulvinatum (Yoshim. Doi) Jaklitsch & Voglmayr, p. 152

Trichoderma pulvinatum (Fuckel) Jaklitsch & Voglmayr, p. 152

Trichoderma rhododendri (Jaklitsch & Voglmayr) Jaklitsch & Voglmayr, p. 152

Trichoderma rodmanii (Samuels & P. Chaverri) Jaklitsch & Voglmayr, p. 152