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Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales)

January 2013

January 2013
46(7):358–380

Authors:

Ivan V. Zmitrovich

Russian Academy of Sciences

Vera Malysheva

Komarov Botanical Institute of the Russ. Acad. Sc. (BIN RAS), St Petersburg

Рассмотрены вопросы молекулярной таксономии и традиционной (основанной на анализе комплекса морфологических признаков) систематики трутовых грибов, характеризующихся траметоидным морфотипом. Представлен обзор истории таксономического «дробления» рода Trametes. Реконструировано молекулярное филогенетическое древо порядка Polyporales c использованием данных по первичной структуре регионов большой субъединицы и внутренних транскрибируемых спейсеров (ITS) рибосомальной ДНК. Подробно рассмотрены морфологические особенности грибов траметоидного морфотипа, и полученные молекулярные данные сопоставлены с имеющимися морфологическими сведениями; очерчены границы родов. Показано, что в порядке Polyporales выделяются две основные филогенетические радиации уровня семейства – Meruliaceae и Polyporaceae, обе из которых содержат роды траметоидных грибов. Показано также, что род Panus, характеризующийся настоящим пластинчатым гименофором, оказался дистанцированным от родов Lentinus and Polyporus, но близким к Cerrena и Steccherinum – родами «мерулиоидной радиации». Роды траметоидных грибов семейства Polyporaceae оказались в составе «траметоидного» (Pycnoporus, Cubamyces, Cellularia, Coriolopsis s. str., Sclerodepsis, Artolenzites, Lenzites, Trametes s. str.) и «полипороидного» (Datronia, Earliella, Microporus, Daedaleopsis, Hexagonia s. str., Pogonomyces, Funalia) кластеров. Сделано 7 новых комбинаций: Daedaleopsis nitida (Dur. et Mont.), Cellulariella acuta (Berk.), Cellulariella warnieri (Dur. et Mont.), Funalia aspera (Jungh.), F. caperata (Berk.), F. floccosa (Jungh.), and F. sanguinaria (Klotzsch). Описана новая разновидность Trametes pubescens var. anthopora c Дальнего Востока России. Описан род Cellulariella. Составлена диагностическая таблица родов и представлен конспект видов траметоидных грибов.

The best tree of large-scale Polyporales obtained from the ML analysis of the nLSU dataset. The red colour indicate the subclades with BS values from MP analysis 870 and PP 80.95 from the BY analysis. Others values from MP and BS analyses are shown under and above the branches. The trametoid genera are marked with asterisk.

…

The best tree of the core-Trametes obtained from the ML analysis of the ITS dataset. Bootstrap values (%) from MP and Posterior probability from BY analyses are shown under and above the branches respectively.

…

Trametes pubescens var. anthopora (LE 286999). Above-external habitus of basidiomata: a-upperside, bhymenophore side; scale bar-10 mm. Below-sclerohyphae and basidiospores; scale bar-10 mm.

…

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ÓÄÊ 582.284.99:57.065

TOWARDS A PHYLOGENY OF TRAMETES ALLIANCE

(BASIDIOMYCOTA, POLYPORALES)

ÇÌÈÒÐÎÂÈ× È. Â., ÌÀËÛØÅÂÀ Â. Ô. Î ÔÈËÎÃÅÍÈÈ TRAMETES

È ÐÎÄÑÒÂÅÍÍÛÕ ÒÀÊÑÎÍÀÕ (BASIDIOMYCOTA, POLYPORALES)

Komarov Botanical Institute RAS, St. Petersburg

iv_zmitrovich@mail.ru

The paper deals with morphology-based and molecular taxonomy of the polypores having the trameto-

id morphotype. The history of Trametes restriction was presented. The molecular phylogeny of Polypora-

les highlighting all the trametoid nodes was reconstructed using nLSU and ITS rDNA datasets. Fine mor-

phological markers of trametoid morphotype were revised. All the obtained molecular entities were charac-

terized morphologically and the generic limits were discussed. It was shown that two large radiations of

family level, the Meruliaceae and Polyporaceae contain the trametoid nodes. The lamellate genus Panus

has appeared to be distant from Lentinus and Polyporus, but closely related to trametoid Cerrena and Stec-

cherinum (merulioid radiation). Trametoid genera of Polyporaceae have appeared to be disseminated wi-

thin two subradiations — polyporoid and trametoid ones. The polyporoid radiation includes such genera as

Datronia, Earliella, Microporus, Daedaleopsis, Hexagonia s. str., Pogonomyces, and Funalia. The trame-

toid radiation is presented by such genera as Pycnoporus, Cubamyces, Cellularia, Coriolopsis s. str., Scle-

rodepsis, Artolenzites, Lenzites, and Trametes s. str. 7 new combinations were made as Daedaleopsis niti-

da, Cellulariella acuta, C. warnieri, Funalia aspera, F. caperata, F. floccosa, F. sanguinaria. A new varie-

ty, Trametes pubescens var. anthopora from the Russian Far East was described, as well. The genus Cellu-

lariella was formally described for substitution of Cellularia nom. ambig. The generic diagnostical key of

trametoid fungi and their species conspects were proposed.

Key words: molecular taxonomy, morphology, phylogeny.

Ðàññìîòðåíû âîïðîñû ìîëåêóëÿðíîé òàêñîíîìèè è òðàäèöèîííîé (îñíîâàííîé íà àíàëèçå êîì-

ïëåêñà ìîðôîëîãè÷åñêèõ ïðèçíàêîâ) ñèñòåìàòèêè òðóòîâûõ ãðèáîâ, õàðàêòåðèçóþùèõñÿ òðàìåòî-

èäíûì ìîðôîòèïîì.

Ïðåäñòàâëåí îáçîð èñòîðèè òàêñîíîìè÷åñêîãî «äðîáëåíèÿ» ðîäà Trametes.

Ðåêîíñòðóèðîâàíî

ìîëåêóëÿðíîå ôèëîãåíåòè÷åñêîå äðåâî ïîðÿäêà Polyporales c èñïîëüçîâàíèåì äàííûõ ïî ïåðâè÷-

íîé ñòðóêòóðå ðåãèîíîâ áîëüøîé ñóáúåäèíèöû è âíóòðåííèõ òðàíñêðèáèðóåìûõ ñïåéñåðîâ (ITS)

ðèáîñîìàëüíîé ÄÍÊ. Ïîäðîáíî ðàññìîòðåíû ìîðôîëîãè÷åñêèå îñîáåííîñòè ãðèáîâ òðàìåòîèäíîãî

ìîðôîòèïà, è ïîëó÷åííûå ìîëåêóëÿðíûå äàííûå ñîïîñòàâëåíû ñ èìåþùèìèñÿ ìîðôîëîãè÷åñêèìè

ñâåäåíèÿìè; î÷åð÷åíû ãðàíèöû ðîäîâ. Ïîêàçàíî, ÷òî â ïîðÿäêå Polyporales âûäåëÿþòñÿ äâå îñíîâ-

íûå ôèëîãåíåòè÷åñêèå ðàäèàöèè óðîâíÿ ñåìåéñòâà — Meruliaceae è Polyporaceae, îáå èç êîòîðûõ

ñîäåðæàò ðîäû òðàìåòîèäíûõ ãðèáîâ. Ïîêàçàíî òàêæå, ÷òî ðîä Panus, õàðàêòåðèçóþùèéñÿ íàñòîÿ-

ùèì ïëàñòèí÷àòûì ãèìåíîôîðîì, îêàçàëñÿ äèñòàíöèðîâàííûì îò ðîäîâ Lentinus and Polyporus,íî

áëèçêèì ê Cerrena è Steccherinum — ðîäàìè «ìåðóëèîèäíîé ðàäèàöèè». Ðîäû òðàìåòîèäíûõ ãðè-

áîâ ñåìåéñòâà Polyporaceae îêàçàëèñü â ñîñòàâå «òðàìåòîèäíîãî» (Pycnoporus, Cubamyces, Cellula-

ria, Coriolopsis s. str., Sclerodepsis, Artolenzites, Lenzites, Trametes s. str.) è «ïîëèïîðîèäíîãî» ( Datro-

nia, Earliella, Microporus, Daedaleopsis, Hexagonia s. str., Pogonomyces, Funalia) êëàñòåðîâ. Ñäåëàíî

7 íîâûõ êîìáèíàöèé: Daedaleopsis nitida, Cellulariella acuta, C. warnieri, Funalia aspera, F. caperata,

F. floccosa, and F. sanguinaria. Îïèñàíà íîâàÿ ðàçíîâèäíîñòü Trametes pubescens var. anthopora c

Äàëüíåãî Âîñòîêà Ðîññèè. Îïèñàí ðîä Cellulariella. Ñîñòàâëåíà äèàãíîñòè÷åñêàÿ òàáëèöà ðîäîâ è

ïðåäñòàâëåí êîíñïåêò âèäîâ òðàìåòîèäíûõ ãðèáîâ.

Êëþ÷åâûå ñëîâà: ìîëåêóëÿðíàÿ òàêñîíîìèÿ, ìîðôîëîãèÿ, ôèëîãåíèÿ.

ÌÈÊÎËÎÃÈß È ÔÈÒÎÏÀÒÎËÎÃÈß

Òîì 47 2013 Âûï.6

ÁÈÎÐÀÇÍÎÎÁÐÀÇÈÅ ÑÈÑÒÅÌÀÒÈÊÀ, ÝÊÎËÎÃÈß

An interest to the genus Trametes increases during se

veral last decades. This genus contains one of the most im

portant medicinal mushrooms Trametes versicolor or Tur

key tail; the fungus has been actively studied during the

last 20 years as a main producer of «Polysaccharide-K»,

protein-bound polysaccharide with anticancer activity

(Stamets, 2005; Wasser, Didukh, 2005) and producer of

oxidoreductases for such biotechnological properties as

delignification of wood pulp, cellulose fiber bleaching, de-

gradation of organic pollutants — pentachlorophenols, po-

lychlorinated biphenyls, polycyclic aromatic hydrocarbons

and others (Heinfling et al., 1997; Nyanhongo et al., 2007;

Moturi et al., 2009).

However, the taxonomical content of Trametes rema-

ins unstable in all the times. Till now, this genus is accep-

ted in extremely wide sense by some authors (Corner,

1989; Zmitrovich et al., 2012) whereas the others prefer

rather fine generic splitting of alliance of trametoid fungi

(Ryvarden, 2000; Bernicchia, 2005).

Historically, the genus Trametes

was finally segrega-

ted from Polyporus by Fries (1838) for type P. suaveolens

and also to some similar species, characterized by pileate

sessile basidiocarps with densely fibrous context and

one-layered tubes. Later, Fries (1874) added more species

to this genus. In the period of artificial Polyporus splitting,

the colour and comparative thickness of the context were

basic diagnostical features. Therefore, a lot of generic units

arose parallelly to Trametes, first of all Pycnoporus with

thin rufous to carmine red tissues (Karsten, 1881), Coriolus

with thin white tissues (Quelet, 1886), Poronidulus — the

same as previous with peculiar nodes outside pileus (Mur-

rill, 1904), and Coriolopsis with thin brown tissues (Murrill,

1905). Lloyd (1921) and later Pil*at (1939) merged the genus

Coriolus into Trametes, since the basidiocarps thickness in

the fungi discussed varied in rather wide range.

The following four decades were characterized by cer-

tain complication of the Trametes concept. The genera

Pseudotrametes (Bondarzew, Singer, 1941; Singer, 1944)

and Funalia (Patouillard, 1900; Bondarzew, Singer, 1941)

were described for Trametes gibbosa and T. trogii, respec-

tively. Donk (1966) described the genus Datronia for Frie-

sian Trametes mollis due to its double context and rather

sinuose pores. This author united the genera Pseudotrame-

tes and Trametes, whereas the genus Coriolus he conside-

red separately (Donk, 1974).

Corner (1989), basing on net of characters revealed af-

ter tropical account, united within enlarged Trametes all

the aforementioned generic units, plus such genera as Mic-

roporus (Beauvois, 1804), Cerrena (Gray, 1821), Hexago-

nia, Lenzites (Fries, 1838), Daedaleopsis (Schroeter,

1889), Cryptoporus (Shear, 1902), Earliella (Murrill,

1905), Mollicarpus (Ginns, 1984), and even brown-rot Fo-

mitopsis (Karsten, 1881). Some additional combinations in

such enlarged Trametes were made by Zmitrovich et al.

(2012). This solution certainly removed a contradiction

between tropical and boreal generic concepts, but has not

made a mycological mainstream, which was fixed by Gil

bertson and Ryvarden (1986; Ryvarden, Gilbertson, 1994).

The last authors received the genus Trametes as

inclusive

for Coriolus and Pseudotrametes, but kept separately such

white-rot genera as Cerrena, Coriolopsis, Lenzites, Hexa

gonia, and Pycnoporus.

In the recent decade, some molecular approaches in the

Trametes-conglomerate were realized (Tom&sovsky et al.,

2006; Cui et al., 2011; Malysheva, Zmitrovich, 2011), and

the most complete dataset was presented by Justo and Hib-

bett (2011) and Welti et al. (2012).

According to Justo and Hibbett (2011), this conglome-

rate has appeared to be disseminated within «large polypo-

roid» and «large trametoid» clades. In the «large polyporo-

id clade», the Trametes-like fungi were dispersed within

the following clusters:

— the Lentinus-clade (Trametes trogii, T. gallica,

T. aspera);

— the Polyporus—Datronia-clade (Trametes mollis

and T. scutellata);

— the Daedaleopsis—Earliella-clade (T. confragosa,

T. scabrosa, T. hirta);

— the «Ganodermataceae»-clade (Trametes cf. byrsi-

na, Cryptoporus volvatus).

The large trametoid clade in Justo and Hibbett’s tree,

comprised core Trametes (T. ectypa, T. versicolor, T. och-

racea, T. conchifera, T. villosa) and neighbouring clusters

as:

— the Lenzites-clade (T. betulina, T. gibbosa, T. pavo-

nia, T.

membranacea);

— the Coriolopsis-clade (T. polyzona);

— the Artolenzites-clade (T. maxima, T. elegans);

— the Pycnoporus-clade (T. cinnabarina, T. sangui-

nea, T. cubensis).

Such genera as Cerrena and Fomitopsis in research

under consideration have left far beyond core polyporoid

radiation to Meruliaceae- and Antrodia-clades, respective-

ly, that corresponded to other molecular phylogenies. The

morphological separation of parallel morphotypes in «lar-

ge polyporoid» and «large trametoid» clades as well as the-

ir subclades is recognized by quoted authors as «still prob-

lematic». The research by Welti et al. (2012) was focused

to «large trametoid» clade, initially. Three main lineages

were revealed here by these authors: 1) a monospecific li-

neage represented by Artolenzites elegans, 2) a lineage inc-

luding the genus Pycnoporus in its traditional sense plus

Trametes cingulata, T. lactinea, T. ljubarskyi, T. menziesii

and 3) a lineage corresponding to the core genus Trametes,

including type species of Trametes, Coriolopsis and Lenzi-

tes.

The present paper is devoted to further nuclear rDNA

and morphological study of conglomerate of trametoid

fungi, with involving to our research of additional (predo-

minantly southern) hexagonoid, favoloid and lentinoid re-

presentatives of both polyporoid and trametoid radiations.

Since a selection of the objects for analysis affects the re-

sults very strongly, our purpose was to involve in the rese-

arch such a non-trametoid taxa, which are identical to Tra-

metes micromorphologically, e. g. Polyporus, Lentinus

s. l., and Panus (Singer, 1962; Corner, 1981, 1989; Ryvar

den, 1991) what, having affected on additional node pro

duction, leads to highlighting of some generic borders. On

the other hand, we highlighted only few regions of co

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 359

* The authors of fungal names are presented in the taxonomical

part of the paper.

re-Trametes clade, in contrast to Justo and Hibbett (2011),

who overviewed all the core-Trametes content. Also, in

our recent work (Malysheva, Zmitrovich, 2011) we alrea

dy clearly separated T. hirsuta from T. pubescens-conglo

merate. However, the last one remains poorly resolved.

Therefore, we have involved some material from the Rus-

sian Far East in the present analysis, particularly, a certain-

ly deviant forms of T. conchifera and atypical Far Eastern

T. pubescens. Third our purpose was the morphological

testing of units appeared in frameworks of rDNA-phylo-

geny.

Materials and methods

Taxa selection. A selection of taxa for the present ana-

lyses was based on our knowledge on overall molecular

phylogeny of the Polyporales and elaboration of individual

areas of global tree (Kim, 1998; Kim, Jung, 2000; Kruger,

Gargas, 2004; Tom&sovsky et al., 2006; Dai et al., 2007;

Sotome et al., 2008; Lee, Lim, 2010; Cui et al., 2011; Jus-

to, Hibbett, 2011; Miettinen et al., 2011; Welti et al., 2012)

with account of capture of some new nodes. In our opinion,

some additional key nodes can be captured by involving of

several taxa of such artificial genera as Hexagonia, Corio-

lopsis, Lentinus, and Polyporus.

Taxon sampling. Fourteen specimens of Trametes

examined in the morphological study were selected for

molecular analysis. We generated a total of 15 ITS and

4 nLSU sequences for this study.

75 additional ITS sequences and 63 additional LSU

sequences of other polyporoid/merulioid genera were re-

trieved from GenBank based on BLAST results

(http:/www.ncbi.nlm.nih.gov/BLAST/) with polyporoid/

merulioid species as well as the data of the Justo and Hib-

bett (2011) paper.

The final dataset consisted of 92 ITS sequences and 68

nLSU sequences.

For all analyses the sequences of Thelephora terrestris

and Exidia glandulosa retrieved from GenBank were cho-

sen as outgroup based on BLAST homology. In analyses

for LSU-dataset E. glandulosa was selected as an outgroup

and E. glandulosa plus Thelephora terrestris — for

ITS-dataset.

An overview of all taxa studied is given in Table,

which shows the names of species, GenBank accession

numbers, Herbarium numbers and collection particulars.

DNA extraction, PCR and sequencing. DNA was

extracted from herbarium material using 2 % CTAB ex-

traction buffer with following steps of consecutive additi-

on of chloroform-isoamyl alcohol mixture (24:1), then

isopropyl alcohol and 3M sodium acetate solution for pre-

cipitation, 70 % ethanol for washing and finally water for

dissolution. The alternative method of extraction DNA

was using NucleoSpin

Plant II Kit (Macherey-Nagel).

The ribosomal ITS1—5.8S—ITS2 region was amplified

by PCR with the fungal specific primers ITS1F and ITS4B

(Gardes, Bruns, 1993) and 28S region was amplified by

PCR with the fungal specific primers LROR and LR7 (Vil

galys Laboratory, http:/www.biology.duke.edu/fungi/my

colab/primers). The PCR products were purified using Fer

mentas Genomic DNA Purification Kit (Thermo Scienti

fic). Sequencing of this strand was performed with ABI

model 3130 Genetic Analyzer (Applied Biosystems) using

BigDye™ Terminator Cycle Sequencing Ready Reaction

Kit (Applied Biosystems) with the same primers. The raw

data were processed using Sequencing Analysis 5.3.1 (Ap-

plied Biosystems).

Alignments and phylogenetic analysis. The ITS and

LSU sequences were aligned with the web tools MAFFT

(http:/align.bmr.kyushu-u.ac.jp/mafft/online/server/) with

Q-INS-i strategy and default settings for other options. The

final alignment was corrected manually using MEGA 5

(Tamura et al., 2011).

Phylogenetic reconstructions were performed with

maximum parsimony (MP), maximum likelihood (ML)

and Bayesian (BA) analyses.

A MP analysis was performed using PAUP*4.0.b10

(Swofford, 2002). One hundred heuristic searches were

conducted by stepwise addition with random sequence

addition and tree bisection-reconnection (TBR) branch-

swapping algorithm. One tree was held at each step during

stepwise addition and the number of trees retained was

limited to 100. Parsimony bootstrap analysis was perfor-

med with 1000 replicates. Gaps were treated as missing

characters. Clades with only a support 8 50 % were retai-

ned.

ML was run in the RAxML server, v.7.2.8 (http:/

http:/phylobench.vital-it.ch/raxml-bb/; Stamatakis et al.,

2008), under GTR model with one hundred rapid bootstrap

replicates.

Bayesian analysis was performed using MrBayes 3.1

for two independent runs, each with 10 000 000 generati-

ons with sampling every 100 generations, with GTR model

and four chains. Posterior probability (PP) value 8 0.95

are considered as significant.

Morphological elaboration. Microscopical study of

basidiomata was carried out as described by Gilbertson

and Ryvarden (1986). Freehand sections and squash mo-

unts of basidiomata were examined in the Melzer’s rea-

gent, 5 % KOH and 2 % Cotton Blue.

Results and discussion

New sequences and alignments. A total of 19 new se-

quences were reconstructed during our study: 15 ITS and

4 nLSU (see Table). Two new sequences of Trametes ‘pu-

bescens’ (LE 286996 and LE 286999) belonging to origi-

nally collected material from the Russian Far East, appea-

red to be phylogenetically independent lineage within the

T. pubescens—T. ochracea-clade. Below in the text, a uni-

que morphological pattern of representatives of this linea-

ge will be considered, and a new variety, T. pubescens var.

anthopora, will be described.

A total of 141 sequences were obtained from Gen-

Bank: 77 ITS and 64 nLSU. All these were checked by

BLAST-inquiries. The sequence AF 163576 marked as

«Hexagonia hydnoides»(Polyporaceae) in GenBank was

recognized as sister lineage to Irpex lacteus and Emmia la

temarginata (Meruliaceae). The sequences FJ 810183 and

FJ 481044 marked as «Lenzites betulina» (trametoid radia

360 I. V. Zmitrovich, V. F. Malysheva

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 361

Collections used in this study, their country, herbarium, GenBank accession number

Taxon Country Herbarium GenBank accession No

ITS

Cerrena unicolor China FJ 810175

Òî æå » FJ 478121

» » Poland DQ 056858

‘Coriolopsis byrsina’ USA FJ 810520

‘Coriolus’ caperata India JF 283779

Òî æå Cuba: Bayamo AB 158316

‘Coriolus’ gallica USA: Arizona JN 165013

‘Coriolus’ rigida USA JF 894113

Òî æå Jamaica JF 894112

‘Coriolopsis’ sanguinaria China FJ 627251

‘Coriolus’ trogii » HM 050415

Òî æå China: Mogao Grottoes HQ 380781

» » China GU 199348

Daedalea dickinsii Japan AB 733159

Daedaleopsis confragosa France JX 082375

Datronia mollis USA: New York JN 165007

Diplomitoporus flavescens Germany FN 907908

Earliella scabrosa Venezuela FJ 711056

Òî æå JN 165006

» » Puerto Rico JN 165009

Exidia glandulosa Canada AY 509555

Flavodon flavus Myanmar JQ 638521

Òî æå China FJ 478126

Ganoderma applanatum United Kingdom: England AY 884178

G. applanatum Germany FR 686556

‘Hexagonia’ nitida France: Corsica JN 645082

‘Hexagonia’ tenuis Taiwan JN 003683

‘Hexagonia hydnoides’(Irpex) Korea AF 163576

‘Lentinus’ lecomtei Japan JQ 955726

Lentinus tigrinus Mongolia AF 516518

Òî æå DQ 056860

‘Lentinus’ strigellus Brazil JQ 955722

‘Lenzites betulina’(Daedaleopsis) China FJ 810183

Òî æå » FJ 481044

Lopharia cinerascens USA: Mississippi JN 165019

Microporus affinis Vietnam LE 269654 KC 503507

M. ‘luteus’ Russia: AP LE 289450 KC 503504

Òî æå Òî æå LE 231601 KC 503505

» » » » LE 289449 KC 503503

M. subaffinis

China FJ 627249

xanthopus Cameroon LE 232511 KC 503506

Neolentinus cyathiformis Germany EF 524038

N. lepideus HM 536098

Panus conchatus Finland JN 710579

Phlebia tremellosa France: Orne, Belleme JX 082340

Phyllotopsis nidulans China GQ 142019

Polyporus arcularius Canada: Ontario AF 516522

P. badius Canada HQ 604799

‘Polyporus grammocephalus’ Paraguay: Alto Parana AF 516563

Polyporus squamosus Germany FR 686579

Òî æå Bulgaria HQ 439363

P. varius USA: North Carolina AF 516578

Òî æå Germany AF 516574

Pycnoporus cinnabarinus AF 363769

Òî æå AF 363757

»» AF 363765

Taiwanofungus camphoratus Taiwan JN 165025

362 I. V. Zmitrovich, V. F. Malysheva

Continued table

Taxon Country Herbarium GenBank accession No

Taiwanofungus camphoratus Taiwan DQ 455776

Thelephora terrestris HM 189965

Trametes conchifera USA: New York JN 164988

Òî æå USA: South Carolina JN 164925

» » USA: Mississippi JN 164924

» » USA: New York JN 164926

» » USA: Virginia JN 164939

» » USA: Wisconsin JN 164987

» » Russia: AP LE 286986 KC 503502

» » Òî æå LE 19127 KC 503501

T. cubensis Venezuela JN 164989

Òî æå » JN 164922

» » USA: Mississippi JN 164923

T. elegans JN 164985

T. maxima Venezuela JN 164957

T. ochracea Czech Republic AY 684177

Òî æå Canada: Ontario JN 164948

» » USA: Alaska JN 164976

T. polyzona Venezuela JN 164979

T. pubescens Ukraine LE 231728 HQ 435864

Òî æå Karelia LE 208081 HQ 435858

» » Russia: AP LE 231718 HQ 435847

» » Russia: EP LE 257070 HQ 435845

» » Òî æå LE 262759 HQ 435851

» » » » LE 254528 HQ 435870

» » Russia: AP LE 286996 KC 503499

» » Òî æå LE 286999 KC 503500

T. suaveolens USA: Wisconsin JN 164967

T. versicolor Argentina JN 164965

Òî æå Brazil JN 164975

» » Japan AB 592335

Trametopsis cervina Czech Republic AY 684175

Òî æå USA: Florida JN 164991

Schizophyllum commune Indonesia: Java AJ 537501

Steccherinum ochraceum Norway: Vestfold AF 126906

nLSU

Cerrena unicolor Finland FJ 821524

Òî æå Sweden FJ 821525

» » South Korea FJ 821526

‘Coriolopsis byrsina’ AY 351955

‘Coriolopsis’ caperata AY 351947

‘Coriolopsis’ gallica USA: Arizona JN164814

‘Coriolopsis’ sanguinaria Taiwan AY 351949

Òî æå » AY 351950

‘Coriolopsis’ trogii USA: New York JN 164808

Daedalea dickinsii Japan AB 733338

Daedaleopsis confragosa AF 261542

Òî æå AJ 406547

Datronia mollis USA: Montana JN 164791

Diplomitoporus flavescens China GU 566006

Earliella scabrosa » AY 351946

Òî æå Puerto Rico JN 164793

Exidia glandulosa AY 293179

Ganoderma applanatum EU 232274

‘Hexagonia’ hirta China AY 351943

Òî æå » AY 351944

‘Hexagonia’ hydnoides » AY 351942

tion) were associated to Daedaleopsis-assemblage (poly-

poroid radiation).

Nuclear LSU dataset of the Polyporales. The data

obtained (186 parsimony informative characters; 0.3617/

0.6311 consistency/retention index) show that trametoid

polypores are distributed within several clades of Polypo

rales within two basic crown radiations of this order, cor

responded to families Meruliaceae and Polyporaceae

(Fig. 1). Besides, some trametiod polypores appeared to be

distributed within the «lower» areas of Polyporales-tree.

These areas (comprised mainly various groups of «brown-

rot polypores») were not highlighted by the present rese

arch and were visualized as «long branch assemblage» in

the cladorgam, where were intermixed with some agaric li

neages (Phyllotopsis nidulans, Schizophyllum commune)

and Gloeophyllales (Neolentinus).

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 363

Continued table

Taxon Country Herbarium GenBank accession No

‘Hexagonia’ tenuis China AY 351937

Òî æå AY 351936

‘Lentinus’ lecomtei Japan JQ 955733

‘Lentinus’ strigellus Brazil JQ 955731

L. tigrinus AF 518627

Òî æå AF 135173

Lenzites betulina AF 261543

Microporus affinis Vietnam LE 269654 KC 503509

Òî æå China AY 351931

M. xanthopus » AY 333799

Òî æå Japan AB 566341

Neolentinus lepideus » AB 733312

Òî æå HM 536077

Panus conchatus Mexico AY 616004

Phlebia tremellosa AY 293200

Phyllotopsis nidulans Germany AY 207278

‘Polyporus’ arcularius South Korea HM 003896

P. badius Japan AB 368083

P. grammocephalus » AB 587619

P. squamosus USA AF 135181

Òî æå AF 393069

P. varius Japan AB 368111

P. ‘varius’ AF 261540

Pycnoporus cinnabarinus AF 393074

Òî æå AF 261536

»» HQ 891298

Taiwanofungus camphoratus Taiwan EU 232280

Òî æå » EU 232281

‘Trametes’ elegans USA AY 855912

T. conchifera USA: Mississippi JN 164797

Òî æå Japan AB 368116

T. ‘conchifera’ Russia: AP LE 286999 KC 503510

‘Trametes’ cubensis USA: Florida JN 164787

Òî æå USA: Mississippi JN 164798

‘Trametes’ gibbosa Peru FJ 890419

Òî æå United Kingdom: England JN 164800

T. maxima Venezuela JN 164804

T. ochracea USA: Minnesota JN 164812

‘Trametes’ polyzona Ghana JN 164790

T. pubescens Ukraine LE 231728 KC 503508

Òî æå Russia: EP LE 257070 KC 503511

T. suaveolens Czech Republic AY 855909

T. versicolor USA: New York JN 164809

Òî æå AJ

406538

Trametopsis cervina USA: Mississippi JN 164796

Schizophyllum commune AF 261587

Steccherinum ochraceum Sweden EU 118670

Note. EP — European part of Russia, AP — Asian part of Russia.

364 I. V. Zmitrovich, V. F. Malysheva

Fig. 1. The best tree of large-scale Polyporales obtained from the ML analysis of the nLSU dataset. The red colour indicate the

subclades with BS values from MP analysis 870 and PP 80.95 from the BY analysis. Others values from MP and BS analyses are

shown under and above the branches. The trametoid genera are marked with asterisk.

The structure of merulioid radiation as a whole is cor

responded well to the current trees (Larsson, 2007; Lee,

Lim, 2010; Miettinen, Larsson, 2011). Particularly, this ra

diation includes such trametoid genera as Diplomitoporus

related to Steccherinum, Junghuhnia and Antrodiella (Mi-

ettinen, 2011), Trametopsis related to Phlebia-clade

(Tom&sovsky, 2008), and Cerrena, occupying an intermedi-

ate position between Phlebia- and Steccherinum-clades

(Ko, Jung, 1999; Lee, Lim, 2010). The sufficiently new

data were obtained by us for structure of Meruliaceae tree:

the genus Panus (type Panus conchatus plus several stri-

gose southern representatives currently known as Lenti-

nus-members) has appeared to be a sister taxon of Cerre-

na. Both taxa form a single robust clade.

Crown Polyporaceae (bordered by Lopharia basal

branch) comprises two large radiations, polyporoid and

trametoid ones.

The trametoid members of polyporoid radiation were

successfully distributed within such entities as Earliella

(contains Earliella scabrosa and an ambiguous sequence

marked as «Coriolopsis byrsina» FJ 810520), core Hexa-

gonia (H. hirta), Daedaleopsis (contains properly Daeda-

leopsis plus ‘Hexagonia’ tenuis), Microporus (well corres-

ponded to the current genus concept), and Funalia (suffici-

ent part of Coriolopsis-tail excluding type C. polyzona).

Also, the trametoid Datronia mollis occupies the position

within internal radiation of Polyporus, whereas ‘Coriolop-

sis’ caperata is clustering with ganodermoid lineage. As a

whole,

all these data are corresponded to Justo and Hib-

bett’s (2011) phylogeny, with small addition: ‘Hexagonia’

hydnoides (type of old Pogonomyces) appeared to be sister

taxon of Lentinus s. str., the union, comprising Lentinus

sect. Tigrini sensu Pegler (1983) and Polyporellus in sense

of old authors (including such taxa as Polyporus tricholo-

ma, P. arcularius, P. brumalis).

The trametoid radiation is densely branched. This inc-

ludes four well-supported clades as core-Trametes (conta-

ins Trametes pubescens—T. ochracea complex, T. con-

chifera, T. suaveolens, etc.), Lenzites (contains type of

Lenzites and T. gibbosa — such an arrangement was de-

monstrated by Tom&sovsky et al., 2006), Pycnoporus (well

corresponded to current genus concept), and Cubamyces

(contains ‘Trametes’ cubensis and ‘Trametes’ elegans; the

position of second one is unstable in comparison with ITS

dataset). Also, two lineages in basal multifurcation of co-

re-Trametes are presented by ‘Trametes’ polyzona and

‘Trametes’ maxima. Morphologically, these lineages cor-

responded to genera Coriolopsis s. str. and Sclerodepsis

Cooke sensu Ryvarden (1972).

ITS dataset of the Polyporales. The data obtained

(340 parsimony informative characters; 0.3470/0.6923

consistency/retention index) corresponds well to nLSU da-

taset (Fig. 2).

Both Meruliaceae and Polyporaceae-superclades are

steadily allocated on ITS tree as well as the polyporoid and

trametoid radiations in crown region of Polyporaceae-su-

perclade.

Similarly to nLSU-tree, the Meruliaceae superclade

contains Trametopsis in

the phlebioid clade, Cerrena and

Panus within the steccherinoid clade and Diplomitoporus

as basal to the latter.

The polyporoid radiation of Polyporaceae-superclade

gives the same lineages as in nLSU tree, namely Micropo

rus (contains species of the genus Microporus), Earliella

(corresponded to the genus Earliella with addition of am

biguous sequence of «Coriolopsis byrsina» FJ 810520).

The Lentinus-clade has the same position as in nLSU-phy-

logeny, having the ‘Polyporus’ arcularius taken away. The

clade Daedaleopsis includes, besides the ‘Hexagonia’ te-

nuis, also ‘Hexagonia’ nitida. The Funalia-clade compri-

ses such taxa as ‘Coriolopsis’ caperata,‘Coriolopsis’ rigi-

da, and also ‘Coriolopsis’ gallica together with ‘Coriolop-

sis’ trogii inside a proof subcluster. The position of

‘Coriolopsis’ sanguinaria can be considered as incongru-

ent to nLSU-topology element, which is uncertain and ra-

ther distant from Funalia-body. The Polyporus-clade cor-

responds to traditional restricted concept of the same genus

but, like nLSU phylogeny, comprises the Datronia mollis

as a terminal lineage of Polyporus squamosus-subclade.

Within the general Polyporales ITS-tree, the trametoid

radiation was collapsed, as it seen on Fig. 2. There was a

separate reconstruction of this radiation during the present

research.

ITS dataset of the trametoid radiation. The general

topology of trametoid radiation (Fig. 3) demonstrates high

degree of congruency with those on nLSU tree: three ro-

bust

clades are distinguishable. These are core Trametes,

Pycnoporus and Cubamyces. However, unlike to

nLSU-phylogeny, the position of the ‘Trametes’ elegans

here is distant from Cubamyces and is rather uncertain.

Additionally, some lineage approximately of generic rank

as Coriolopsis s. str. (containing ‘Trametes’ polyzona) and

Sclerodepsis (containing ‘Trametes’ maxima) are identifi-

ed, like nLSU-phylogeny. An ambiguous lineage ‘Polypo-

rus grammocephalus’ (AF 516563) supplements a basal

multifurcation of the core-Trametes, but this sequence ne-

eds to be checked. Within the core-Trametes clade two

species complexes are highlighted: the Trametes conchife-

ra, which gave two sublades corresponded to North Ame-

rican and Far East Russian material, and the T. pubescens,

which contains easily allocated lineage (Trametes pubes-

cens LE 286996, LE 286999), which can be interpreted as

an independent variety T. pubescens var. anthopora (see

taxonomical part).

Morphological dataset of the trametoid fungi

(Fig. 4). The following range of morphological variability

of taxa of the trametoid morphotype included into analysis

has been presented (terminology according to Zmitrovich

et al., 2011).

General characteristics. The trametoid mor-

photype unites a wood-inhabiting fungi, which forming the

annual or perennial sessile or shortly-stipitate tough (con-

sisted mainly by sclerohyphae), not xanthochroic basidio-

mata with one-layered hymenophores with pores, rarely

cellars or lamellae; generative hyphae with clamps; spores

smooth, thin-walled, inamyloid: cylindrical to ellipsoid;

the fungi cause an active white rot.

Habitus:a)trametoid — sessile, with sterile context

thicker than hymenophoral layer at the base, but dimini

shes towards the margin; b) corioloid — sessile, with steri

le context equal to hymenophoral layer at the base and di

minishes towards the margin; c) scenidioid — sessile, with

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 365

sterile context equal or thinner than hymenophoral layer,

of the same thickness everywhere; d) polyporoid — stipi

tate, with sterile context thicker than hymenophoral layer

at the base, diminishes toward the margins (Zmitrovich et

al., 2012).

Sizes: a) minute (1—3 cm in greatest dimension);

b) medium-sized (3—10 cm in greatest dimension); c) lar

ge-sized (over 12 cm in greatest dimension).

Upperside: a) tomentose — with outer loose layer

composed by numerous small erect hairs; as a rule, tomen

tose surfaces are furnished by dense subpellis, but in some

cases can be gradually related to context; b) villose — co

vered by sparse thin hairs, without formation of loose lay

er; subpellis present or not; c) strigose — covered by rough

erect pillars (agglutinated fascicles of sclerohyphae);

d) hispid — covered by rough hairs, pillars and crests; sub

pellis present or not; e) squalid — covered by rough crests

only; without subpellis; f) glabrous — i. e. pellis or subpel

lis without hairs and crests: (i ) crustose — as cornescent

cuticle, (ii) matt — as wooly field.

366 I. V. Zmitrovich, V. F. Malysheva

Fig. 2. The best tree of large-scale Polyporales obtained from the ML analysis of the ITS dataset. Bootstrap values (%) from MP and

Posterior probability from BY analyses are shown under and above the branches respectively. The trametoid clade is collapsed (see

Fig. 3). The trametoid genera are marked with asterisk.

Upperside general sculpture: a) scrupose —

with irregular wart-like inflations; b) ridged — with radial

ly arranged ectal folds; c) sulcate — with concentrical

rows.

Context composed by densely packed hyphal mass

with prevalence of sclerohyphae. When sclerohyphae

are hyaline, the context is white to cream; when sclerohyp

hae are pigmented, the context is tan to brown. In section:

a) homogeneous; b) duplex: (i) in density, (ii) in colo

ration.

Hymenophore: a) poroid — tubular with deep

roundish or elongated pores; b) irpicoid — toothed resul

ted of pore splitting; c) hexagonoid — consisting of short

tubes (cellars) with large hexagonal pores; d) favoloid —

the same as hexagonoid, but with radially elongated cel

lars; e) labyrinthine (daedaleoid) — with radially-arranged

elongated pores, sometimes intermixed with lamellae;

f) lamellate — with radially or fountain-like arranged la

mellae.

Hyphal system. In various researches it has been

characterized as dimitic (generative and skeletal hyphae

present) or trimitic (generative, skeletal and binding hy

phae present) in various members of the genus (Corner,

1932, 1989). The recent revision (Zmitrovich, 2010a) eli

minates a principal distinction between skeletal and bin

ding hyphae (both types substitute a pseudoskeletal hyphae

in core of basidiomata or its marginal areas, respectively).

Therefore, the hyphal system is basically dimitic — with

arboriform or sympodially branched sclerohyphae. How

ever, superficially sclerified hyphae in some trametoid

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 367

Fig. 3. The best tree of the core-Trametes obtained from the ML analysis of the ITS dataset. Bootstrap values (%) from MP and

Posterior probability from BY analyses are shown under and above the branches respectively.

368 I. V. Zmitrovich, V. F. Malysheva

fungi bear clamp-connections, i. e. the hyphal system is

pseudodimitic. Some intermediate states between pseudo

dimitic and dimitic hyphal systems can be found in old

specimens and in the herbarium.

Sclerohyphae: a) sympodially branched — more

or less dichotomously with moderate branching in a dend-

rite center and increasingly branching at the dendrite peri-

phery; there are two variants of sympodial branching pa-

ttern: regularly branched (i), and occasionally branched

(ii); b) arboriform — with strong axial element and regular

appendage axes; c) seemingly unbranched (steccherino-

id) — uninflated skeletals with collapsed appendages; the

branching of steccherinoid hyphae we can see on the stage

of pseudoskeletal precursors.

Sterile hymenial elements:a)leptocystidia —

obligate element intermixed in basidial palisade: fusoid

cells of the same length as basidia; b) hyphal pegs — rather

regular element through the genus: the fascicles of agglutina-

ted generative hyphae protruded the hymenium; c) sclerids —

unbranched or branched setiform elements connected to scle-

rohyphae, protruded the hymenium, rarely pileus cover.

Basidia:uniform element, typically clavate with inf-

lated epibasidial segment, 4-spored, clamped at the base,

4.5—6.5 mm wide, 12—30 mm in length.

Basidiospores: typically cylindrical, but also el-

lipsoid to ovoid; thin-walled in the most of representatives.

The generic concept and morphological holdings.

Each taxonomical interpretation of rRNA gene trees makes

it necessary to revise a genus idea. In due time, Prof. L. Ry-

varden has told the follows: «To me, genera exist only in

our imagination, and the concept is mostly used for two

purposes; as a handy and distinct tool for sorting species

into entities, and in an attempt to illustrate phylogeny and

evolutionary principles» (Ryvarden, 1991, p. 17). It seems

to be, the first purpose is connected with adaptive aspect of

generic units. It is rather obvious that genera represent an

adaptive character’s patterns — a certain morphotypes.

The second purpose is connected with phylogenetic aspect

of these units: the genera represent a certain lineages. As a

whole, a natural genus must combine a certain morphotype

and lineage, simultaneously. rRNA-trees, as with tubu-

lin-based, RPB-based, TEF-based and other conservative

cell components-trees reflect time frames of divergence,

but are not sensitive to adaptive morphogenetic events, es-

pecially in multicellulars. Therefore, their meaning for

taxa separation is connected to revealing of phylogenetic

events (lineage reconstruction), not taxa ranking. For gene-

ric ranking, a revealing of certain adaptive characters’ pat-

tern corresponded to morphotype is needed. rRNA phylo-

geny reveals within convergent units a certain lineages,

which needs a precise morphological revision before a ran-

king. As a rule, all the lineages revealed can be characteri-

zed morphologically, because some fine features come to

light after repeated precise testing.

As sufficient for trametoid morphotype characterizati

on, the following gross and fine morphology features were

listed in previous section.

Lineage overview and its taxonomical interpreta

tion. The fact that both nLSU- and ITS-trees are highly

congruent forces us to consider the results received as so-

und and good material for further morphological testing. A

total of 16 persistent entities of trametoid fungi will be dis-

cussed here.

Merulioid radiation

Trametopsis. The genus was described for one species,

Trametopsis cervina. It was segregated from Trametes ba-

sing on nLSU and mitSSU research (Tom&sowsky, 2008),

being clustered with Ceriporiopsis-species. Later, after wi-

dening the range of merulioid taxa in analyses, the Trame-

topsis was attracted to Irpex s. str., near to the phlebioid

clade (e. g., Justo, Hibbett, 2011). Their position around

the Irpex s. str. keeps in the present research. Morphologi-

cally, Trametopsis cervina shares the genus Diplomitopo-

rus (Spirin, Zmitrovich, 2008), having sympodially (occa-

sionally) branched, strongly cyanophilic sclerhyphae and

slightly curved basidiospores. However, for Diplomitopo-

rus-species the shorter basidia (up to 20 mm) and basidio-

spores (up to 7 mm) are characteristic. As macromorpholo-

gical differences of Trametopsis from Diplomitoporus the

squalid (vs strigose-tomentose) upperside and irpicoid ten-

dency of hymenophore should be mentioned. Therefore,

we consider the Trametopsis as a good genus both in mole-

cular and morphological perspective.

Diplomitoporus. This genus restricted to Diplomitopo-

rus crustulinus and D. flavescens in boreal mycotas was

persistent between a basal steccherinoid nodes since Lars-

son’s (2007) and Miettinen’s (2011) analyses. Such a posi-

tion keeps in our analysis. Morphologically, this unit gra-

vitates to Antrodiella in having a small basidiomata with

minute pores and short basidia (up to 17 mm), but being se-

parated by bean-like spores and occasionally (vs regularly)

sympodially branched sclerohyphae, which are weakly cy-

anophilic in both genera. The Trametopsis differs by larger

basidia and basidiospores, and also squalid (vs tomento-

se-strigose in pileate Diplomitoporus flavescens) uppersi-

de. As a whole, in restricted sense (2 boreal, plus some tro-

pical species, as D. allanosporus, etc.) this seems to be a

good generic entity.

Cerrena—Panus. A stable position of this grouping

within the steccherinoid clade is supported here by both

nLSU and ITS datasets. Concerning Cerrena, this position

has been keeping stable since Ko and Jung’s (1999) mole-

cular phylogeny, whereas Panus till now wasn’t conside-

red in connection with merulioid radiation. In current cir-

cumscription, the unit Panus is

principally correspondent

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 369

Fig. 4. The representatives of genera comprising Trametes alliance: a — Diplomitoporus flavescens, b — Cerrena unicolor, c —

Panus similis (photo A. V. Alexandrova), d — Microporus xanthopus (photo A. V. Alexandrova), e — Daedaleopsis confragosa,

f — Hexagonia apiaria (photo A. V. Alexandrova), g — Funalia trogii, h — Pogonomyces hydnoides, i — Pycnoporus

cinnabarinus, j — Cellulariella acuta (photo A. V. Alexandrova), k — Earliella scabrosa (photo A. V. Alexandrova), l — Datronia

mollis, m — Coriolopsis polyzona, n — Sclerodepsis maxima (photo A. V. Alexandrova), o — Trametes pubescens. The order of

taxa as it presented in the «Diagnostical key».

to Lentinus subgen. Panus sensu Pegler (1983). Morpholo

gically, both taxa are well united by steccherinoid (seemin

gly unbranched) pattern of the sclerohyphae and the pre

sence of skeletals terminations in hymenium as «sclerocys

tidia» and «metuloids» in Panus and the same structures

named as «pseudocystidia» in Cerrena. The differences of

two taxa are the sessile basidiomata with poroid to labyrin

thine hymenophore and ovoid basidiospores in Cerrena

versus the sitpitate basidiomata with strictly lamellate hy

menophore and cylindrical basidiospores in Panus. The

absence of any inflated hyphal elements, like Corner’s

(1984) «skeleto-binding cells» in Panus, together with the

ir slow, Trametes-like basidiome lengthening allow to con

sider this morphotype as variety of trametoid one rather

than clitocyboid and even polyporoid-lentionoid (Fig. 5).

SEM researches of basidiome ontogeny and hymenophore

development of Panus conchatus, P. rudis, and P. fulvus

show its clear difference from those of Lentinus crinitus

(genus type) and L. tigrinus: the lamellae and initial veils

in Panus and Lentinus are not homologous (Hibbett et al.,

2008). Indeed, the large genus Lentinus should be splitted

with shifting of its Panus-associated part from Polypora

ceae to Meruliaceae family. Morphogenetically, we consi

der as correct the parallel between Cerrena—Panus and

Steccherinum—Mycorrhaphium pairs, where stipe-like

trace seems to be a secondary phenomenon. If so, the la

mellae/lamellulae system of Panus, originates parallelly to

other Agaricales, Boletales and Polyporaceae, represents a

peculiar example of a deep convergence (Fig. 5).

Polyporoid radiation

Microporus. This lineage, stable in various phylogeni

es (Sotome et al., 2008; Justo, Hibbett, 2011; the present

paper), is well corresponded to morphological generic con

cept. Superficially, it is difficult to show a principal diffe

rence of this unit from Trametes s. str., because the bran

ching pattern of sclerohyphae is similar (regular sympodi

al, as in «trimitic» polypores) and the basidiospores are

analogous (short-cylindrical to exceptionally ovoid). How

ever, a re-examination of Microporus hyphal system by

Dutta and Roy (1985) showed that «trimiticism» of this po

lypore is rather conventional, and the most of sclerified

hyphae bear clamp-connections, i. e. the system is basical

ly pseudodimitic. Of course, some terminal segments of

370 I. V. Zmitrovich, V. F. Malysheva

Fig. 5. A hypothetical scheme which reflects the main directions of morphological evolution in the Polyporales. The scheme shows

that process of «trametization» has occurred independently two-three times in the Meruliaceae (Diplomitoporus, Trametopsis,

Cerrena) and two-three times in the Polyporaceae (Microporus, Datronia, the «large Trametes»). Initially, the sclerohyphae (S) in

«large Trametes» ancestor were dichophytic, on the manner of Datronia.

sclerohyphae are devoid of protoplast (the picture, similar

to Antrodiella), but on live material the most of hyphae are

clamped and septate. Also, the genus slightly deviates

from Trametes s. str. by their pigmented skeletals, by the

presence of «coralloid dichophytic elements» in hymeni-

um (Zhishu et al., 1993), and by smaller pores on generally

stipitate basidiomata (stipe often with discoid foot). As a

whole, this is a good genus, demonstrating strict paralle-

lism with Trametes s. str. (trametoid radiation).

Earliella. This lineage is well supported in Justo and

Hibbet’s (2011) as well as the present analyses. We exclu-

de from interpretation the attraction of an ambiguous sequ-

ence «Coriolopsis byrsina» FJ 810520 in our nLSU- and

ITS-trees. In monotypic circumscription, this genus is har-

dly separated from Datronia, at least superficially, because

having broadly decurrent pilei with sinuose pores together

with black-line on the context. However, the sclerohyphae

are quite different: they are subsolid and actively sympodi-

ally branched in Earliella («trimitic hyphal system» in tra-

ditional sense) and having wide lumina and rarely sympo-

dially branched in Datronia (see Roy, Mitra, 1986). The

sclerohyphae in Earliella are hyaline in contrast to the

most of tropical Coriolopsis s. l. and Hexagonia s. l. The

genus Trametes s. str., having a hyaline skeletals, differs

by their moderate branching pattern, whereas skeletals of

Earliella shares those of Dichomitus-species. The other

difference of Earliella from Trametes s. str. is the presence

of superficial crust with sclerids. Therefore, the Earliella

seems to be useful generic entity.

Hexagonia s. str. On nLSU-tree this clade represented

by single species, Hexagonia hirta, was as sister to Earliel-

la. Morphologically, this polypore is quite different: the

pores are hexagonal, the surface is hispid and the contextu-

al hyphae are pigmented, sympodially branched. The ge-

nus Hexagonia in their classical interpretation (Fidalgo,

1968) is obviously heterogeneous. The results of our inves-

tigation show that at least three entities — Daedaleopsis,

Hexagonia s. str. and Pogonomyces — are appropriate for

natural re-arrangement of Hexagonia-conglomerate.

Daedaleopsis. The clade Daedaleopsis in our

nLSU-tree comprises the Daedaleopsis in traditional volu-

me and ‘Hexagonia’ tenuis. On the ITS tree this union

comprises also ‘Hexagonia’ nitida.

These polypores have

trametoid habitus and brown-colored sclerohyphae with

regular sympodial branching. The spores are cylindrical

and slightly curved. The hymenophore varies from angu-

lar-poroid to lamellate in Daedaleopsis-part to hexago-

nal-poroid to cellar-like in Hexagonia-part. It is obviously

that hymenophore configuration cannot be a key morpho-

logical marker for trametoid fungi.

Lentinus s. str. The lineage, comprising Lentinus cri-

nitus—L. tigrinus-group with Polyporus arcularius-group

(Polyporellus in sense of old authors) was revealed in

works by Krüger with co-authors (Kruger, Gargas, 2004;

Krüger et al., 2006). Later, the clusterizing of Lentinus

squarrosulus, L. tigrinus and L. polychrous with ‘Polypo-

rus’ arcularius,‘Polyporus’ brumalis and ‘Polyporus’

tricholoma was showed by Sotome et al. (2008). In Justo

and Hibbett’s (2011) phylogeny the corresponding clade

contained Lentinus squarrosulus, L. tigrinus and ‘Polypo

rus’ arcularius,‘Polyporus’ brumalis,‘Polyporus’ tricho

loma, respectively. Both ITS- and nLSU-trees in the

present paper unite Lentinus tigrinus and ‘Polyporus’ ar

cularius, too. It is obviously that ‘Polyporus’ ciliatus and

‘Polyporus’ tenuiculus are additional members of this unit.

The lamellae-bearing part of this union is rather exactly

corresponded to Lentinus subgen. Lentinus sensu Pegler

(1983). All these representatives are characterized by the

presence of «skeleto-binding cells» — dichotomously ra-

mified hyphal segments with an inflated axial areas. Also,

the

large, elongated-cylindrical basidiospores are characte-

ristic for the species. An upperside varies from tomento-

se-strigose to squalid or squamulose. The poroid part of

this union bears an identical microsctructures. For enlar-

ged description of «skeleto-binding cells» of these polypo-

res see Corner (1984). The proper combinations of the po-

lypores under Lentinus name were made by Krüger and

Gargas (2004) and Zmitrovich (2010b). The presence of

inflated hyphal elements is an evidence for non-trametoid

nature of the Lentinus-morphotype. Bjerkanderoid-grifolo-

id morphotypes with abundance of physalohyphae within

stump-like base of multipileate basidiome can be possible

precursors for lentinoid and polyporoid ones (Fig. 5).

Pogonomyces. The lineage ‘Hexagonia’ hydnoides

has appeared on our nLSU-tree as basal for Lentinus s. str.

Superficially, this species sharply different from lentini,

but resembles Hexagonia s. str., where it was classified till

now. However, this possesses arboriform sclerohyphae

with more or less inflated axial element (the hyphae of He-

xagonia s. str. are uninflated). The fungus has a peculiar

strigose upperside with prominent pillars resembling the

thorns. As a whole, this entity is quite distinguishable both

on molecules and morphology.

Funalia. This clade contains species, traditionally

considered under the name Coriolopsis, excluding the type

species, Coriolopsis polyzona (trametoid radiation).

nLSU-tree comprises ‘Coriolopsis’ trogii,‘Coriolopsis’

gallica, and ‘Coriolopsis’ sanguinaria; on ITS-tree ‘Co-

riolopsis’ rigida and ‘Coriolopsis’ caperata are also pre-

sented, whereas ‘Coriolopsis’ sanguinaria have an uncer-

tain position. All these species are characterized by corio-

loid morpohotype with colored context. Context coloration

varies from wood-coloured in ‘Coriolopsis’ trogii to deep

brown. The sclerohyphae are regularly sympodially bran-

ched, cyanophilous. Their colour varies from golden-yel-

low to deep-brown. Both the nLSU- and ITS-analyses and

recently published data (Dai et al., 2007) show that ‘ Corio-

lopsis’ sanguinaria is rather distant from Polyporus s. str.,

what is obviously on the basis of morphology. nLSU-tree

demonstrates an attraction of ‘Coriolopsis’ caperata (spe-

cies

which keeps the Funalia-morphology) to ganodermo-

id lineage similar to Justo and Hibbett’s (2011) tree. Beca-

use ITS-tree is well includes this taxon, the position of se-

quence AY 351947 is seemingly ambiguous.

As a whole, the ganodermoid lineage is a stable branch

of polyporoid radiation (Krüger, Gargas, 2004; Dai et al.,

2007; Moncalvo, Buchanan, 2008; Hibbett, Justo, 2011).

Polyporus. This clade well corresponds to Polyporus

s. str., but with merging of Datronia (and Dichomitus)asit

show both our analyses and also several recent phylogeni

es (Dai et al., 2007; Sotome et al., 2008; Justo, Hibbett,

2011). The Polyporus-part of this clade is characterized by

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 371

polyporoid morphotype and inflated hyphae, characteristic

for Lentinus s. str. The Datronia-part is corioloid to sceni

dioid with sclerohyphae reminiscent those of Polyporus,

but with uninflated axial elements. They are precisely des

cribed by Roy and Mitra (1986). The fact that Datronia

and Polyporus squamosus form robust clade distant from

Polyporus varius-group indicates on heterogeneity of tra-

ditional Polyporus rather than incorporation of Datronia

into the genus. In the future, a precise molecular analysis

of relationships between Datronia, Dichomitus and all the

sections of Polyporus is necessary. Before this, we prefer

to consider the Datronia-unit as a trametoid genus, the sis-

ter to Polyporus squamosus. It is obviously that a separate

genus should be described for P. varius-group, but it falls

outside the present frameworks.

Trametoid radiation

Pycnoporus—Cubamyces—Artolenzites. The lineage

Pycnoporus in our nLSU- and ITS-trees is strictly corres-

ponded to morphological limits of the genus. Morphologi-

cally, the genus is characterized by corioloid habitus, a su-

perficial crust in maturity, red-colored context and mode-

rately sympodially branched sclerohyphae with weakly

amyloid walls and red-colored external deposites. In our

nLSU-tree this lineage forms a single clade with ‘Trame-

tes’ cubensis with rather insufficient support. Rather si-

milar results are presented by Welti et al. (2012) and

Justo and Hibbett (2011), where this low-supported clade

includes also ‘Trametes’ ljubarskyi,‘Trametes’ elegans,

‘Trametes’ lactinea and ‘Trametes’ cubensis. Our ITS-tree

show that at least ‘Trametes’ elegans and ‘Trametes’

cubensis are rather isolated lineages of the trametoid ra-

diation and such an entities as Artolenzites and Cubamyces

respectively are appropriate for these taxa. Artolenzites

elegans is corioloid with matt upperside and lamellate

hymenophore; the sclerohyphae are actively sympodially

branched with formation of dentate sclerids in the hyme-

nium. Cubamyces cubensis is corioloid with superfi-

cial crust; the sclerohyphae of arboriform branching

pattern; the spores are cylindrical. ‘Trametes’ ljubarskyi

is similar excepting the lacrymoid tendency in spore

outline.

Lenzites. This lineage have a strong support in our

nLSU-tree that is well corresponded to the recent Trame-

tes-phylogenies (Tom&sovsky et al., 2006; Justo, Hibbett,

2011, etc.). This includes Lenzites betulina and ‘Trametes’

gibbosa. Morphologically, this unit is characterized by tra-

metoid habitus with matt upperside, white context and ob-

long-poroid to lamellate hymenophore; the sclerohyphae

are colorless and having a sympodial branching pattern.

Two other lenzitoid species, ‘Lenzites’ warnieri and ‘Len-

zites’ acuta (widely known as Daedalea flavida) in phylo-

geny of Welti et al. (2012) and our unpublished data are

distant from Lenzites-group. It is an independent lineage of

trametoid

radiation, and formally re-description of Cellu

laria for the accommodation of the two species seems to be

better solution.

Coriolopsis s. str. This lineage is presented by single

species ‘Trametes’ polyzona, known also as Coriolopsis

occidentalis. The habit is corioloid to scenidioid with to

mentose to hispid upperside, tan context and large pores.

The sclerohyphae are moderately sympodially branched,

subhyaline. The spores are cylindrical and rather elonga

ted. Morphologically similar species is ‘Hexagonia’ varie-

gata. On other molecular trees (Justo, Hibbett, 2011; Welti

et al., 2012) this lineage is also well supported. In his book

on Tanzanian fungi, Niemelä (Härkonen et al., 2003) com-

bines ‘Trametes’ polyzona into Funalia due to its «cyano-

philous» sclerohyphae. However, on p. 144 the skeletal

hyphae of ‘Funalia’ polyzona are characterized as «green

in Cotton Blue», whereas the binding hyphae are marked

as acyanophilous.

Sclerodepsis. Such species as ‘Trametes’ maxima lo-

oks like an isolated member of trametoid multifurcation in

our tree, slightly gravitating to Artolenzites in Justo and

Hibbett’s (2011) tree and slightly gravitating to core-Tra-

metes in Welti’s collective (2012) tree. Morphologically

this entity can be described as having scenidioid to coriolo-

id habitus with sessile or decurrent, often marginally proli-

ferating basidiomata with hispid to matt upperside and of-

ten splitting to dentate hymenophore; basidiospores ellip-

soid and sclerohyphae 3—7 mm are moderately

sympodially branched, hyaline. Closely related, if not con-

specific, is ‘Trametes’ meyenii; both species are well stac-

ked into Sclerodepsis as it has outlined by Ryvarden

(1972). The generic type, ‘Trametes’ sclerodepsis, was

re-examined by Ryvarden (1984), and despite of the ab-

sence of basidiospore throughout specimen, a general con-

clusion was about clear relationships with ‘Trametes’ ma-

xima: «The species is related to Cerrena maxima (Mont.)

Hansen of the Caribbean area, which, however, is beco-

ming distinctly hydnoid with age and is paler and more

densely

hirsute». Since the type of 1856 cannot be a matter

for molecular study, the consensus concerning practical

use of this entity will be, obviously, incomplete.

Trametes s. str. The genus in restricted sense stays a

large unit, where our purpose was to test a congeneric na-

ture of Poronidulus. Actually, the Trametes conchifera has

generated as a separate clade within Trametes s. str. Mor-

phologically, this unit comprises fungi with trametoid to

corioloid habitus with strigose to matt upperside, white

context and poroid hymenophore; sclerohyphae are active-

ly sympodially branched, hyaline and acyanophilous; the

basidiospores are cylindrical. Some species, as T. hirsuta,

are well delimited (Malysheva, Zmitrovich, 2011), where-

as the complex T. versicolor—T. ochracea—T. pubescens

is rather problematic for species recognition (Tom&sovsky,

Homolka, 2004). Despite of a rather low species resolution

in this zone, we could catch a sufficient distance of Far

Eastern part of the T. pubescens conglomerate. This minu-

te trametes have thinner tube dissepiments and narrower

basidiospores, what is a reason of its segregation on a spe-

cies level (see description of T. anthopora in a taxonomical

part).

Taxonomy. The following part contains a diagnostical

table for trametoid genera and species conspect. The speci

es needed for further studies are included in those or other

units provisionally.

I. Sclerohyphae seemingly unbranched (steccherinoid) —

uninflated with collapsed appendages; their abundant

372 I. V. Zmitrovich, V. F. Malysheva

terminations in the hymenium are recognized as pse

udocystidia.

A. Habitus corioloid; hymenophore poroid to daedaleoid

or irpicoid; basidiospores ovoid .....Cerrena.

B. Habitus lentinoid; hymenophore strictly lamellate, with

lamellae and lamellulae; basidiospores cylindrical . .

....................Panus.

II. Sclerohyphae occasionally sympodially branched,

strongly cyanophilous and slightly amyloid.

A. Habitus trametoid; upperside squalid; basidia over 20

mm long and basidiospores over 7 mmlong.........

Trametopsis.

B. Habitus corioloid to tyromycetoid; upperside villose to

tomentose; basidia up to 20 mm and basidiospores up

to 7 mm............Diplomitoporus.

III. Sclerohyphae sympodially branched or arboriform,

mostly inamyloid,

cyanophilous or not.

A. Sclerohyphae clamped and in a central parts of tissues

with a living protoplast .......Microporus.

B. Sclerohyphae devoid of clamps, with empty lumina.

1. Sclerohyphae pigmented, cyanophilous; basidiospores

cylindrical.

a. Branching pattern sympodial.

a. Context tan to cinnamon, upperside glabrous, pores me-

dium-sized hexagonal to daedaleoid or hymenophore

lamellate.............Daedaleopsis.

b. Context deeply brown, upperside strigose, pores large,

hexagonal ...........Hexagonia s. str.

g. Context tan to deeply-brown, upperside hispid to matt,

pores medium- to small-sized, more or less circular

...................Funalia.

b. Branching pattern arboriform. Sclerohyphae with infla-

ted axial element. Basidiomata scenidioid to trameto-

id with small hexagonal pores; upperside strigose

.................Pogonomyces.

2. Hyphae hyaline or nearly so, acyanophilous or weakly

cyanophilous.

a. Sclerohyphae weakly branched (looks to be unbranched

in a dendrite center).

a. Corioloid, medium-sized; hymenophore poroid; hyphae

slightly amyloid, with external deposites; context

bright colored; basidiospores slightly curved....

.................Pycnoporus.

b. Trametoid, large-sized; hymenophore daedaleoid to la-

mellate; hyphae inamyloid, without deposites, in

some cases with crystalline encrustation; context cre-

am................Cellulariella.

b. Sclerohyphae regularly branched.

a. The branching dichopodial, under a sharp angle; the dia-

meter falls appreciably.

2 The lumina are distinguishable; there are coralloid ele-

ments in the hymenium; no surface sclerids .....

...................Datronia.

2 The hyphae are subsolid; there are surface sclerids; no

coralloid elements in the hymenium . . . Earliella.

b. The branching under right angle or nearly so, the diame-

ter falls insufficiently.

2 Upperside crustose; habit ostreiform; hymenophore po

roid; basidiospores cylindrical ....Cubamyces.

2 Upperside hispid to villose; habitus corioloid; hymenop

hore poroid; basidiospores cylindrical with fusoid

tendency ...........Coriolopsis s. str.

2 Upperside hispid to matt; habitus corioloid to scenidioid,

marginally proliferating; hymenophore poroid with

irpicoid tendency; basidiospores ellipsoid......

.................Sclerodepsis.

2 Upperside matt to subcrustose with intermixed hyphal

structure (type d according to Welti et al., 2012); ha-

bitus

ostreiform; hymenophore daedaleoid-lamella-

te; basidiospores cylindrical .....Artolenzites.

2 Upperside tomentose — trichodermoid with differentia-

ted subpellis (type b according to Welti et al., 2012);

habitus trametoid; hymenophore radially arranged

poroid to lamellate; basidiospores cylindrical......

...................Lenzites.

2 Upperside strigose to tomentose; subpellis differentiated

or not; habitus corioloid to trametoid; hymenophore

poroid; basidiospores cylindrical or ellipsoid . . . .

................Trametes s. str.

POLYPORALES Gäum., Vergl. Morphol. Pilze: 503,

1926.

MERULIACEAE Rea, British Basid.: 620, 1922

(incl. Steccherinaceae Parmasto, Consp. syst. Cort.: 169,

1968).

TRAMETOPSIS Tom&sovsk*y, Czech Mycol. 60: 8,

2008.

Trametopsis cervina (Schwein.) Tom&sovsk*y, Czech

Mycol. 60: 8, 2008 (Syn.: Boletus cervinus Schwein.,

1822; Polyporus squarrosus Berk. et M. A. Curtis, 1872;

Trametes populina Bres., 1896; Coriolus orizabensis Mur-

rill, 1912).

DIPLOMITOPORUS Doma*nski, Acta Soc. Bot. Po-

lon. 39: 191, 1970.

Diplomitoporus crustulinus (Bres.) Doma*nski, Acta

Soc. Bot. Polon. 39: 191, 1970 (Bas.: Poria crustulina

Bres., 1925).

D. flavescens (Bres.) Doma*nski, Acta Soc. Bot. Polon.

39: 191, 1970 (Bas.: Trametes flavescens Bres. 1903; Syn.:

Polyporus winogradowii Bondartsev, 1912).

Provisional position: D. allantosporus Ryvarden et

Iturr., 2003; D. cunninghamii P. K. Buchanan et Ryvarden,

1998; D. hondurensis (Murrill) Ryvarden, 2000; D. mic-

rosporus Iturr. et Ryvarden, 2010; D. navisporus Gibertoni

et Ryvarden, 2004; D. overholtsii (Pil*at) Gilb. et Ryvar-

den, 1985; D. rimosus (Murrill) Gilb. et Ryvarden, 1985;

D. stramineus Ryvarden et Iturr., 2003; D. venezuelicus

Ryvarden et Iturr., 2003.

CERRENA Gray, Nat. Arr. British Pl. 1: 649, 1821.

Cerrena

aurantiopora J. S. Lee et Y. W. Lim, Mycolo-

gia 102: 212—213, 2010.

C. consors (Berk.) K. S. Ko et H. S. Jung, FEMS Mi-

crobiol. Lett. 170: 185, 1999 (Bas.: Irpex consors Berk.,

1877).

C. cystidiata Rajchenb. et De Meijer, 1990.

C. unicolor (Bull.) Murrill, J. Mycol. 9: 91, 1903

(Bas.: Boletus unicolor Bull., 1785; Syn.: Sistotrema cine

reum Pers., 1794; Hericius strigiliformis Dubois, 1803;

Polyporus latissimus Fr., 1815; P. argyraceus Pers., 1825;

Phyllodontia magnusii P. Karst., 1883; Daedaleopsis inca

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 373

* Weakly amyliod in Pycnoporus.

na P. Karst., 1904; Buillardia nigrozonata L*azaro Ibiza,

1916; Polyporus rohlenae Velen., 1922).

PANUS Fr., Epicr. Syst. Mycol.: 396, 1838.

Panus ciliatus (L*ev.) T. W. May et A. E. Wood, Myco

taxon 54: 148, 1995 (Bas.: Lentinus ciliatus L*ev., 1844;

Syn.: L. melanophyllus L*ev., 1844; L. setiger L*ev., 1844;

L. echinopus L*ev., 1846; L. braccatus L*ev. in Zolling.,

1854; L. dichrous L*ev. in Zolling., 1854; L. zonifer Berk.

et Broome, 1873; L. egregius Massee, 1910; Panus brun

neipes Corner, 1981).

P. conchatus (Bull.: Fr.) Fr., Epicr.: 397, 1838 (Bas.:

Agaricus conchatus Bull., 1786; Syn.: A. torulosus Pers.,

1801; A. inconstans Pers., 1801; Panus monticola Berk.,

1851; P. vaporarius Baglietto, 1865; Lentinus percomis

Berk. et Broome, 1873; L. divisus Schulz., 1879; L. bresa

dolae Schulz., 1885; L. obconicus Peck, 1906).

P. fasciatus (Berk.) Singer, Agaricales mod. Tax. 2nd

ed.: 172, 1962 (Bas.: Lentinus fasciatus Berk., 1840; Syn.:

L. dealbatus Fr. in Lehmann, 1847; L. fuscopurpureus

Kalchbr., 1880; L. holopogonius Berk. ex Cooke, 1892;

L. terrestris Lloyd, 1925).

P. hookerianus (Berk.) T. W. May et A. E. Wood, My

cotaxon 54: 148, 1995 (Bas.: Lentinus hookerianus Berk.,

1851).

P. lecomtei (Fr.) Corner, Beih. Nova Hedw. 69: 90,

1981 (Bas.: Lentinus lecomtei Fr., 1825; Syn.: Agaricus

flabelliformis Schaeff., 1762 non Bolton ex Fr., 1821;

A. strigosus Schwein., 1822; Panus rudis Fr., 1838; Agari-

cus sainsonii L*ev., 1842; Lentinus chaetophorus L*ev.,

1844; Panus lamyanus Mont., 1856; P. hoffmannii Fr.,

1865; Lentinus sparsibarbis Berk. et M. A. Curtis, 1868;

L. substrigosus P. Henn. et Shirae, 1900; L. metatensis

Bacc., 1917; Panus semirudis Singer, 1936; P. fragilis

O. K. Miller, 1965).

P. similis (Berk. et Broome) T. W. May et A. E. Wood,

Mycotaxon 54: 148, 1995 (Bas.: Lentinus similis Berk. et

Broome, 1873; Syn.: L. velulinus Fr. var. africanus

P. Henn. in Engler, 1893; L. erringtohnii Pat., 1900; L. sa

murensis Pil*at, 1941).

P. strigellus (Berk.) Chardon et Toro, Monogr. Univ.

Porto Rico Ser. B 2: 315, 1934 (Bas.: Lentinus strigellus

Berk., 1868; Syn.: Panus guaraniticus Speg., 1883; Len

tinus crispus Pat., 1889; L. tubarius Pat., 1899; Pocil

laria palmeri Earle, 1906; Lentinus subglaber Lloyd,

1917).

P. tephroleucus (Mont.) T. W. May et A. E. Wood,

Mycotaxon 54: 148, 1995 (Bas.: Lentinus tephroleucus

Mont., 1851; Syn.: L. leprieurii Mont., 1854; L. siparius

Berk. et M. A. Curtis, 1868; L. dentatus Bres., 1925).

P. velutinus (Fr.) Overh., J. Dept Agr. Porto Rico 14:

353, 1930 non Fr., 1838 (Bas.: Lentinus velutinus Fr.,

1830; Syn.: Lentinus fulvus Berk., 1842; L. coelopus L*ev.,

1846; L. nepalensis Berk., 1854; L. blepharodes Berk. et

M. A. Curtis, 1868; L. fastuosus Kalchbr. et MacOwan,

1881; L. fallax Speg., 1883; L. castaneus Ellis et Macbr.,

1896; L. holumbrinus De Seynes, 1897; L. fissus Henn.,

1897; L. natalensis

Van der Byl, 1924; L. thomensis Cou

tinho, 1925; L. pseudociliatus Raithelhuber, 1974).

Provisional position: Lentinus martianoffianus

Kalchbr. in Thüm., Bull. Mosk. Obshch. Ispyt. Prir. 52:

144, 1877; L. hirtiformis Murrill, N. Amer. Fl. 9: 293,

1915; L. courtetianus Har. et Pat., Bull. Mus. Hist. Nat. Pa

ris 15: 88, 1909.

POLYPORACEAE Corda, Ic. Fung. 3: 49, 1839.

POLYPOROIDEAE Fr., Syst. Orbis Veg.: 79, 1825

(corresponded to «polyporoid radiation» in molecular

trees).

DATRONIA Donk, Persoonia 4: 337, 1966.

Datronia mollis (Sommerf.) Donk, Persoonia 4: 338,

1966 (Bas.: Daedalea mollis Sommerf., 1826; Syn.: Poly

porus cervinus Pers., 1825 non Schwein.; Trametes ser

pens Fr., 1849 non Polyporus serpens Fr., 1818; Polyporus

sommerfeldtii P. Karst., 1878).

D. scutellata (Schwein.) Gilb. et Ryvarden, Mycota

xon 22: 364, 1985 (Bas.: Polyporus scutellatus Schwein.,

1834; Syn.: Fomes atroalbus Henn. et E. Nyman, 1900;

Trametes nigrescens Bres., 1905).

D. stereoides (Fr.) Ryvarden, Blyttia 25: 168, 1967

(Bas.: Polyporus stereoides Fr., 1818; Syn.: Trametes epi

lobii P. Karst., 1868; Polyporus planus Peck, 1878; Dae

dalea lassbergii Allesch., 1889; Coriolus planellus Mur

rill, 1905; Coriolellus planellus Murrill, 1906; Trametes

sendaiensis Yasuda, 1923).

Provisional position: D. decipiens (Bres.) Ryvarden,

Mycotaxon 33: 308, 1988.

EARLIELLA Murrill, Bull. Torrey Bot. Cl. 32: 478,

1905.

Earliella scabrosa (Pers.) Gilb. et Ryvarden, Mycota-

xon 22; 364, 1985 (Bas.: Polyporus scabrosus Pers., 1827;

Syn.: P. fuscobadius Pers., 1827; P. corrugatus Pers.,

1827; Daedalea sanguinea Klotzsch, 1833; Polyporus per-

soonii

Mont., 1834; P. indecorus Jungh., 1838; P. venulo-

sus Jungh., 1838; Daedalea microsinuosa Klotzsch et

Berk., 1839; Polyporus tostus Berk., 1844; Hexagonia cru-

enta Mont., 1845; Polyporus tegularis L*ev., 1846; Trame-

tes hookeri Berk., 1854; Daedalea emodensis Berk., 1854;

Trametes versiformis Berk. et Broome, 1872; Hexagonia

picta Berk., 1878; Polyporus aruensis Berk., 1878; Trame

tes moselei Berk., 1878; Polystictus parishii Berk. ex Coo

ke, 1886; Trametes nitida Pat., 1890; Daedalea conchata

Bres., 1890; D. andamani Berk. ex Cooke, 1891; Polystic

tus rufocinerescens Henn., 1899; Earliella cubensis Mur

rill, 1905; Favolus subrigidus Murrill, 1908; Trametes

zimmermanii Bres., 1920 ut ‘zimmermani’).

MICROPORUS Beauv., Fl. Oware Benin Afr. 1: 12,

1805.

Microporus affinis (Blume et T. Nees) Kuntze, Rev.

Gen. Pl. 3: 494, 1898 (Bas.: Polyporus affinis Blume et

T. Nees, 1826; Syn.: P. luteus Blume et T. Nees, 1826;

P. flabelliformis Klotzsch, 1833; P. crenatus Berk., 1843;

P. microloma L*ev., 1844; P. squamaeformis Berk., 1854;

P. carneoniger Berk. ex Cooke, 1883; P. trizonatus Coo

ke, 1883; Polystictus subaffinis Lloyd, 1916).

M. quarrei (Beeli) D. A. Reid, Microscopy 32: 453,

1975 (Bas.: Polyporus quarrei Beeli, 1922).

M. vernicipes (Berk.) Kuntze, Rev. Gen. Pl. 3: 497,

1898 (Bas.: Polyporus vernicipes Berk., 1878; Syn.: Cori

olus langbianensis Har. et Pat., 1914; C. subvernicipes

Murrill, 1908).

M. xanthopus (Fr.) Kuntze, Rev. Gen. Pl. 3: 494, 1898

(Bas.: Polyporus xanthopus Fr., 1818; Syn.: P. saccatus

Pers., 1827; P. pterygodes Fr., 1838).

374 I. V. Zmitrovich, V. F. Malysheva

DAEDALEOPSIS J. Schröt. in Cohn, Krypt. Fl.

Schles. 3: 492, 1888.

Daedaleopsis confragosa (Bolton) J. Schröt. in Cohn,

Krypt. Fl. Schles. 3: 492, 1888 (Bas.: Boletus confragosus

Bolton, 1792; Syn.: B. labyrinthiformis Bull., 1790; B. an

gustatus Sowerby, 1799; Daedalea suaveolens Pers.,

1801; Boletus horizontalis Thore, 1803; Daedalea rubes

cens Alb. et Schwein., 1805; D. pruinata Secr., 1833; Tra

metes erubescens S. Schulz., 1882; T. zonata Wettst., 1885

non Nees; Daedalea ochracea Velen., 1922).

Daedaleopsis nitida (Durieu et Mont.) Zmitr. et

V. Malysheva comb. nov. (Bas.: Hexagonia nitida Durieu

et Mont., Syll. Gen. Sp. Crypt.: 170, 1856; MB 803404).

D. purpurea (Cooke) Imazeki et Aoshima in Hara, Fl.

East. Himal.: 619, 1966 (Bas.: Trametes purpurea Cooke,

1882; Syn.: Daedaleopsis nipponica Imazeki, 1943).

D. septentrionalis (P. Karst.) Niemelä, Karstenia 22:

11, 1982 (Bas.: Lenzites septentrionalis P. Karst., 1866).

D. tenuis (Hook. in Kunth) Imazeki, Bull. Tokyo Sci.

Mus. 6: 78, 1943 (Bas.: Boletus tenuis Hook. in Kunth,

1822).

D. tricolor (Bull ex M*erat) Bondartsev et Singer, Ann.

Mycol. 39: 64, 1941 (Bas.: Agaricus tricolor Bull ex

M*erat, 1791; Syn.: Lenzites atropurpurea Sacc., 1873).

Provisional position: D. papyroresupinata (S. Ito et

S. Imai) Imazeki, Bull. Tokyo Sci. Mus. 6: 78, 1943;

D. pergamenea (Berk. et Broome) Ryvarden, Mycotaxon

20: 350, 1984; D. sinensis (Lloyd) Y. C. Dai, Fungal Sci-

ence, Taipei 11: 90, 1996; Trametes mimetes (Wakef.) Ry-

varden, Norw. J. Bot. 19: 236, 1972; T. salina Corner,

Beih. Nova Hedw. 97: 149, 1989.

HEXAGONIA Fr., Fl. Scan.: 339, 1835; nom.con-

serv.

Hexagonia apiaria (Pers. in Freyc.) Fr., Epicr.: 497,

1838 (Bas.: Polyporus apiarius Pers. in Freyc., 1827; Syn.:

Hexagonia koenigii Berk., 1843; H. henshallii

Lloyd,

1910).

H. hirta (P. Beauv.) Fr., Epicr.: 496, 1838 (Bas.: Favo

lus hirtus P. Beauv., 1805; Syn.: Boletus favus L., 1753;

Polyporus sinensis Fr., 1821; Trametes crassa L*ev., 1844;

T. hystrix Cooke, 1881; T. adelphica Cooke, 1891; Hexa

gonia atra Lloyd, 1822; H. pectinata Lloyd, 1825).

H. niam-niamensis Henn., Engler’s Bot. Jahrb. 14:

348, 1892 (Syn.: H. nigrocincta Pat., 1906).

H. speciosa Fr., K. Vet. Akad. Handl.: 137, 1848

(Syn.: H. obversa Pat., 1891; H. crassispora Lloyd, 1917;

H. subvelutina Wakef., 1917; H. sericata Wakef., 1922).

POGONOMYCES Murrill, Bull. Torrey Bot. Cl. 31:

609, 1904.

Pogonomyces hydnoides (Sw.) Murrill, Bull. Torrey

Bot. Cl. 31: 609, 1904 (Syn.: Boletus hydnoides Sw., 1806;

B. ursinus Link, 1809; B. fibrosus Hook. in Kunth, 1822;

Trametes ocellata Berk. et M. A. Curtis, 1868; Polyporus

verrucosohirtus Speg., 1884).

FUNALIA Pat., Essai taxon. Hym.: 95, 1900.

Funalia aspera (Jungh.) Zmitr. et V. Malysheva

comb. nov. (Bas.: Polyporus asper Jungh., Praem Fl.

Cryptog. Javae: 60, 1838; Syn.: P. koenigii Berk., 1839;

Trametes badia Berk., 1842; Polyporus confertus L*ev.,

1844; P. cohaerens L*ev., 1846; P. strigatus Berk., 1847;

P. fuscellus L*ev., 1854; Trametes pyrrhocreas Berk.,

1872; Polyporus lineatoscaber Berk. et Broome, 1882;

P. arenosus Cooke, 1884; P. curreyi Berk. ex Cooke,

1886; P. olivaceus Rostr., 1902; P. zebra Lloyd, 1915;

P. rugososporus Lloyd, 1915; Hexagonia lignosa Lloyd,

1919;

Polyporus fijii Lloyd, 1922; MB 803405).

Funalia caperata (Berk.) Zmitr. et V. Malysheva

comb. nov. (Bas.: Polyporus caperatus Berk., Grevillea 9:

98, 1881; Syn.: Polyporus vespilloneus Berk., 1856; P. cir

rifer Berk. et M. A. Curtis, 1868 ut ‘cirriferus‘; Trametes

dibapha Berk., 1880; Polyporus purpureobadius Pat.,

1892; Polystictus ekunduensis Henn., 1895; P. fischeri

Henn., 1898; Coriolopsis fulvocinerea Murrill, 1908;

C. subglabrescens Murrill, 1908; Polystictus umbrinus

Bres., 1912; Coriolopsis caperatiformis Murrill, 1912; Po

lyporus euzonatus Lloyd, 1919; MB 803406).

Funalia floccosa (Jungh.) Zmitr. et V. Malysheva

comb. nov. (Bas.: Polyporus floccosus Jungh., Praem. Fl.

Crypt. Javae: 49, 1838; Syn.: P. proteus Berk., 1843;

P. nigrocinctus Berk., 1843; P. capitosus Mont., 1847;

Trametes rigida Berk. et Mont., 1849; Polyporus excur

rens Berk. et M. A. Curtis, 1868; Trametes acuta Cooke,

1882; Polystictus ecklonii Berk. ex Cooke, 1886; Trametes

carteri Berk. ex Sacc., 1891; Coriolus chudaei Pat., 1907;

Polyporus rusticus Lloyd, 1918; P. flexilis Van der Byl,

1922; Hexagonia velutinoglabra Lloyd, 1923; Polyporus

livingstoniensis Van der Byl, 1925; Trametes felliponei

Bres., 1926; Fomes palustris Rick, 1960; MB 803407).

F. gallica (Fr.) Bondartsev et Singer, Ann. Mycol.

339: 62, 1941 (Bas.: Polyporus gallicus Fr., 1821; Syn.:

Boletus favus Bull., 1788 nom. inval.; Poria vulgaris Gray,

1821; Trametes stuppea Berk., 1841; Polyporus extenua-

tus Durieu et Mont., 1856; Trametes hispida Bagl., 1865;

Polyporus schulzeri Kalchbr., 1868; P. pulcher Speg.,

1880; Trametes peckii Kalchbr. in Peck, 1881; Hexagonia

favus Qu*el., 1888 nom. inval.).

F. leonina (Klotzsch) Pat., Essai.: 95, 1900 (Syn.: Po-

lyporus leoninus Klotzsch, 1839; P. mons-veneris Jungh.,

1838).

Funalia sanguinaria (Klotzsch) Zmitr. et V. Maly

sheva comb. nov. (Bas.: Polyporus sanguinarius Klotzsch,

Linnaea 8: 484, 1833; Syn.: P. bicolor Jungh., 1838; P. ru

gulosus L*ev., 1844; P. anisopilus L*ev., 1844; P. vulnera

tus L*ev., 1844; P. trigonus L*ev., 1846; P. anebus Berk.,

1847; P. plebeius Berk., 1855 ut ‘plebejus’; P. beckleri

Berk., 1872; P. polytropus Berk. et Broome, 1875; Fomes

oblinitus Berk. ex Cooke, 1886; Hapalopilus subrubidus

Murrill, 1907; Polyporus subpruinatus Bres., 1912; Tra

metes lignicolor Bres., 1920; Polyporus brunneomacula

tus Lloyd, 1920; P. biogilvus Lloyd, 1920; P. loreus Beeli,

1922; P. retroater Lloyd, 1922; Trametes retropicta Llo

yd, 1922; Polyporus bosei Bres., 1926; MB 803408).

F. trogii (Berk. in Trog) Bondartsev et Singer, Ann.

Mycol. 39: 62, 1941 (Bas.: Trametes trogii Berk. in Trog,

1850; Syn.: Polyporus ozonoides Berk., 1851; Daedalea

trametes Speg., 1880; Inodermus maritimus Qu*el., 1887;

Trametes tucumanensis Speg., 1899).

Provisional position: Trametes biogilvoides Corner,

Beih. Nova Hedwigia 97: 81, 1989; T. brunneoleuca

(Berk.) Corner, Beih. Nova Hedw. 97: 84, 1989;

T. byrsina

(Mont.) Pat., Essai.: 93, 1900; T. drummondii (Klotzsch)

Ryvarden, Mem. N. Y. bot. Gdn 28: 202, 1976; T. glabro

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 375

rigens (Lloyd) Zmitr., Wasser et Ezhov, Int. J. Med. Mush

rooms 14: 315, 2012; T. hostmannii (Berk.) Zmitr., Wasser

et Ezhov, Int. J. Med. Mushrooms 14: 317, 2012; T. mo

desta (Kunze ex Fr.) Ryvarden, Norw. J. Bot. 19: 236,

1971; T. paxillosa Corner, Beih. Nova Hedw. 97: 130,

1989; T. strumosa (Fr.) Zmitr., Wasser et Ezhov, Int. J.

Med. Mushrooms 14: 318, 2012; T. telfairii (Klotzsch)

Corner, Beih. Nova Hedw. 97: 167, 1989.

TRAMETOIDEAE Pinto-L., Mem. Soc. Brot. 8: 161,

1952 (corresponded to ‘trametoid radiation’ in molecular

trees).

PYCNOPORUS P. Karst., Rev. Mycol. 3: 18, 1881.

Pycnoporus cinnabarinus (Jacq.) P. Karst., Rev. My-

col. 3: 18, 1881 (Bas.: Boletus cinnabarinus Jacq., 1776;

Syn.: B. coccineus Bull. 1790 non Fr., 1838; B. miniatus

Libosch., 1817).

P. puniceus (Fr.) Ryvarden, Norw. J. Bot. 19: 236,

1972 (Bas.: Trametes punicea Fr., 1851).

P. sanguineus (L.) Murrill, Bull. Torrey Bot. Cl. 31:

421, 1904 (Bas.: Boletus sanguineus L., 1763; Syn.: B. ru-

ber Lam., 1783; Polyporus coccineus Fr., 1851).

CUBAMYCES Murrill, Bull. Torrey Bot. Cl. 32: 480,

1905.

Cubamyces cubensis (Mont.) Murrill, Bull. Torrey

Bot. Cl. 32: 480, 1905 (Bas.: Polyporus cubensis Mont.,

1837; Syn.: Polyporus lactineus Berk., 1843; P. ostreatus

L*ev. 1846; Trametes levis Berk., T. subcubensis Murrill,

1939.

Provisional position: Trametes menziesii (Berk.)

Ryvarden, Norw. J. Bot. 19(3): 236, 1972; Trametes lju-

barskyi Pil*at, Bull. trimestr. Soc. Mycol. France 52: 309,

1937; T. marianna (Pers.) Ryvarden, Persoonia 7: 309,

1973.

CELLULARIELLA Zmitr. et V. Malysheva gen.

nov.

Trametoidea, eusomica, cum hymenophorum daeda-

leoideum ad lenzitoideum; sclerohyphae hyalinae, occasio

sympodialiter ramosae, sine depositae sed irregulariter in-

crustatae. Basidia clavata, sporae cylindraceae.

Typus: Lenzites acuta Berk., London J. Bot. 1: 146,

1842.

Trametoid, large-sized; hymenophore daedaleoid to

lamellate; sclerohyphae hyaline, sympodially and rarely

branched, inamyloid, without deposites, in some cases

with crystalline encrustation; context cream. Basidia cla-

vate, basidiospores cylindrical.

Cellulariella acuta (Berk.) Zmitr. et V. Malysheva

comb. nov. (Bas.: Lenzites acuta Berk.,

London J. Bot. 1:

146, 1842; Syn.: Daedalea inaequabilis Berk., 1843; Poly-

porus poecilus Berk., 1843; Daedalea flavida L*ev., 1844;

D. pruinosa L*ev., 1844; D. lurida L*ev., 1844; Lenzites mu-

rina L*ev., 1844; Sistotrema ochroleucum L*ev., 1846; Dae-

dalea microzona L*ev., 1846; Hexagonia glabra L*ev.,

1846; Trametes colliculosa Berk., 1847; T. lobata Berk.,

1851; Polyporus lenziteus L*ev. in Zolling., 1854; Lenzites

ochrophylla Berk., 1854; Daedalea aulacophylla Berk.,

1872; D. hobsonii Berk., 1872; Lenzites beckleri Berk.,

1872; Trametes rugosa Berk. et Broome, 1875; Daedalea

subsulcata Berk. et Broome, 1875; Lenzites guilfoylei

Berk. ex Cooke, 1881; Daedalea scalaris Berk. et Broome,

1882; Lenzites alutacea Cooke, 1883; Daedalea schom

burgkii Berk. ex Sacc., 1888; D. flabellum Berk. ex Cooke,

1891; Lenzites submurina Murrill, 1908; L. clemensiae

Murrill, 1908; Daedalea subconfragosa Murrill, 1908;

Lenzites adusta Massee, 1910; Daedalea ridlei Lloyd,

1920; Trametes transmutans Lloyd, 1920; Lenzites albolu-

tea Lloyd, 1920; Daedalea bosei Lloyd, 1922; Hexagonia

flavofusca Lloyd, 1922; Lenzites huensis Lloyd, 1922; He-

xagonia murina Lloyd, 1923; Daedalea glabra Lloyd,

1924).

Cellulariella

warnieri (Dur. et Mont.) Zmitr. et

V. Malysheva comb. nov. (Bas.: Lenzites warnieri Durieu

et Mont., Ann. Sci. Nat., Bot. 14: 182, 1860; Syn.: L. fa-

ventina Caldesi, 1869; L. reichardtii Schulz. in Thum.,

1880).

CORIOLOPSIS Murrill, Bull. Torrey Bot. Cl. 32:

358, 1905.

Coriolopsis polyzona (Pers.) Ryvarden, Norw. J. Bot.

19: 230, 1972 (Syn.: Polyporus polyzonus Pers., 1827;

P. lundii Fr., 1828; P. torridus Fr., 1828; P. occidentalis

Klotzsch, 1833; Trametes lanata Fr., 1838; P. scytinus

Berk., 1843; P. lenis L*ev., 1848; Trametes scalaris Fr.,

1849; Polystictus cyclodes Fr., 1851; P. scorteus Fr., 1851;

Trametes devexa Berk., 1872; T. heteromalla Cooke,

1882; T. andina Pat., 1895; T. ochrolignea Lloyd, 1916;

Fomes wombaliensis Beeli, 1822; Trametes tomentosa

Van der Byl, 1922).

Provisional position: Trametes variegata (Berk.)

Zmitr., Wasser et Ezhov, Int. J. Med. Mushrooms 14: 315,

2012.

SCLERODEPSIS Cooke, Grevillea 19: 49, 1890.

Sclerodepsis berkeleyi Cooke, Grevillea 19: 49, 1890.

S. maxima (Mont.) Ryvarden, Norw. J. Bot. 19: 236,

1972 (Bas.: Irpex maximus Mont., 1837).

S. meyenii (Klotzsch) Ryvarden, Norw. J. Bot. 19: 236,

1972 (Bas.: Polyporus meyenii Klotsch, 1843; Syn.: Dae-

dalea dregeana Mont., 1847; D. ochracea Kalchbr., 1876;

D. incompta Berk., 1882; T. obstinata Cooke, 1883; T. cor-

nea Pat., 1890; Daedalea papyracea Massee, 1907;

D. hobbsii Van der Byl, 1924; Trametes capensis Lloyd,

1950).

ARTOLENZITES Falck, Hausswammforsch 3: 37,

1909.

Artolenzites elegans (Spreng.) Teixeira, Rev. Bras.

Bot. 13: 138, 1986 (Syn.: Daedalea elegans Spreng.,

1820;

D. palisoti Fr., 1821; D. levis Hook. in Kunth, 1822; Bole-

tus aesculi-flavae Schwein., 1822; Daedalea repanda

Pers. in Gaudich., 1827; Polyporus aesculi Fr., 1828; Dae-

dalea polita Fr., 1830; D. applanata Klotzsch, 1833; Len-

zites pallida Berk., 1842).

LENZITES Fr., Fl. Sc.: 339, 1835.

Lenzites betulina (L.) Fr., Epicr.: 405, 1838 (Bas.:

Agaricus betulinus L., 1753; Syn.: A. hirsutus Schaeff.,

1774; A. tomentosus Lam., 1778; A. flaccidus Bull., 1788;

A. coriaceus Bull., 1792; Boletus versicolor Vent., 1812;

Daedalea variegata Fr., 1818; D. interrupta Fr., 1828;

Lenzites umbrina Fr., 1838; L. junghuhnii L *ev., 1844;

L. berkeleyi L*ev., 1846; L. cinnamomea Fr., 1851; L. pi

nastri Kalchbr., 1874; L. sorbina P. Karst., 1881; L. cyc

logramma Pat., 1907; L. subbetulina Murrill, 1912; L. con

nata L*azaro Ibiza, 1916; L. hispida L*azaro Ibiza, 1916;

L. ochracea Lloyd, 1922; L. pertenuis Lloyd, 1922).

376 I. V. Zmitrovich, V. F. Malysheva

L. gibbosa (Pers.) Hemmi, Ann. Phytopath. Soc. Japan

9: 12, 1939 (Bas.: Merulius gibbosus Pers., 1795; Syn.:

Agarico-suber scalptum Paulet, 1793; Trametes crenulata

Berk., 1854; T. kalchbrenneri Fr., 1868; Bulliardia vires

cens L*azaro Ibiza, 1916).

Provisional position: Lenzites vespacea (Pers.) Pat.,

Essai.: 91, 1900; Trametes barbulata Corner, Beih. Nova

Hedwigia 97: 75, 1989; T. benevestita Corner, Beih. Nova

Hedwigia 97: 77—78.

TRAMETES Fr. Fl. Sc.: 339, 1835.

Trametes cingulata Berk., Hooker’s J. Bot. 6: 164,

1854 (Syn.: Polyporus palliseri Berk. ex Cooke, 1882; Fo-

mes virginiae Mancini et Sacc., 1888; Trametes cyclo-

phaea Pat., 1906; Coriolus decorsei Har. et Pat., 1909; Po-

lyporus griseus Bres., 1912; Trametes jalapensis Murrill,

1912; Polyporus granulatus Lloyd, 1922).

T. conchifera (Schwein.) Pil*at in Kavina et Pil*at, Atl.

Champ. Eur. 3: 264, 1939 (Bas.: Boletus conchifer Schwe-

in., 1822).

T. ectypa (Berk. et M. A. Curtis) Gilb. et Ryvarden,

N. Amer. Polyp. 2: 740, 1987 ut ‘ectypus’ (Bas.: Polypo-

rus ectypus Berk. et M. A. Curtis, 1872; Syn.: P. farinosus

Rick, 1907; P. farinosus Lloyd, 1915).

T. hirsuta (Wulfen) Lloyd, Mycol. writ. 7: 1319, 1924

(Bas.: Boletus hirsutus Wulfen in Jacq., 1788; Syn.: B. ve-

lutinus J. J. Planer, 1788; B. wulfenii Humb., 1793; B. nig-

romarginatus Schwein., 1822; Polyporus vellereus Berk.,

1842; P. galbanatus Berk., 1843; P. cinerescens L*ev.,

1844; P. cinereus L*ev., 1846; P. gourliei Berk., 1860; Co-

riolus

velutinus P. Karst., 1906; Trametes porioides L*aza-

ro Ibiza, 1917; Polyporus fagicola Velen., 1922).

T. junipericola Manjon, Moreno et Ryvarden, Boll.

Soc. Micol. Castellana 8: 47, 1984.

T. manila¸nsis (Lloyd) Teng, Fung. China: 763, 1963

(Bas.: Polyporus manila¸nsis Lloyd, 1918).

T. membranacea (Sw.) Kreisel, Monogr. Cien. Univ.

Havana 16: 83, 1971 (Bas.: Boletus membranaceus Sw.,

1788).

T. multicolor (Schaeff.) Julich, Persoonia 11: 427,

1982 (Bas.: Boletus multicolor Schaeff., 1774; Syn.:

B. ochraceus Pers., 1794; B. zonatus Nees, 1817; Coriolus

concentricus Murrill, 1907; C. lloydii Murrill, 1907; Bulli-

ardia rufescens L*azaro Ibiza, 1916; Polyporus aculeatus

Velen., 1922; Trametes zonatella Ryvarden, 1978).

T. orientalis (Yasuda) Imazeki, Bull. Tokyo Sci. Mus.

6: 73, 1943 (Bas.: Polystictus orientalis Yasuda, 1918).

T. pocas (Berk.) Ryvarden, Mycotaxon 20: 351, 1984

(Bas.: Polyporus pocas Berk., 1878; Syn.: P. subpelluci-

dus Berk., 1878).

T. pubescens (Schumach.) Pil*at in Kavina et Pil*at, Atl.

Champ. Eur. 3: 268, 1939 (Bas.: Boletus pubescens Schu-

mach., 1803; Syn.: B. velutinus Pers., 1794; Polyporus sul-

livantii Mont., 1842; P. molliusculus Berk., 1847; Hanse-

nia imitata P. Karst., 1886; Coriolus applanatus P. Karst.,

1904).

T. sediliensis Corner, Beih. Nova Hedw. 97: 159,

1989.

T. suaveolens (Fr.) Fr., Epicr.: 491, 1838 (Bas.: Poly

porus suaveolens Fr., 1828; Syn.: Boletus discoideus

Dicks., 1793; Polyporus itoi Lloyd, 1924; Trametes radia

ta Burt, 1931).

T. subectypa (Murrill)

Gilb. et Ryvarden, N. Amer. po

lyp. 2: 758, 1987 ut ‘subectypus’ (Bas.: Coriolus subecty

pus Murrill, 1907).

T. supermodesta Ryvarden et Iturr., Mycologia 95:

1074, 2003.

T. tephroleuca Berk., Hooker’s J. Bot. 6: 165, 1854

(Syn.: T. immutata Berk., 1854).

T. versicolor (L.) Lloyd, Mycol. notes 65: 1045, 1921

(Bas.: Boletus versicolor L., 1753; Syn.: Polyporus fusca-

tus Fr., 1818; P. nigricans Lasch in Rabenh., 1859).

T. villosa (Sw.) Kreisel, Monogr. Cien. Univ. Havana

16: 83, 1971 (Bas.: Boletus villosus Sw., 1806; Syn.: Poly-

porus pinsitus Fr., 1828; P. sericeohirsutus Klotzsch,

1833; P. gibberulosus L*ev., 1846; P. tener L*ev., 1846; Po-

lystictus barbatulus Fr., 1851; P. umbonatus Fr. in Speg.,

1851; Hexagonia friesiana Speg., 1884; Polystictus con-

sobrinus Speg., 1889; Polyporus sericeus Pat., 1895; Co-

riolus fulvoumbrinus Murrill, 1907; Favolus dussii Lloyd,

1923).

Trametes pubescens (Schumach.) Pil *at var. anthopo-

ra Zmitr., N. Bukharova et V. Malysheva var. nov. —

MB 803403; Fig. 6.

Basidiomata copiosa, corioloidea, leptosomica ad

eusomica (15—60 % 2—9 mm), infundifuliformia, flabelli-

formia, conchata, spathulata vel petaloidea, semicirculata

ad reniformia, normaliter basi attenuata ad substipitata.

Superficio abhymenophoralis sine subpellis, villosa, radi-

aliter rugosa et sulcata, cremea ad lignosa, cum zonatio

obscura. Caro nivea, homgenea, ad 4 mm crass. Hyme-

nophorum poroideum, tenuiseptatum, unistratosum, ad

6 mm crass. Porae normaliter 4—6(7 ) per mm, quandoque

disruptae ad 1—3 per mm, angulatae ad anthomorphosae,

sinuosae, cremicolor (citrescens) ad carnosae et lignicolor

in statu sicco. Systema hypharum dimiticum cum sclero-

hyphae sympodialiter ramosae. Hyphae generatoriae

2—4 mm diam., fibulatae, tenuitunicatae. Sclerohyphae

2—9.5 mm diam., crassitunicatae, hyalinae, acyanophilae,

inamyloideae. Basidia 14—20 % 4—6 mm,

clavatae, 4-ste-

rigmatae, basi fibulatae. Cystidia nulla. Sporae

4.5—5.5 % 1.5—2.2 mm, cylindraceae ad suballantoideae

vel lacrymoideae, tenuitunicatae, hyalinae, inamyloideae,

acyanophilae.

Forma typica similis est, sed porae euis 3—5 per mm

et sporae 5—6 % 2—2.5 mm.

Basidiomata of corioloid habitus, small- to medium-si-

zed (15—60 % 2—9 mm), rosette-like, flabelliform, conc-

hate, spathulate or petaloid, semicircular to reniform in

outline, in most of cases with narrow base; as a rule cluste-

red, sessile or stipitate. Upperside without a differentiated

subpellis, velvety (slightly villose), radially ridged and

concentrically sulcate, cream to tan, with obscure pale-co-

lored zones. Context purely white, rather loose, homoge-

neous, up to 4 mm thick. Hymenophore poroid, as a single

layer up to 6 mm thick. Pores normally 4—6(7) per mm, in

some regions collapsed to 1—3 per mm, with very thin (up

to 0.2 mm) pergameneous dipssepiments, angular to flo-

wer-like, sinuose in some regions, initially cream, then

with carnose or citrine tints, pale-colored at the maturity.

Hyphal system dimitic with sympodially branched scle

rohyphae. Generative hyphae 2—4 mm in diam., clamped,

thin-walled, rarely branched. Sclerohyphae 2—9.5 mmin

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales) 377

378 I. V. Zmitrovich, V. F. Malysheva

Fig. 6. Trametes pubescens var. anthopora (LE 286999). Above — external habitus of basidiomata: a — upperside, b —

hymenophore side; scale bar — 10 mm. Below — sclerohyphae and basidiospores; scale bar — 10 mm.

diam., thick-walled to subsolid, hyaline, acyanophilic, ina

myloid; the branching pattern sympodial, moderate, most

ly rectangular. Basidia 14—20 % 4—6 mm, clavate, 4-spo

red, with a basal clamp. Cystidia none. Basidiospores

4.5—5.5 % 1.5—2.2 mm, cylindrical, slightly curved or

with a certain lacrymoid tendency, thin-walled, hyaline,

inamyloid, acyanophilous.

Etymology: named after its thin-walled tubes with

some flower-like pores with crispate dissepiments.

Type material:Russia, Far East, Evreiskaya Terri

tory, Bastak Reserve, N 49°02R033S, E 132R56R814S,onfal

len branch of Betula sp. Coll., N. V. Bukharova, 19 Aug.

2010, LE 286999 (holotype); ibid., N 49°02R172S,

E 132°57R194S, on fallen log of Betula sp. Coll., 19 Aug.

2010, N. V. Bukharova, LE 286999 (holotype).

Note.Typical form of T. pubescens is similar, but dif

fers by larger pores (3—5 per mm) with normally thi

ckened dissepiments (more than 0.2 mm) and slightly lar

ger basidiospores (5—6 % 2—2.5 mm). The molecular di

vergence is well corresponded to a species level.

T. orientalis is more distant and differs by more intensive

upperside coloration and subellipsoid basidiospores

5—6 % (2.5)3—3.5 mm.

We thank Dr. E. F. Malysheva for help in molecular

study, Dr. N. V. Bukharova for field collections and dis-

cussion on new species, Dr. A. V. Alexandrova for illu-

strative material of tropical trametoid fungi and Dr.

T. S. Bulgakov and I. A. Kazartsev for some sources re-

commendation.

The financial support for this work was provided by

the Russian Foundation for Basic Research (project

N 12-04-31315 and 12-04-33018).

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Jan 2012
INT J MED MUSHROOMS

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Systematic revisit of Sparsitubus (Basidiomycota, Aphyllophorales), an unusual cyphelloid polypore from China

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Apr 2007
FUNGAL DIVERS

Sparsitubus is a monotypic genus found in East Asia that has a bizarre cyphelloid hymenophore. The position of Sparsitubus in the Polyporaceae has been suggested previously based on morphological characters. To study the relationships of Sparsitubus among other Homobasidiomycetes, rDNA sequences of Sparsitubus and polypores representatives were generated or downloaded from GenBank. Parsimony analyses using rDNA data suggest that Sparsitubus is in a clade of polypores producing a white rot and maybe closely related to the Ganodermataceae. Characters of fruiting body, hyphal systems, and basidiospore were compared and discussed between S. nelumbiformis and related polypores or similar cyphelloid fungi.

The Taxonomical and Nomenclatural Characteristics of Medicinal Mushrooms in Some Genera of Polyporaceae

Article

Full-text available

Jan 2010
INT J MED MUSHROOMS

Ivan V. Zmitrovich

In this article, the taxonomical status of medicinal mushrooms containing taxa representative of Polyporaceae is considered. The results of this study confirmed the genus Ganoderma as a member of the family Polyporaceae. The heterogeneity of the genus Polyporus is also considered. Current literature examination has shown that three more natural genera can be found within the genus Polyporus. Seven new combinations are proposed for the genera Lentinus and Polyporus.

Phylogenetic classification of Trametes (Basidiomycota, Polyporales) based on a five-marker dataset

Article

Full-text available

Dec 2011

The phylogeny of Trametes and related genera was studied using molecular data from ribosomal markers (nLSU, ITS) and protein-coding genes (RPB1, RPB2, TEF1-alpha) and consequences for the taxonomy and nomenclature of this group were considered. Separate datasets with rDNA data only, single datasets for each of the protein-coding genes, and a combined five-marker dataset were analyzed. Molecular analyses recover a strongly supported trametoid clade that includes most of Trametes species (including the type T. suaveolens, the T. versicolor group, and mainly tropical species such as T. maxima and T. cubensis) together with species of Lenzites and Pycnoporus and Coriolopsis polyzona. Our data confirm the positions of Trametes cervina (= Trametopsis cervina) in the phlebioid clade and of Trametes trogii (= Coriolopsis trogii) outside the trametoid clade, closely related to Coriolopsis gallica. The genus Coriolopsis, as currently defined, is polyphyletic, with the type species as part of the trametoid clade and at least two additional lineages occurring in the core polyporoid clade. In view of these results the use of a single generic name (Trametes) for the trametoid clade is considered to be the best taxonomic and nomenclatural option as the morphological concept of Trametes would remain almost unchanged, few new nomenclatural combinations would be necessary, and the classification of additional species (i.e., not yet described and/or sampled for molecular data) in Trametes based on morphological characters alone will still be possible. Alternative scenarios to divide the trametoid clade in five or ten genera were considered but if any of these options were to be adopted morphological distinction of the segregated genera and the ascription of additional species to any of these genera would be very difficult and in some cases impossible. The genera Artolenzites, Coriolopsis (as currently typified), Coriolus, Cubamyces, Cyclomycetella, Lenzites, Poronidulus, Pseudotrametes and Pycnoporus are considered synonyms of Trametes.

Oxidoreductases from Trametes spp. in Biotechnology: A Wealth of Catalytic Activity

Article

Full-text available

Jul 2007

Those oxidoreductases that are part of the ligninolytic complex of basidiomycete and ascomycete fungi have played an increasingly important role in biotechnological applica-tions during the last decade. The stability of these extracellular enzymes, their good solu-bility, and a multitude of catalyzed reactions contribute to this trend. This review focuses on a single genus of white-rot basidiomycetes, Trametes, to highlight the numerous possi-bilities for the application of this microorganism as well as three of its enzymes: laccase, cellobiose dehydrogenase, and pyranose 2-oxidase. Whereas laccase is without doubt a major player in biotechnology, the two other enzymes are less well known, but represent emerging biocatalysts with potential. Both cellobiose dehydrogenase and pyranose 2-oxi-dase are presumed to participate in lignin breakdown and will be used to exemplify the potential of less prominent oxidoreductases from this genus.

Check List of European Polypores

Article

Jan 1975

A critical checklist of the Polyporaceae in tropical East Africa

Article

Jan 1972

L. Ryvarden

North American Polypores

Article

Sep 1986

Check List of European Polypores

Article

Sep 1975

The basidiomycete genus Polyporus - An emendation based on phylogeny and putative secondary structure of ribosomal RNA molecules

Article

Nov 2004

The fungal genus Polyporus is an assemblage of white-rotting lignicolous basidiomycetes. It has undergone considerable expansion and contraction over a period of two and three quarter centuries. Current generic circumscription of Polyporus has kept the genus non-monophyletic. Species of Polyporus infrageneric group Polyporellus are closely related to some species of Lentinus. We introduce data for ITS2 spacer rRNA secondary structure evolution by quasi-independent comparison with large subunit rRNA phylogeny, and suggest a fraction of primary nuclear rDNA ITS sequence data as novel taxonomic character. A major taxonomic shift is suggested, supported by molecular and morphological characters, and allowing inclusion of species with gilled hymenophores in Polyporus. Two new names are proposed: Polyporus phyllostipes D.Krüger, nom. nov. and Polyporus gerdai D.Krüger, nom. nov. Die Gattung Polyporus (Basidiomycetes) – eine Emendation auf der Basis von Phylogenie und mutmaßlicher sekundrer Struktur der ribosomalen RNA-MoleküleDie Pilzgattung Polyporus, eine Gruppe Weißfule erregender holzbewohnender Basidiomyceten, wurde über nahezu drei Jahrhunderte vielfach expandiert und verkleinert. Bei der derzeitigen Gattungsumschreibung von Polyporus gilt die Gattung als nicht-monophyletisch. Arten der Polyporus-Gruppe Polyporellus sind eng verwandt zu einigen Lentinus-Arten. Anhand quasi-unabhngigem Vergleich mit der Phylogenie der rRNA der großen Untereinheit (LSU) stellen wir Daten zur Evolution der ITS2 Spacer rRNA vor, und schlagen ein ITS Kern-rDNA-Fragment als taxonomisches Merkmal vor. Unterstützt mit molekularen und morphologischen Daten wird eine grundlegende taxonomische Verschiebung vorgeschlagen, welche Arten mit Lamellen-Hymenophoren in Polyporus erlaubt. Zwei neue Namen werden vorgeschlagen: Polyporus phyllostipes D.Krüger, nom. nov. und Polyporus gerdai D.Krüger, nom. nov. (© 2004 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Towards a phylogeny of Trametes alliance (Basidiomycota, Polyporales)

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