Phylogenetic classification of Trametes (Basidiomycota, Polyporales) based on a five-marker dataset

Abstract

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.

Full text

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INTRODUCTION
The genus Tramete s Fr. (Polyporales, Basidiomycota) is
characterized by the combination of pileate basidiocarps, po-
roid hymenophore, trimitic hyphal system, thin-walled smooth
basidiospores not react ing in presence of Melzer’s reage nt and
production of a white-rot type of wood decay (Gilbertson
& Ryva r den , 1987 ). It is a cosm opolit an genus prese nt in vi r t u -
ally any type of forest ecosystem in temperate, boreal, and trop-
ical areas (Gilbertson & Ryvarden, 1987). Trametes versicolor
(“turkey tails”) is among the most common and widespread
species of mushroom-forming fungi (Fig. 1) and its whole ge-
nome is currently being sequenced (http://www.jgi.doe.gov/).
To give an idea of how widespread and ecologically important
Trametes species are the USDA databases (http://nt.ars-grin
.gov/fungaldatabases/fungushost/FungusHost.cfm) currently
register 295 plant species in which Trametes versicolor has
been recorded, including angiosperms (e.g., Acer, Betula, Eu-
calyptus, Malus, Populus, Quercus) and conifers (e.g., Abies,
Cupressus, Larix, Pinus). While the concept of Trametes just
outlined is widely accepted (e.g., Ryvarden, 1991; Ryvarden
& Gilbertson, 1994; Bernicchia, 2005) it is not the only one that
has been proposed. Corner (1989) had a much wider generic
concept that synonymized under Trametes 15 other genera of
Polyporales with trimitic hyphal systems (Table 1), including
taxa with stipitate basidiocarps (e.g., Microporus) and genera
causing a brown-rot type of decay (Daedalea, Fomitopsis).
Though he used the name Trametes for this genus the oldest
generic name for Tr a metes sensu Corner would be Daeda-
lea. Ryvarden (1991) defined the Trametes-group of the fam-
ily Polyporaceae Corda by the combination of trimitic hyphal
syste m and whit e -rot ty pe of dec ay an d ther e he in clude d all th e
genera synonymized by Corner (1989) with the exception of the
brown-rot Daedalea and Fomitopsis and the sti pitate Lignosus.
Ryvarden (1991) also included some genera not considered by
Corner (1989) like Cryptoporus, Elmerina and Fomitella in the
Trametes-group of the Polyporaceae (Table 1).
The conflicting concepts about the generic limits of Tram -
etes and the lack of diagnostic morphological features in the
group of trimitic Polyporales have resulted in a large number of
nomenclatural combinations in the genus. The MycoBank da-
tabase (Crous & al., 2004, http://www.MycoBank.org) registers
Phylogenetic classification of Trame tes (Basidiomycota, Polyporales)
based on a five-marker dataset
Alfredo Justo & David S. Hibbett
Clark University, Biology Department, 950 Main St., Worcester, Massachusetts 01610, U.S.A.
Author for correspondence: Alfredo Justo, ajusto@clarku.edu
Abstract:
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 t rametoid 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 par t 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
nomenclatu ral 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 mo-
lecular 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 cu r r ent ly ty pif ied), Coriolus, Cubamyces, Cyclomycetella, Lenzites,
Poronidulus, Pseudotrametes and Pycnoporus are considered synonyms of Trametes.
Keywords
generic concepts; large-scale phylogeny; multi-marker analysis; nomenclature; Polyporaceae; Polyporales;
taxonomy; Trametes
Supplementary Material
Figures S1–S5 are available in the free Electronic Supplement to the online version (Supplementary
Data section) of this article (http://ingentaconnect.com/content/iapt/tax).
SyStematicS and Phylogeny

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Phylogenetic classication of Trametes
Fig. 1.
Morphological overview of the t rametoid clade.
A,
Trametes versicolor;
B,
hymenophore of Trametes suaveolens;
C,
Lenzites betulinus;
D,
sword-like hyphae of L . betulinus;
E,
hymenophore of L. betulinus;
F,
Trametes elegans;
G,
hymenophore of T. el ega n s;
H,
Pycnoporus cin-
nabarinus;
I,
hymenophore of P. cinnabarinus;
J,
Coriolopsis polyzona;
K,
hymenophore of C. polyzona.

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Tab le .
Overview of the genera of Trametes sensu Corner (1989) and the Trametes-group sensu Ryvarden (1991).
Genus (year of publication)
Trametes
sensu Corner
(1989)
Trametes group
sensu Ryvarden
(1991) Phylogenetic placement
Cerrena Gray (1821) Yes Yes Polyporales /phlebioid clade (Ko & Jung, 1999a;
Lee & Lim, 2010; present study)
Coriolopsis Murrill (1905) Yes Yes Polyporales /core polyporoid clade/ Type species (C. poly-
zona) in the /trametoid clade. At least two other indepen-
dent lineages in the /polyporus clade (present study)
Cryptoporus (Peck) Shear (1902) No Yes Polyporales /core polyporoid clade /polyporus clade
(Ko & Jung, 1999b; Binder & al., 2005; present study)
Daedalea Pers. (1801) Yes No Polyporales /antrodia clade (Ko & Jung 1999b; Binder
& al., 2005; present study)
Daedaleopsis J. Schröt. (1888) Yes Yes Polyporales /core polyporoid clade /polyporus clade
(Ko & Jung, 1999b; Binder & al., 2005; present study)
Datronia Donk (1966) Yes Ye s Polyporales /core polyporoid clade /polyporus clade
(Ko & Jung, 1999b; Hibbett & Donoghue, 2001; Binder
& al., 2005; present study)
Earliella Murrill (1905) Yes Yes Polyporales /core polyporoid clade /polyporus clade
(present study)
Elmerina Bres. (1912) No Ye s Tremellales (Larsson & al., 2004)
Fomitella Murrill (1905) No Yes Unknown; no molecular data are available for the type
species (Fomitella supina)
Fomitopsis P. Karst. (1881) Yes No Polyporales /antrodia clade (Ko & Jung, 1999b; Hibbett
& Donoghue, 2001; Binder & al., 2005; present study)
Hexagonia Fr. (1838) Yes Yes Polyporales /core polyporoid clade /polyporus clade
(Ko & Jung 1999b; present study)
Lenzites Fr. (1836) Yes Yes Polyporales /core polyporoid clade /trametoid clade
(Ko & Jung, 1999b; Binder & al., 2005; Tomsovský & al.,
2006; present study)
Lignosus Lloyd ex Torrend (1920) Yes No Polyporales /core polyporoid clade /polyporus clade
(Sotome & al., 2008; present study)
Megasporoporia Ryvarden & J.E. Wright (1982) Yes Yes Polyporales /core polyporoid clade /polyporus clade
(Ko & Jung, 1999b; present study)
Microporus P. Beauv. (1805) Yes Yes Polyporales /core polyporoid clade /polyporus clade
(Sotome & al., 2008; present study)
Mollicarpus Ginns (1984) Yes Yes Unknown; no molecular data are available for the type
species (Mollicarpus cognatus)
Pycnoporus P. Karst. (1881) Yes Yes Polyporales /core polyporoid clade /trametoid clade
(Tomsovský & al., 2006; present study)
Trichaptum Murill (1904) Ye s Ye s Hymenochaetales (Hibbett & Donoghue, 1995; Binder
& al., 2005; Larsson & al., 2006)

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750 specific and infraspecific names in Tramete s and Index
Fungorum (http://www.indexfungorum.org) registers 818, but
the actual number of species that belong in Tramete s sensu
Ryvarden (1991) is much lower, with 18 species reported from
North America, 9 from Europe, and ca. 20 species in the Neo-
tropics (Gilber t son & Ryvar den , 1987; Ryva rde n & Gilb ertso n,
1994; Quante n, 1997; Ryvarden, 200 0). Kirk & al. (2008) esti-
mate the global diversity of the genus is ca. 50 species.
The most inclusive phylogenetic analysis of the order
Polyporales (Binder & al., 2005) used four ribosomal DNA
(rDNA) markers (nuclear ribosomal large subunit [nLSU], mi-
tochondrial large subunit [mtLSU], nuclear ribosomal small
subunit [nSSU], mitochondrial small subunit [mtSSU]) in ap-
prox. 124 species of Polyporales. According to their results,
the order Polyporales was divided into four informally named
clades, viz. antrodia clade, core polyporoid clade, phlebioid
clade, and residual polyporoid clade (throughout the paper clade
names are written with no capitals and not italicized to avoid
confusion with formal taxonomic names). The type species of
Tra m e tes (T. suaveolens) was placed in the core polyporoid
clade together with many of the genera traditionally classified
in the families Polyporaceae and Ganodermataceae (Donk)
Donk (Ryvarden, 1991).
Recent mole cula r studi es have helped to cla r i f y the highe r-
level and generic relationships of polyporoid fungi, and some
of the genera considered as part of the Trametes-group have
been placed outside Polyporales: Trichapt u m in Hymeno chae-
tales (Hibbett & Donoghue, 1995; Binder & al., 2005; Larsson
& al., 2006), and Elmerina in Tre mel lales (La r sso n & al., 20 04).
Other genera in the Trametes-group or Trametes sensu Corner
(1989) belong in Polyporales but are not closely related to
Trametes: Cerrena belongs in the phlebioid clade (Ko & Ju ng,
1999a; Lee & Lim, 2010) and the brown-rot genera Daedalea
and Fomitopsis are placed in the antrodia clade (Ko & Jung,
1999b; Hibbett & Donoghue, 2001; Binder & al., 2005). Based
on the studies just mentioned, the articles by Tomšovský & al.
(2006) and Sotome & al. (2008), and the results presented here
the remaining genera of the Trametes-group belong in the core
polyporoid clade (Table 1; Fig. 1).
The most inclusive phylogenetic studies with a focus on
Trametes are those of Tomšovský & al. (2006), who studied
ITS (internal transcribed spacers 1 and 2 including 5.8S nu-
clear rDNA) and nLSU sequences in 11 isolates representing
seven European species, and Ko & Jung (1999b), who studied
mtSSU sequences in seven isolates of Trametes. Collectively,
these studies support the placement of most Trametes species
in the core polyporoid clade, but also suggested that Trametes
is paraphyletic or polyphyletic, with Lenzites and Pycnoporus
species intermixed with Trametes species.
Molecular data also showed that some species classified
in Trametes by Corner (1989), Gilbertson & Ryvarden (1987),
and Ryvarden & Gilbertson (1994) do not belong in the ge-
nus: (i) Trametes trogii is more closely related to Coriolop-
sis gallica than to the other species of Trametes (Ko & Jung,
1999b; Tomšovský & al., 2006); (ii) Trametes consors belongs
in the genus Cerrena (Ko & Jung, 1999a); (iii) Tomšovský
(2008) showed that Trametes cervina is probably related to
Ceriporiopsis and has been transferred to the new genus Tr a -
metopsis (Tomšovský, 2008).
All the phylogenetic studies mentioned so far, with the
exception of Sotome & al. (2008) and Lee & Lim (2010), rely
exclusively on rDNA data. Protein-coding genes such as the
RNA polymerase II largest subunit (RPB1), RNA polymerase
II second-largest subunit (RPB2), and translation elongation
factor 1-alpha (TEF1) have proven to be useful for phylogenetic
reconstr uction at different taxonomic levels in Agaricomycetes
(e.g., Matheny & al., 2002, 2007; Frøslev & al., 2005) but data
from these genes are lacking for many species of Trametes
and putatively related taxa. The purpose of the present study
is to perform a phylogenetic study focused on Trametes and
related genera using data from ribosomal markers (nLSU, ITS)
and protein-coding genes (RPB1, RPB2, TEF1), examine and
discuss the taxonomic and nomenclatural implications of the
results and finally propose a formal taxonomic arrangement
for the trametoid polypores at the generic level.
MATERIALS AND METHODS
Fungal isolates and DNA extraction. —
Specimens and
culture collections were retrieved from the Forest Products
Laboratory (USDA Forest Service, Madison, Wisconsin,
U.S.A.) supplemented with new collections made in the north-
easter n and southeaster n United States, with a total of 117 iso-
lates (Appendix). Sampling focused on species of Tr a metes
sensu Gilbertson & Ryvarden (1987), with special attention to
type species of genera currently believed to be synonymous
with Trametes, and genera considered as part of the Trametes-
group by Ryvarden (1991).
Cu lture s wer e grow n for 2–3 wee ks at 25°C–30° C on soli d
media (MEA: 20 g malt extract, 0.5 g yeast extract, 20 g agar in
1 l of water). When the plate was totally covered with new my-
celium the tissue was scrapped with sterile scalpels and trans-
ferred to a 1.5 ml microtube and ground with a sterile plastic
pestle. In the case of specimens a small portion of the fruiting
body was ground with liquid nitrogen. Cell lysis proceeded 1 h
at 65°C with the addition of 600 ml extraction buffer (50 mM
EDTA, 50 mM Tris-HCl, 3% SDS, pH 8). Cell debris, poly-
saccharides, and proteins were separated from aqueous DNA
portions through two purification steps with equal volumes
of phenol : chloroform (1 : 1) and chloroform : isoamylalcohol
(24 : 1). Total DNA was precipitated with the addition of 3 M
sodium acetate (0.1 Vol.%) and isopropanol (0.54 Vol.%) and
incubation for 30 to 60 min at −20°C. The DNA pellets were
washed in 1 ml 70% EtOH, dried at 65°C for 5 to 15 minutes
and resu spended in 100 mL ste rile H
2
O. Dilutions of the origi-
nal DNA extraction, usually ranging from 1 : 10 to 1 : 500, were
used in the PCR experiments.
PCR amplification and sequencing. —
The ITS region
(approx. 600–700 bp) was amplified for all 117 samples studied
here using the primer pair ITS-1F/ITS4 (White & al., 1990;
Gardes & Bruns, 1993). Sequence data of nLSU, RPB1, RPB2,
and TEF1 were obtained for a subset of 28 taxa. We succes-
fully amplified these four genes for all selected taxa, except for

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the TEF1 sequence of Coriolopsis gallica. An RPB1 sequence
was also generated for the AFTOL (http://aftol.org/) isolate of
Pycnoporus cinnabarinus, for which data of all other mark-
ers studied here have been already generated (Matheny & al.,
2007). For nLSU (approx. 1300 bp) the primer pair LR0R/
LR7 (Vilgalys Lab, http://www.biology.duke.edu/fungi/myco
lab/primers) was used. For rDNA markers the following PCR
protocol was used: (1) initial denaturation at 95°C for 2 min, (2)
denaturation at 94°C for 45 s, (3) annealing at 50°C for 1 min
10 s, (4) extension at 72°C for 2 min, (5) repeat for 34 cycles
starting at step 2, (6) leave at 72°C for 10 min (Binder & al.,
2010). Sequencing primers for ITS and nLSU were the same
used for PCR and in the case of nLSU with two additional
internal primers: LR3R and LR5 (Vilgalys Lab).
RPB1, RPB2, and TEF1 we re ampli f ied with the followi ng
specifications. The area between conserved domains A and C
of RPB1 (approx. 1400 bp) was amplified using the primer pair
RPB1-Af and RPB1-Cr (Stiller & Hall, 1997; Matheny & al.,
2002). In some cases the primer RPB1-2.2f (Binder & al., 2010)
was used as an alternative to RPB1-Af, giving a slightly shorter
product (approx. 1000 bp). Additional sequencing primers
were: RPB1-2f, RPB1-2.1f, RPB1-2.2f, and RPB1-2.1r (Frøslev
& al., 2005). The area between conserved domains 5 and 11 of
RPB2 (approx. 2100 bp) was amplified in two separate reac-
tions using the primer pairs RPB2-f5F/RPB2-7.1R (Liu & al.,
1999; Matheny, 2005) and RPB2-6.9F/RPB2-b11R1 (Matheny
& al., 2007). Additional sequencing primers were RPB2-b6F/
RPBb6R2 (Matheny, 2005; Matheny & al., 2007) for the first
PCR and RPB2-f7cF/RPB2-b8.2R (Liu & al., 1999; Matheny
& al., 2007) for the second PCR. Approximately 900–1200 bp
of TEF1 was amplified using the primer pair EF1-983F/EF1-
2212R (Rehner & Buckley, 2005). Additional sequencing prim-
ers were EF1-1577F and EF1-1567R (Rehner & Buckley, 2005).
For all protein-coding genes the following “touchdown”
PCR protocol was used: (1) initial denaturation at 94°C for
2 min, (2) denaturation at 94°C for 40 s, (3) annealing at 60°C
for 40s (minus 1 C per cycle), (4) extension at 72°C for 2 min,
(5) repe at for 9 cycles st ar ting at ste p 2, (6) de natura tio n at 94°C
for 45 s, (7) annealing at 53°C for 1 min 30 s, (8) extension at
72°C for 2 min, (9) repeat for 36 cycles starting at step 6, (10)
leave at 72°C for 10 min.
The amplification products for all markers were sequenced
using BigDye 3.1 terminator sequencing chemistry (Applied
Biosystems, Foster City, California, U.S.A.) and run on an
Applied Biosystems 3130 Genetic Analyzer. Raw data were
processed using Sequencher v.4.7 (GeneCodes, Ann Arbor,
Michigan, U.S.A.). Expected fragment lengths and degree of
primer overlap for each one of the protein-genes, as well as
possible alternatives to the primers used here, is posted at http://
wordpress.clarku.edu/polypeet/datasets/primer-information/.
Sequence alignment and phylogenetic analyses. —
In
addition to the sequences generated here an additional 111 se-
quences were retrieved from GenBank (Benson & al., 2011)
and come mainly from the molecular studies mentioned in the
introduction. Accession numbers of ITS sequences used in the
analysis of the trametoid clade are given in Fig. 3 and all other
accesion numbers are listed in the Appendix. The sequences
were aligned using MAFFT v.6 (Katoh & Toh, 2008; http://
mafft.cbrc.jp/alignment/server/). The strategy G-INS-i was se-
lected for all genes except for the alignment of ITS sequences
across the Polyporales where the strategy Q-INS-i was used.
The alignments were manually corrected using MacClade
v.4.08 (Maddison & Maddison, 2002; http://macclade.org/).
For the combined datasets each marker was aligned separately
an d then concate nate d in Mac Cla de. Six dif fer ent datase ts were
assembled for the phylogenetic analyses (Table 2).
Three different phylogenetic analyses were performed in
all the datasets: (i) Maximum likelihood analyses (ML) were
run in the RAxML servers, v.7.2.8 (http://phylobench.vitaleit
.ch/r axmlebb /in dex.p hp; Sta mata k is & al., 20 08), under a GTR
model with one hundred rapid bootstrap replicates. (ii) Equally
weighted parsimony analyses were performed using PAUP*
Table 2.
Description and comparison of the datasets used for the phylogenetic analysis. The Trametes-group is considered in the sense of
Ryvarden (1991).
Dataset Ingroup sequences Outgroup
Total char-
acters (gaps
included)
Parsimony-
informative
characters
Consistency
index/reten-
tion index
Well-supported
nodes
nLSU+ITS 41 Trametes-group taxa
and 24 other Polyporales
Boletopsis leucomelaena and
Hydnellum geogenium (Thelephorales) 1825 345 (19%) 0.38/0.62 28 out of 52 (54%)
ITS 155 trametoid clade
sequences
Dentocorticium sulphurellum and
Lopharia cinerascens (Polyporales) 565 157 (28%) 0.51/0.91 22 out of 71 (31%)
RPB1 28 Trametes-group taxa
and 8 other Polyporales
Camarophyllus basidiosus and
Cantharelulla umbonata (Agaricales) 1178 561 (48%) 0.32/0.43 21 out of 33 (64%)
RPB2 28 Trametes-group taxa
and 19 other Polyporales
Boletopsis leucomelaena and
Hydnellum geogenium (Thelephorales) 2096 814 (39%) 0.25/0.36 24 out of 46 (52%)
TEF1 25 Trametes-group taxa
and 11 other Polyporales
Boletopsis leucomelaena and
Hydnellum geogenium (Thelephorales) 1012 321 (31%) 0.35/0.37 10 out of 25 (40%)
nLSU+ITS+RPB1
+RPB2+TEF1
28 Trametes-group taxa
and 21 other Polyporales
Boletopsis leucomelaena and
Hydnellum geogenium (Thelephorales) 6022 1981 (33%) 0.30/0.40 41 out of 47 (87%)

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v.4.0.b10 (Swofford, 2002). One thousand heuristic search
replicates were performed with starting trees generated by
stepwise addition with random addition sequences followed
by tree bisection reconnection branch swapping. Up to two
trees were kept in each replicate. Parsimony bootstrap analy-
sis was performed with 1000 replicates, each with 10 random
taxon addition sequences and branch swapping set to subtree
pruning and regrafting. (iii) Bayesian analyses (BY) were run
using MrBayes v.3.1 (Ronquist & Huelsenbeck, 2003) at the
Cipres Science Gateway (Miller & al., 2010; http://www.phylo
.org/) for 10 million generations, under a GTR model, with four
chains, and trees sampled every 100 generations. The initial
burn-in phase was set to 2.5 million generations and after ex-
amining the graphic representation of the likelihood scores of
the sampled trees that was confirmed to be an adequate value
for all datasets. A fifty percent majority-rule consensus tree
was computed using the remaining trees. A node is considered
to be well supported if it is supported in at least two of the three
analyses by a bootstrap (BS) value equal or greater than 75%
and/or a posterior probability (PP) equal or greater than 0.95.
For RPB1, RPB2, and TEF1 two alternative datasets were
analyzed using ML, one excluding 3rd codon positions in cod-
ing regions, which are prone to saturation, and another using
amino acid instead of nucleotide data. For all three genes the
phylogenetic resolution of the resulting trees and their support
values decreased with the exclusion of 3rd codon positions or
the use of amino acid data (data not shown), therefore nucleotide
data including 3rd codon positions were used for these genes.
A search for potential conflicts between the rDNA
(nLSU + ITS) dataset and each of the protein-coding genes
was performed by comparing the resulting trees for each da-
taset and looking for strongly supported positive conflict. No
conflicts were detected between the datasets analyzed in the
present study.
RESULTS
New sequences and alignments. —
A total of 230 new
sequences were generated for this study: 117 ITS, 28 nLSU, 29
RPB1, 29 RPB2, and 27 TEF1. A comparative overview of the
different datasets used for the phylogenetic analyses is given
in Table 2. The alignments have been deposited in TreeBASE
(http://purl.org/phylo/treebase/phylows/study/TB2:S11678).
nLSU + ITS dataset of Polyporales (Fig. 2). —
Three major
lineages, the core polyporoid, phlebioid and antrodia clades,
appear as well supported (Fig. 2). The genus Grifola G r ay,
which in some analyses (Binder & al., 2005; García-Sandoval
& al., 2011) appears as a member of the antrodia clade, appears
in our analyses as sister to the core polyporoid clade. The latter
can be divided in three well-supported subclades, here named
as trametoid clade, polyporus clade, and dentocorticium clade.
The trametoid clade includes all sampled species of Tra m -
etes as def i ned by Gilbertson & Ryvarden (1987) and Ryvard en
& Gilbertson (1994) with the exception of Trametes cervina
(= Trametopsis cervina), placed in the phlebioid clade and
Trametes trogii (= Coriolopsis trogii) placed in the polypor us
clade. It also includes Lenzites betulinus, Coriolopsis polyzona,
and the genus Pycnoporus. Some groups in this clade are well
supported: (i) The type of the generic name (T. suaveolens)
is grouped with the T. versicolor group; (ii) L. betulinus is
grouped with T. gibbosa, T. membranacea, T. pavonia, and
T. pocas; (iii) the three Pycnoporus species also are grouped
together. However, relationships between these groups and the
placement of other species of Trametes are unresolved in the
analyses. Our results from the nLSU + ITS dataset are in general
agreement with the data presented by Ko & Jung (1999b) and
Tomšovský & al. (2006).
The polyporus clade includes the type genus of the fam-
ily (Polypor us), which in its current taxonomic concept (e.g.,
Bernichia, 2005) is clearly polyphyletic. Six genera of the
Tra m e tes-group (Ryvarden, 1991) are placed in this clade:
Cryptoporus, Daedaleopsis, Datronia, Earliella, Megaspo-
roporia, and Microporus. Coriolopsis species appear in two
separate groups, and also separate from the type of the genus
(C. polyzona) that is placed in the trametoid clade. In general
the internal relationships of this clade receive low support.
The dentocorticium clade includes Dentocorticum sul-
phurellum and Lopharia cinerascens. It is placed , with no sig-
nificant support, as sister to the trametoid clade.
The remaining member of the Trametes-group sampled
here, the genus Cerrena, is placed in the phlebioid clade.
ITS dataset of the t rametoid clade (Fig. 3). —
All members
of the trametoid clade present in the nLSU+ITS dataset, except
T. pocas for which no ITS is available, are represented here,
together with two additional taxa that were not included in the
nLSU+ITS dataset as no nLSU data is available: Trametes sp.
(collection AJ354) and Lenzites warnieri.
Eighteen major lineages, representing morphological
species or monophyletic species-complexes (e.g., T. versicolor
group), are recovered in the analyses. The sequences of T. ju-
nipericola and Trametes sp. appear in an unresolved position
together with T. conchifer and the T. versicolor group. With
the exception of Pycnoporus sanguineus (including P. coccin-
eus) and T. hirsuta each one of these 18 major lineages is well
supported. However, only the sister-taxa relationship between
L. betulinus an d T. gibbosa and th e groupi ng of the T. versicolor
group and related species receive significant statistical support.
RPB1, RPB2, and TEF1 datasets of the Polyporales (Figs.
S1–S3). —
With the exception of the 5′ end of intron 2 of RPB1
(approx. 470 bp) all intron regions for the three protein-genes
had to be excluded almost entirely because of ambiguous align-
ment. The original alignments including all introns are avail-
able at http://wordpress.clarku.edu/polypeet/datasets/.
The general topology of the RPB1 (Fig. S1) and RPB2 (Fig.
S2) trees is similar to the nLSU + ITS trees (Fig. 2). Sup port val-
ues and the overall resolution are generally better in the RPB1
trees than in the nLSU + ITS or the RPB2 trees (Table 2). The
dentocorticium clade appears as sister to the clade including the
polyporus and trametoid clades, with good statistical support,
in the RPB1 trees. It appears as sister to the polyporus clade,
with no statistical support, in the RPB2 tree. The TEF1 trees
(Fig. S3) are very poorly resolved. Because of this lack of reso-
lution a four-marker dataset (nLSU + ITS + RPB1 + RPB2) was

1573
Justo & Hibbett •
Phylogenetic classication of Trametes
TAX ON
60 (6) • December 2011: 1567–1583
Fig. .
Fift y percent majority-rule consensus tree from the BY analysis of the nLSU+ ITS dataset of the Polyporales. BS values from the ML
and MP analyses ≥ 75 (ML/MP) and PP ≥ 0.95 from the BY analysis for nodes supported in at least two of the analyses are given on or below
the branches.
0.04
Trametes versicolor
Trametes membranacea
Ganoderma lucidum
Trametes pavonia
Phlebia radiata
Lentinus squarrosulus
Trametes pubescens
Pycnoporus puniceus
Trametes gibbosa
Polyporus umbellatus
Cerrena consors
Pseudofavolus cucullatus
Trametes ectypa
Trametes maxima
Megasporoporia setulosa
Pycnoporus cinnabarinus
Hexagonia hirta
Polyporus arcularius
Microporus xanthopus
Earliella scabrosa
Amauroderma rugosum
Coriolopsis aspera
Polyporus udus
Lopharia cinerascens
Hydnellum geogenium
Trametes aff. maxima
Polyporus squamosus
Trametes ljubarskii
Microporus vernicipes
Trametes ochracea
Lentinus tigrinus
Cryptoporus volvatus
Trametes hirsuta
Fomitopsis pinicola
Lenzites betulinus
Boletopsis leucomelaena
Coriolopsis trogii
Trametes cubensis
Trametes elegans
Trametes conchifer
Trametes junipericola
Irpex lacteus
Coriolopsis cf. caperata
Ganoderma tsugae
Datronia mollis
Dentocorticium sulphurellum
Grifola frondosa
Polyporus tricholoma
Pycnoporus sanguineus
Cerrena aurantipora
Coriolopsis polyzona
Phanerochaete chrysosporium
Datronia scutellata
Coriolopsis sanguinaria
Cerrena unicolor
Trametes suaveolens
Coriolopsis gallica
Polyporus brumalis
Daedaleopsis confragosa
Trametopsis cervina
Coriolopsis cf. byrsina
Trametes villosa
Climacodon septentrionalis
Daedalea quercina
Trametes pocas
Grifola sordulenta
Lignosus rhinocerus
1
100/100
87/-
1
0.96
85/93
1
98/79 0.95
94/-
92/-
1
1
81/91
1
99/92
81/88
1
1
92/82
1
1
1
a
a = 86/-
b = 88/78
c = 80/-
b
c
97/87
1
Polyporales
phlebioid
clade
core polyporoid clade
100/100
1
100/96
1
1
98/83
1
100/99
1
100/99
89/77
1
97/94
1
97/97
1
81/- 1
100/100
0.96
86/84 1
100/100
94/99
1
94/93/0.99
antrodia clade
Grifola
dentocorticium clade
trametoid
clade
polyporus clade
75/-
0.98
100/100
1
1
= Type species of Trametes
= Type species of a genus of
Trametes sensu Corner (1989) and/or
the Trametes group sensu Ryvarden (1991)
= Other species in those genera
0.04
Trametes versicolor
Trametes membranacea
Ganoderma lucidum
Trametes pavonia
Phlebia radiata
Lentinus squarrosulus
Trametes pubescens
Pycnoporus puniceus
Trametes gibbosa
Polyporus umbellatus
Cerrena consors
Pseudofavolus cucullatus
Trametes ectypa
Trametes maxima
Megasporoporia setulosa
Pycnoporus cinnabarinus
Hexagonia hirta
Polyporus arcularius
Microporus xanthopus
Earliella scabrosa
Amauroderma rugosum
Coriolopsis aspera
Polyporus udus
Lopharia cinerascens
Hydnellum geogenium
Trametes aff. maxima
Polyporus squamosus
Trametes ljubarskii
Microporus vernicipes
Trametes ochracea
Lentinus tigrinus
Cryptoporus volvatus
Trametes hirsuta
Fomitopsis pinicola
Lenzites betulinus
Boletopsis leucomelaena
Coriolopsis trogii
Trametes cubensis
Trametes elegans
Trametes conchifer
Trametes junipericola
Irpex lacteus
Coriolopsis cf. caperata
Ganoderma tsugae
Datronia mollis
Dentocorticium sulphurellum
Grifola frondosa
Polyporus tricholoma
Pycnoporus sanguineus
Cerrena aurantipora
Coriolopsis polyzona
Phanerochaete chrysosporium
Datronia scutellata
Coriolopsis sanguinaria
Cerrena unicolor
Trametes suaveolens
Coriolopsis gallica
Polyporus brumalis
Daedaleopsis confragosa
Trametopsis cervina
Coriolopsis cf. byrsina
Trametes villosa
Climacodon septentrionalis
Daedalea quercina
Trametes pocas
Grifola sordulenta
Lignosus rhinocerus
1
100/100
87/-
1
0.96
85/93
1
98/79 0.95
94/-
92/-
1
1
81/91
1
99/92
81/88
1
1
92/82
1
1
1
a
a = 86/-
b = 88/78
c = 80/-
b
c
97/87
1
Polyporales
phlebioid
clade
core polyporoid clade
100/100
1
100/96
1
1
98/83
1
100/99
1
100/99
89/77
1
97/94
1
97/97
1
81/- 1
100/100
0.96
86/84 1
100/100
94/99
1
94/93/0.99
antrodia clade
Grifola
dentocorticium clade
trametoid
clade
polyporus clade
75/-
0.98
100/100
1
1
= Type species of Trametes
= Type species of a genus of
Trametes sensu Corner (1989) and/or
the Trametes group sensu Ryvarden (1991)
= Other species in those genera

1574
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60 (6) • December 2011: 1567–1583Justo & Hibbett •
Phylogenetic classication of Trametes
analyzed and the resulting trees (results not shown) compared
to the same dataset with TEF1 data. Inclusion or exclusion of
TEF1 does not significantly change the topology or support
values in the resulting trees, therefore TEF1 sequences were
used in the combined five-marker dataset though it appears that
this gene contributes very little to resolving the higher-level
phylogeny of Polyporales.
Combined (nLSU + ITS + RPB1 + RPB2 + TEF1) dataset of th e
Polyporales (Fig. 4). —
Fomitopsis (antrodia clade) appears as
sister to the remaining Polyporales, followed by the split be-
tween the phlebioid clade and the remaining taxa. Grifola ap-
pears as sister to the core polyporoid clade, which again can be
subdivided in three subclades: trametoid clade and polyporus
clade as sister taxa and the dentocorticium clade as sister to
the clade containing both.
The internal topology of the trametoid clade is much better
resolved than in the nLSU + ITS or any of the single gene data-
sets with 87% of the nodes receiving strong support (Table 2).
Five major lineages are recovered. To facilitate the discussion
each lineage is here informally named using the oldest generic
name available for that group: (i) The pycnoporus clade includes
Pycnoporus and T. cubensis, which are placed as sister to all
other members of the clade. (ii) The artolenzites clade includes
Trametes elegans a nd the T. maxima group. (iii) The co r iolo psi s
clade includes only Coriolopsis polyzona. (iv) The len z ite s clade
includes L. betulinus, T. gibbosa, T. pavonia, and T. membrana-
cea. (v) The trametes clade includes T. vil los a, T. h irsuta , T. sua-
veolens, T. conchifer, and the T. versicolor group. Coriolopsis
polyzona is plac ed as sist er to th e len z ite s clade but th i s relat ion -
ship does not get significant statistical support. Except for this
and the relationship between some members of the T. versicolor
group all other sister-taxa relationships are well supported.
The polyporus clade is also better resolved in the com-
bined dataset with three well-supported major lineages: (i) the
ganoderma clade includes Ganoderma, Amauroderma, and
and some Coriolopsis species; (ii) the datronia clade includes
Polyporus p.p. and Datronia; (iii) the lentinus clade includes
Polyporus p.p., Lentinus, Coriolopsis p.p., Daedaleopsis, and
Earliella. The ganoderma clade appears as sister to the clade
containing the lentinus and datronia clades.
Fig. 3 part 1.
Best tree from the ML analysis of the ITS dataset of the trametoid clade. BS values from the ML and MP analyses ≥ 75 (ML/MP)
and PP ≥ 0.95 from the BY analysis for nodes supported in at least two of the analyses are given on or below the branches.
Trametes cubensis
(? = T. lactinea)
0.07 FJ362294 Uncultured root-associated fungus NM
HM756191 Trametes lactinea FL
FJ372686 Pycnoporus puniceus Thailand
FJ234199 Pycnoporus puniceus Cuba
AF363769 Pycnoporus sanguineus Madagascar
FJ234201 Pycnoporus sanguineus Vietnam
AF363761 Pycnoporus coccineus Solomon Islands
DQ411525 Pycnoporus cinnabarinus China
AF363759 Pycnoporus sanguineus India
AF363771 Pycnoporus sanguineus China
Trametes cubensis TJV93 213sp MS
GU731579 Trametes ljubarskii
Dentocorticium sulphurellum FP11801
FJ234204 Pycnoporus sanguineus New Caledonia
GQ982888 Trametes lactinea Thailand
GQ982887 Trametes lactinea Thailand
Trametes cubensis CRM90 Venezuela
FJ362293 Uncultured root-associated fungus NM
AF363766 Pycnoporus cinnabarinus Belgium
HM756192 Trametes lactinea Seychelles
FJ372685 Pycnoporus puniceus Thailand
FJ234206 Pycnoporus cinnabarinus France
Lopharia cinerascens FP105043sp MS
AF363772 Pycnoporus cinnabarinus Finland
AF363758 Pycnoporus sanguineus Sri Lanka
EU520116 Pycnoporus coccineus China
Pycnoporus sanguineus PRSC95 PuertoRico
FJ234198 Pycnoporus puniceus Cuba
FJ372687 Pycnoporus coccineus Thailand
Trametes cubensis AJ177 FL
FJ750266 Pycnoporus coccineus Japan
FJ234205 Pycnoporus cinnabarinus Russia
FJ873395 Pycnoporus coccineus Australia
Pycnoporus sanguineus CR35 Venezuela
Trametes cubensis CR96 Venezuela
AJ537498 Pycnoporus sanguineus Indonesia
AY684174 Trametes ljubarskii France
100/98/1
Outgroup
Trametes ljubarskii
Pycnoporus
puniceus
group
Pycnoporus cinnabarinus
Pycnoporus sanguineus
(? = P. coccineus)
93/100/1
100/100/1
100/100/1
100/100/1
100/100/1

1575
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Phylogenetic classication of Trametes
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60 (6) • December 2011: 1567–1583
DISCUSSION
Taxonomic and nomenclatural overview of the trametoid
clade. —
Background information on the generic names avail-
able for members of the trametoid clade is given here. The
phylogenetic placement of the type species of these genera,
according to the analysis of the five-marker dataset, is high-
lighted in Fig. 4.
Tra metes Fr., Fl. Sca n.: 339. 1836 – Type: T. suaveolens (L . : Fr.)
Fr., Epicr. Syst. Mycol.: 491. 1838 ≡ Boletus suaveolens
L., Sp. Pl.: 1177. 1753 ≡ Polyporus suaveolens (L. : Fr.) Fr.,
Syst. Mycol. 1: 366. 1821.
Diagnostic characters. – Ba sidio carps an n ual to peren n ial,
pileate, sessile, dimidiate to fan-shaped, single or imbricate,
flexible to hard; upper surface hispid to glabrous, often zon-
ate; pore surface white, cream to pale gray; context white to
Trametes aff. maxima FPRI401 Phillipines
Trametes elegans HHB6551 FL
Lenzites betulinus AJ150 MA
Trametes elegans CRM52 Venezuela
AY787683
Trametes hirsuta
Lithuania
GQ280372 Trametes hirsuta Mexico
Trametes elegans FPRI390 Philippines
Trametes hirsuta AJ303 MA
Trametes elegans HHB4626sp FL
Trametes elegans PR1133 PuertoRico
Trametes hirsuta AJ196 MA
Trametes membranacea PRSC82 PuertoRico
Trametes pavonia FP103050sp FL
Trametes maxima OH241sp Venezuela
Trametes maxima OH189sp Venezuela
Trametes hirsuta HHB8591sp AZ
Trametes gibbosa L11664 UK
Trametes villosa FP71974R TN
FJ904854 Coriolopsis polyzona Kenya
GU067734 Lenzites betulinus Finland
Trametes hirsuta L12964sp CostaRica
Coriolopsis polyzona BKW001 Ghana
Trametes elegans FRI437T
Trametes hirsuta FP103959T IL
Trametes elegans FP150762 Belize
Trametes membranacea PRSC69 PuertoRico
Trametes hirsuta RLG5133T NY
Trametes elegans FPRI10 Philippines
EU009976 Polyporales sp. Hawaii
GU062274 Trametes hirsuta Latvia
Lenzites betulinus HHB9942sp FL
AY855916 Lenzites warnieri
Trametes elegans FP105038sp MS
FJ462747 Trametes hirsuta China
AB158315 Trametes maxima Cuba
Trametes hirsuta FP101666sp WI
GQ280373 Trametes hirsuta Mexico
Trametes villosa FP86588sp FL
Trametes pavonia PR2178 PuertoRico
HM756194 Trametes elegans Thailand
Trametes hirsuta DR EST7 Estonia
AY684176 Trametes gibbosa Czech Republic
Trametes membranacea PR3264 PuertoRico
GU731567 Lenzites warnieri
Coriolopsis polyzona OH272sp Venezuela
EU009977 Trametes sp. Hawaii
Coriolopsis polyzona BKW004 Ghana
Trametes aff. maxima FPRI376 Philippines
Trametes membranacea CRM125 Venezuela
Trametes maxima OH172 Venezuela
Coriolopsis polyzona BKW017 Ghana
EU009975 Polyporales Mexico
FJ481048 Trametes gibbosa China
Trametes elegans OH271sp Venezuela
Trametes elegans FP105679 GA
0.07
Trametes elegans
Lenzites warnieri
Coriolopsis polyzona
Trametes maxima group
Trametes pavonia
Trametes membranacea
Trametes gibbosa
Lenzites betulinus
Trametes villosa
Trametes hirsuta
97/96
/1
91/97/1
96/89
1
100/100/1
100
99/1
100/100/1
100/100/1
85/85/1
100/100/1
100/100
/1
100/100
/1
98/96
/1
84/83
/0.98
94/92/0.99 Fig. 3 part 2 .

1576
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60 (6) • December 2011: 1567–1583Justo & Hibbett •
Phylogenetic classication of Trametes
FJ810185 Trametes suaveolens China
Trametes ectypa FP81034R FL
Trametes sp. AJ354 South Korea
Trametes conchifer HHB4940sp NY
Trametes pubescens L15711sp NY
AY686706 Trametes versicolor South Korea
AY684180 Trametes suaveolens Czech Rep.
Trametes conchifer FP86583R VA
AY684173 Trametes pubescens Czech Rep.
Trametes suaveolens HHB14170T AK
Trametes pubescens L15318sp NY
Trametes ectypa FP106037T MS
Trametes versicolor FP133494sp OR
GU067736 Trametes versicolor Finland
Trametes ochracea L6773 CO
Trametes versicolor AJ207 MA
Trametes conchifer FP15064sp NY
Trametes ochracea HHB13429sp AK
FJ903303 Trametes versicolor Latvia
Trametes suaveolens FP102529T WI
Trametes conchifer FP102251sp WI
GU062223 Trametes versicolor Latvia
AM269813 Trametes versicolor VT
FJ591065 Trametes versicolor China
Trametes versicolor AJ190 MA
Trametes versicolor AJ183 FL
Trametes ectypa FP103976sp FL
EU661891 Trametes versicolor China
Trametes pubescens FP101414sp WI
GQ411515 Trametes versicolor New Zealand
FJ810176 Trametes suaveolens China
Trametes versicolor AJ146 FL
Trametes versicolor AJ164 FL
AM981234 Trametes versicolor Slovenia
AY805633 Trametes versicolor Sweden
Trametes pubescens FP101536T WI
AY684177 Trametes ochracea Czech Republic
Trametes versicolor HHB12282sp WI
Trametes pubescens L16093sp NY
AB368486 Trametes versicolor South Korea
Trametes versicolor AJ170 FL
AY684171 Trametes junipericola
FJ591066 Trametes versicolor China
Trametes ochracea MJL2103 CAN Ontario
FJ810188 Trametes suaveolens China
AM269814 Trametes versicolor Switzerland
Trametes versicolor AJ198 MA
Trametes versicolor FP102607sp WI
Trametes pubescens HHB13585sp WI
Trametes conchifer FP106793sp MS
Trametes versicolor FP134940sp ID
Trametes versicolor BAFC285 Argentina
AY684179 Trametes versicolor Czech Republic
Trametes versicolor FP135156sp NY
Trametes versicolor Braz16 Brazil
Trametes ochracea HHB13445sp MI
Trametes pubescens FP101554T WI
Trametes versicolor AJ151 MA
GU067738 Trametes ochracea Finland
Trametes versicolor AJ119 MA
EU771081 Trametes versicolor China
FJ903281 Trametes ochracea Latvia
Trametes versicolor FP1022316sp WI
Trametes conchifer FP103312sp SC
Trametes suaveolens FP102529sp WI
0.07
Trametes versicolor group:
T. versicolor
T. pubescens
T. ochracea
T. ectypa
Trametes conchifer
Trametes junipericola / Trametes sp.
Trametes suaveolens
99/99/1
91/90/1
93/83/1
a
a = 99/94/1
Fig. 3 part 3.

1577
Justo & Hibbett •
Phylogenetic classication of Trametes
TAX ON
60 (6) • December 2011: 1567–1583
Fig. 4.
Fifty percent majority-rule consensus tree from the BY analysis of the n LSU + ITS + RPB1 + RPB2 + TEF1 dataset of the Polyporales. BS
values from the ML and MP analyses ≥ 75 (ML/MP) and PP ≥ 0.95 from the BY analysis for nodes supported in at least two of the analyses are
given on or below the branches.
1
1
1
1
1
1
1
1
1
186/-
/1
1
78/-/0.99
1
1
89
1
1
1
89/-/1
1
1
1
1
1
100/95/1
1
1
1
1
-/75/0.99
81/83/0.99
11
1
1
82/-/0.99
100/100
89/-
100/99
100/94
100/95
92/-
98/-
100/89
100/98
1
100/96
88/-/1
100/100
100/100
80/-
100
100/86
100/100
100/100
100/100
100/100
98/97
99/100
100/100
96/-
100/97/1
92/-/0.99
82/78
100/99
100/100
95/- 100/100
100/100
95/91
Polyporales
phlebioid
clade
Grifola
Fomitopsis
core polyporoid
clade
100/99
trametes clade
lenzites
clade
coriolopsis clade
artolenzites clade
pycnoporus
clade
0.08
Polyporus arcularius
Polyporus brumalis
Lentinus squarrosulus
Lentinus tigrinus
Polyporus tricholoma
Coriolopsis gallica
Coriolopsis trogii
Daedaleopsis confragosa
Earliella scabrosa
Datronia mollis
Polyporus squamosus
Pseudofavolus cucullatus
Datronia scutellata
Polyporus udus
Polyporus umbellatus
Amauroderma rugosum
Coriolopsis cf. byrsina
Ganoderma lucidum
Ganoderma tsugae
Coriolopsis cf. caperata
Trametes ectypa
Trametes versicolor
Trametes ochracea
Trametes pubescens
Trametes conchifer
Trametes suaveolens
Trametes hirsuta
Trametes villosa
Lenzites betulinus
Trametes gibbosa
Trametes pavonia
Trametes membranacea
Coriolopsis polyzona
Trametes aff. maxima
Trametes maxima
Trametes elegans
Pycnoporus cinnabarinus
Pycnoporus sanguineus
Trametes cubensis
Dentocorticium sulphurellum
Lopharia cinerascens
Grifola frondosa
Grifola sordulenta
Climacodon septentrionalis
Phlebia radiata
Phanerochaete chrysosporium
Irpex lacteus
Trametopsis cervina
Fomitopsis pinicola
Hydnellum geogenium
Boletopsis leucomelaena
Trametes
Coriolus
Poronidulus
Lenzites
Pseudotrametes
Cyclomycetella
Coriolopsis
Artolenzites
Pycnoporus
Cubamyces
1
A
B
C
A = dentocorticium clade
B = trametoid clade
C = polyporus clade
lentinus clade
datronia
clade
ganoderma clade
= Type species of Trametes
= Generic type
= Other taxa of the Trametes-group
sensu Ryvarden (1991)
1
1
1
1
1
1
1
1
1
186/-
/1
1
78/-/0.99
1
1
89
1
1
1
89/-/1
1
1
1
1
1
100/95/1
1
1
1
1
-/75/0.99
81/83/0.99
11
1
1
82/-/0.99
100/100
89/-
100/99
100/94
100/95
92/-
98/-
100/89
100/98
1
100/96
88/-/1
100/100
100/100
80/-
100
100/86
100/100
100/100
100/100
100/100
98/97
99/100
100/100
96/-
100/97/1
92/-/0.99
82/78
100/99
100/100
95/- 100/100
100/100
95/91
Polyporales
phlebioid
clade
Grifola
Fomitopsis
core polyporoid
clade
100/99
trametes clade
lenzites
clade
coriolopsis clade
artolenzites clade
pycnoporus
clade
0.08
Polyporus arcularius
Polyporus brumalis
Lentinus squarrosulus
Lentinus tigrinus
Polyporus tricholoma
Coriolopsis gallica
Coriolopsis trogii
Daedaleopsis confragosa
Earliella scabrosa
Datronia mollis
Polyporus squamosus
Pseudofavolus cucullatus
Datronia scutellata
Polyporus udus
Polyporus umbellatus
Amauroderma rugosum
Coriolopsis cf. byrsina
Ganoderma lucidum
Ganoderma tsugae
Coriolopsis cf. caperata
Trametes ectypa
Trametes versicolor
Trametes ochracea
Trametes pubescens
Trametes conchifer
Trametes suaveolens
Trametes hirsuta
Trametes villosa
Lenzites betulinus
Trametes gibbosa
Trametes pavonia
Trametes membranacea
Coriolopsis polyzona
Trametes aff. maxima
Trametes maxima
Trametes elegans
Pycnoporus cinnabarinus
Pycnoporus sanguineus
Trametes cubensis
Dentocorticium sulphurellum
Lopharia cinerascens
Grifola frondosa
Grifola sordulenta
Climacodon septentrionalis
Phlebia radiata
Phanerochaete chrysosporium
Irpex lacteus
Trametopsis cervina
Fomitopsis pinicola
Hydnellum geogenium
Boletopsis leucomelaena
Trametes
Coriolus
Poronidulus
Lenzites
Pseudotrametes
Cyclomycetella
Coriolopsis
Artolenzites
Pycnoporus
Cubamyces
1
A
B
C
A = dentocorticium clade
B = trametoid clade
C = polyporus clade
lentinus clade
datronia
clade
ganoderma clade
= Type species of Trametes
= Generic type
= Other taxa of the Trametes-group
sensu Ryvarden (1991)

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Phylogenetic classication of Trametes
isabelline, homogeneous or duplex, in some species with a dark
line; hyphal system trimitic; generative hyphae hyaline, with
clamps; skeletal hyphae straight, thick-walled to solid, hyaline,
in some species swelling in KOH; binding hyphae tortuous,
solid, hyaline; cystidia absent, in some species pointed (“sword-
like”) hyphal ends may penetrate the hymenium; spores ellip-
soid to allantoid, hyaline, thin-walled, and negative in Melzer’s
re age nt; hete rot hall ic and tetra polar; cause s a white ro t in hard-
wood s, rarely on con ife r wood; co smopolit a n gen us with many
common and widespread species, ca. 50 species worldwide
(Gilbertson & Ryvarden, 1987; Ryvarden & Gilbertson, 1994;
Kirk & al., 2008).
Observations. – Trametes elegans, with a variable hy-
menophore from poroid to lamellate (Fig. 1F–G), was included
in Lenzites by Ryvarden & Johansen (1980) but later in Tr am-
etes by Gilbe r tson & Ry varden (1987), despite the fa ct th at the
lamellate hymenophore is one of the key characters used by
Gilbertson & Ryvarden (1987) to keep Lenzites se para ted from
Trametes. Both names (T. e lega n s and L. elegans) are present
in modern treatments of trametoid fungi: e.g., Quanten (1997)
as L. elegans and Ryvarden (2000) as T. e l egans.
Lenzites Fr., Fl. Scan.: 339. 1836 – Type: L. betulinus (L. : Fr.)
Fr., Epicr. Syst. Mycol.: 405. 1838 ≡ Agaricus betulinus
L., Sp. Pl.: 1176. 1753 ≡ Agaricus betulinus L. : Fr., Syst.
Mycol. 1: 133. 1821.
Diagnostic characters. – Basidiocarps annual, pileate;
upper surface glabrous to hirsute, often in distinct zones; hy-
menophore lamellate to daedaleoid, white to yellowish; con-
text concolorous, tough and moderately thick; hyphal system
trimitic; generative hyphae thin-walled, hyaline, with clamps;
binding hyphae hyaline, strongly branched and tortuous, in
the trama below the hymenium with long, “sword-like” side
branches more or less parallel and pointing towards, and in
parts into, the hymenium, but never above it; skeletal hyphae
hyaline, thick-walled to solid; spores cylindrical, smooth, thin-
walled, hyaline, negative in Melzer’ s reagent; heterothallic and
tetrapolar; on dead hardwoods, more rarely on conifers, causing
a white rot. Cosmopolitan genus, with 3–5 species worldwide.
(Gilbertson & Ryvarden, 1987; Ryvarden & Gilbertson, 1993;
Kirk & al., 2008).
Observations. – Lenzites is separated from Trametes on
the basis of the distinctly lamellate to daedaleoid hymenophore
and the presence of “sword-like” binding hyphae (Fig. 1C–E).
However, a lamellate hymenophore is also present in T. elega ns
and “sword-like” binding hyphae are found in T. cubensis, a
species with poroid hymenophore and otherwise a “typical”
member of Trametes (Gilbertson & Ryvarden, 1987).
Many polyporoid fungi with lamellate/daedaleoid hymeno-
phore have been recombined in Lenzites through the years (e.g.,
Daedalea, Gloeophyllum) resulting in more than 200 nomen-
clatural combinations in the genus but the actual number of
species of Lenzites sensu Ryvarden (1991) is between 3 and 5
(http://www.MycoBank.org/).
Pycnoporus P. Karst. in Rev. Mycol. 3: 18. 1881 – Type: P. cin -
nabarinus (Jacq. : Fr.) P. Karst. in Rev. Mycol. 3: 18. 1881
≡ Boletus cinnabarinus Jacq., Fl. Austriac. 4: 2. 1776 ≡
Polyporus cinnabarinus (Jacq. : Fr.) Fr., Syst. Mycol. 1:
371. 1821.
Diagnostic characters. – Basidiocarps annual, sessile to
effused-reflexed, dimidiate; pileus surface and pore surface
orange-red to cinnabar, color fading on weathering; pores
regular, circular to angular, 3– 4 per mm; context reddish or-
ange, coriaceous; hyphal system trimitic; generative hyphae
with clamps; tramal hyphae with dextrinoid contents; cystidia
absent; basidiospores cylindric, hyaline, smooth, negative in
Melzer’s reagent; heterothallic and tetrapolar; causing a white
rot of dead hardwoods, rarely on conifers. Cosmopolitan ge-
nus, with one species (P. cinnabarinus) in temperate areas and
th r ee (P. coccineus, P. puniceu s, P. sanguine us) in tropical and
subtropical areas (Gilbertson & Ryvarden, 1987; Ryvarden
& Gilbertson, 1994; Kirk & al., 2008).
Observations. – Pycnoporus is separated from Trametes
solely on the basis of the orange-red colors of the basidiocarps
(Fig. 1H–I [for color figure see this article online]).
Coriolopsis Murrill in Bull. Torrey Bot. Club 32: 358. 1905 –
Ty p e: C. occidentalis (Klotzsch) Murrill in Bull. Torrey
Bot. Club 32: 358. 1905 ≡ Polyporus occidentalis Klotzsch
in Linnaea 8: 486. 1833. According to Ryvarden (1972) who
studied the type materials this is a synonym of C. poly-
zona (Pers.) Ryvarden ≡ Polyporus polyzonus Pers. in
Gaudichaud-Beauprét, Voy. Uranie 5: 171. 1827).
Diagnostic characters. – Basidiocarps annual, pileate and
sessile, rarely resupinate; pileus velutinate to hirsute, more
rarely glabrous, zonate to azonate, yellowish to umber brown,
often grayish when hirsute; pore surface concolorous, in some
species with a grayish to blue bloom, pores round to entire,
small to moderately large; context golden to umber brown,
not distinctly delimited toward the pilear cover; hyphal system
trimitic; generative hyphae hyaline, thin-walled, with clamps;
skeletal hyphae thick-walled to solid, cyanophilous, hyaline,
ochrac eous to dee p golde n brown; bindi ng hyp hae th ick-walle d
and concolorous with skeletal hyphae, cystidia absent; spores
hyaline, cylindrical to oblong-ellipsoid, smooth, thin-walled,
negative in Melzer’s reagent, on hardwoods, causing a white
rot. Cosmopolitan genus but most species are restricted to the
tropical zone, ca. 17 species worldwide (Gilbertson & Ryvar-
den, 1986; Ryvarden & Gilbertson, 1993; Kirk & al., 2008).
Observations. – Coriolopsis is separated from Trametes
mainly by the colored hyphae that give the context a brown
color (Fig. 1J–K). The presence of cyanophilic skeletal hyphae
has also been cited as a diagnostic character but it has not been
demonstrated for all the species currently placed in Coriolopsis
(Ryvarden, 1991).
In addition to Trametes, Lenzites, Pycnoporus, and Corio-
lopsis, cur rently accept ed as se para te, there are other si x leg iti -
mate and validly published generic names for taxa included in
the trametoid clade (Fig. 4). None of these generic names, ex-
cept Coriolus, have been widely used by polypore taxonomists
during the 20th century (Ryvarden, 1991), mostly because the
morphological separation from the four main trametoid genera
discussed above is very difficult, if not impossible in some

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Phylogenetic classication of Trametes
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60 (6) • December 2011: 1567–1583
cases. They are listed here because they are relevant for the
nomenclatural and taxonomic discussion of the trametoid clade.
Artolenzites Falck, Hausschwamm-Forschungen 3: 37. 1909 –
Ty p e: Daedalea repanda Pers. in Gaudichaud-Beauprét,
Voy. Uranie 5: 168. 1827 (= Trametes elegans (S p re ng . : Fr.)
Fr., Epicr. Syst. Mycol.: 492. 1838).
Artolenzites elegans (Spreng. : Fr.) Teixeira is the only no-
menclatural combination made in this genus.
Coriolus Quél., Ench i r. Fung.: 175. 1886 – Type: Coriolus ver-
sicolor (L. : Fr.) Quél., Enchir. Fung.: 175. 1886 ≡ Boletus
versicolor L., Sp. Pl.: 1176. 1753 ≡ Polyporus versicolor
(L. : Fr.) Fr., Observ. Mycol. 2: 260. 1818 (sanctioned in
Syst. Mycol. 1: 368. 1821.) ≡ Trametes versicolor (L. : Fr.)
Lloyd, Mycol. Writings 6: 1045. 1920.
Coriolus has been considered by some authors as separate
from Trametes because of the presence of a thin black li ne be-
tween the tomentum and the context, but recent authors (e.g.,
Gilbertson & Ryvarden, 1987) have considered this character
to be very variable and of no taxonomic significance on the ge-
neric level. The genus Cellularia Bul l. (Bulliard, 1789: pl. 414)
is typified by Cellularia cyathiformis Bull., which is cited by
MycoBank (http://www.MycoBank.org) and Index Fungorum
(http://www.indexfungorum.org) as a synonym of Coriolus
versicolor (= T. versicolor). However, if this synonymy is ac-
cepted Coriolus would become an illegitimate name as Cellu-
laria is almost 100 years older. A formal proposal to conserve
Coriolus against Cellularia made by Hawksworth (1984) was
rejected by the Committee of Fungi (Korf, 1988), because the
actual synonymy of Coriolus versicolor and Cellularia cyathi-
formis was con sider ed doubtful . Since this sy nonymy rem ains
as doubtful now as it did at the time of Hawksworth’s proposal
the genus Cellularia will not be considered in the nomencla-
tural discussion.
Cyclomycetella Mur r i ll in Bu ll. Torrey Bot . Club 31: 722 . 1904
– Type: Cyclomycetella pavonia (Hook.) Murrill in Bull.
Torrey Bot. Club 31: 722. 1904 ≡ Boletus pavonius Hook.
in Kunth, Syn. Pl. 1: 10. 1822 ≡ Trametes pavonia (H o ok.)
Ryvarden in Norweg. J. Bot. 19: 237. 1972 [illegitimate
because of the existence of Trametes pavonia (Berk.) Fr.,
a taxonomic synonym of T. e legan s].
Cyclomycetella pavonia is the only nomenclatural combi-
nation made in this genus.
Cubamyces Murrill in Bull. Torrey Bot. Club 32: 480. 1905 –
Ty p e: Cubamyces cubensis (Mont.) Murrill in Bull. Torrey
Bot. Club 32: 480. 1905 ≡ Polyporus cubensis Mont. in
Ann. Sci. Nat., Bot., ser 2, 8: 364. 1837 ≡ Trametes cuben-
sis (Mont.) Sacc., Syll. Fung. 9: 198. 1891.
Cubamyces cubensis is the only nomenclatural combina-
tion made in this genus.
Poronidulus Murrill in Bull. Torrey Bot. Club 31: 425. 1904
– Type: Poroninulus conchifer (Schwein. : Fr.) Murrill in
Bull. Torrey Bot. Club 31: 425. 1904 ≡ Boletus conchifer
Schwein. in Schriften Naturf. Ges. Leipzig 1: 98. 1822 ≡
Polyporus conchifer (Schwein. : Fr.) Fr., Elench. Fung. 1:
96. 1828 ≡ Trametes conchifer (Schwein. : Fr.) Pilát, Atlas
Champ. Eur., Polypor., B 3: 264. 1939.
In its more typical form Poronidulus conchifer is easy to
recog nize beca use of the cup -shaped struct u res in the ba sidio -
carps that produce asexual propagules. However, this character
is not always present and Gil berts on & Ry varden (1987) did not
consider it important enough to maintain T. conchifer in a sep a-
rate genus. The only other taxon in the genus is Poronidulus bi-
valvis Höh n., but the actu al identity of this species is doubt f ul.
Pseudotrametes Bondartsev & Singer ex Singer in Mycologia
36: 68. 1944 – Type: Pseudotrametes gibbosus (Pers. : Fr.)
Bondartsev & Singer ex Singer in Mycologia 36: 68. 1944
≡ Merulius gibbosus Pers. in Ann. Bot. (Usteri) 15: 21.
1795 ≡ Daedalea gibbosa (Pers. : Fr.) Pers., Syn. Meth.
Fung.: 501: 1801 (sanctioned in Syst. Mycol. 1: 338. 1821)
≡ Trametes gibbosa (Pers. : Fr.) Fr., Epicr. Syst. Mycol.:
492. 1838 .
Pseudotrametes was created to accommodate T. gibbosa,
characterized by the radially elongated pores. It is the only
species of the genus.
In view of the results from the phylogenetic analyses and
the nomenclatural and taxonomic history of this group we dis-
cuss below different taxonomic scenarios for Trametes and
related genera.
Phylogenetic classification of the trametoid clade. —
We conclude that all members of the trametoid clade should
be classified under one single generic name (Trametes). The
morphological concept of Trametes of Gilbertson & Ryvarden
(1987) would only have to be slightly expanded to allow spe-
cies with or ang e-red basid ioc a r ps (Pycnoporus) and/or colored
hyphae and context (Pycnoporus, C. polyzona) to be included
in Trametes. Ascription of unsampled and newly described
taxa to this “new” Trametes will be easier than in any of the
alternative taxonomic scenarios considered here and not more
complicated than with the current morphological concept of
Trametes. This solution is straightforward, but it is not totally
free of problems.
The morphological separation of the genus Trametes and
some taxa currently placed in Coriolopsis will still be prob-
lematical. In all phylogenetic analyses Coriolopsis is clearly
polyphyletic (Figs. 2, 4), with two lineages in the polyporus
clade not closely related to each other and away from the type
species (C. polyzona) that will become a member of Trametes.
Morphological characters such as spore length (e.g., above
10 µm in C. aspera, C. byrsina, C. gallica, and C. trogii) or
the tomentose-strigose pileus surface of many species can help
to separate these taxa from Trametes, especially from Tram -
etes polyzona. This difficult morphological separation is not
a consequence of the new taxonomic concept of Trametes but
the result of the lack of diagnostic morphological features in
the group of trimitic polypores (Ryvarden, 1991).
Of all the generic names available for taxa in the trametoid
clade, Lenzites and Tra metes have equal nomenclatural priority

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Phylogenetic classication of Trametes
as they were published on the same year and on the very same
page of the same book (Fries, 1836: 339). For these cases Art.
11.5 of the ICBN. (McNeill & al., 2006) dictates that the first
author treating Lenzites and Trametes as synonyms should be
followed in his choice of name. Pilát (Kavina & Pilát, 1939) was
the first to do so, choosing Trametes over Lenzites, and thus
recombining the type species of Lenzites (L. betulinus) under
Trametes. While some species of Trametes have been alterna-
tively classified under Lenzites (e.g., T. ele gans, T. gibbosa)
the type species (T. suaveolens) has never been recombined in
Lenzites. The name Trametes betulina (L. : Fr.) Pilát should be
used for Lenzites betulinus.
Nomenclatural combinations in Trame t e s already ex-
ist for all species currently placed in Pycnoporus: Trametes
cinnabarina (Jacq. : Fr.) Fr. (= Pycnoporus cinnabarinus),
Trametes punicea Fr. (= Pycnoporus puniceus), and Trametes
sanguinea (L. : Fr.) Lloyd (= Pycnoporus sanguineus). Corner
(1989) recombined Coriolopsis polyzona in Trametes but fai led
to provide a full and direct reference to its basionym therefore
his name is not validly published under Art. 33.4 of the ICBN
(McNeill & al., 2006). The new combination is proposed here:
Trametes polyzona (Pe rs.) Just o, co mb. nov. ≡ Polyporus poly-
zonus Pers. in Gaudichaud-Beaupré, Voy. Ura n i e 5: 171.
MycoBan k 561896
In the case of L. warnieri and Lenzites vespacea (Pe r s .)
Pat. new combinations in Tra m e tes would be necessary but
the separation of these taxa from each other and from Lenz-
ites acuta Berk. is at present doubtf ul based on mor phological
characters (Ryvarden & Johansen,1980) and molecular data
are only available for L. warnieri. Considering that there are
already over 120 nomenclatural and taxonomic synonyms for
L. acuta and L. vespacea listed in MycoBank (http://www
.MycoBank.org) we refrain from proposing any new nomen-
clatural combination for these taxa until molecular data are
available for the three species.
Alternatives to using the name Trametes for all members
of the trametoid clade were also considered (Fig. S4). The most
obvious one would be to recognize the five well-supported lin-
eages recovered in the five-marker dataset analysis (Fig. 4) at
the generic level: Trametes, Lenzites, Coriolopsis, Artolenzites,
and Pycnoporus. If this option is adopted it would be neces-
sary to transfer some species currently placed in Trametes to
Lenzites (T. gibbosa, T. pavonia, T. membranacea) or Pycno-
porus (T. cubensis) and to “resurrect” the name Artolenzites
to accommodate T. eleg a n s and the T. maxima group. Rea sons
not to adopt this option can be readily recognized: (i) These
five genera would be indistinguishable from each other based
on morphology, ecology, and/or geographical distribution. (ii)
Given the homoplasious nature of some important characters
used in the generic distinction of this group (e.g., lamellate vs.
poroid hymenophore, presence of “sword-like” binding hyphae,
nature and distribution of the pigments in the basidiocarps) it
is not possible to determine, based on morphology alone, the
taxonomic position of species not sampled here. (iii) It will not
be possible to ascribe newly described species to any of these
five genera without generating molecular data and even then if
only rDNA is available (e.g., T. pocas, T. ljubarskii, L. warnieri)
it may not be enough for a confident generic classification.
We also considered a third taxonomic scenario: to rec-
ognize the genera Trametes, Coriolopsis (on ly C. polyzona),
Lenzites, and Pycnoporus as defined by Gilbertson & Ryvar-
den (1987), without transferring any species into them, and
adopting new generic names for the taxa that fall outside these
genera. The generic names Pseudotrametes, Cyclomycetella,
Artolenzites, and Cubamyces would accommodate only their
respective type species and two new genera would have to be
created for T. membranacea and the T. maxima group. With
the current sampling a total of ten genera would have to be ac-
cepted for taxa in the trametoid clade (Fig. S4). However the
same problems just outlined for the five-genera option will be
even more serious with this option and very likely the number
of genera in the trametoid clade would increase as new taxa
are sampled for molecular analyses.
Species recognition in Trametes. —
The results of the ITS
(Fig. 3) and five-marker analyses (Fig. 4) offer some insights
about the species-level taxonomy of Trametes.
The separation of the species group around T. versicolor
(T. ectypa, T. ochracea, T. pubescens) is unclear in the ITS
da t ase t. Mor phological separ ation of thes e taxa is also difficult
given the variability of morphological characters used for their
delimitation (Gilbertson & Ryvarden, 1987; Ryvarden & Gil-
bertson, 1994). However there are different lines of evidence
like (i) the results from the five-marker dataset (Fig. 4), (ii)
separate ITS analyses of this complex and the presence of mo-
lecular polymorphisms in the ITS unique to each one of these
taxa (data available at http://wordpress.clarku.edu/polypeet/
datasets/), and (iii) the intercompatibility tests performed by
Tomšovský & Homolka (2004) that suggest that they may actu-
ally be different biological species or at least they are currently
undergoing speciation. The ITS data also suggest the possible
synonymy of Pycnoporus sanguineus and Pycnoporus coc-
cineus, as already proposed by Ryvarden & Johansen (1980)
and of T. cubensis and T. l a ctinea. There is also evidence of
intercontinental distribution of some species of Trametes (e.g.,
T. betulina, T. suaveolens, T. versicolor) as well as their ability
to survive under very different climate conditions, e.g., T. bet-
ulina occurs in boreal (Finland), temperate (Massachusetts,
U.S.A.) and tropical (Hawaii) ecosystems. A more exhaustive
multi-gene analysis of Tramete s is in preparation to address
these questions in detail.
Morphological separation of Trametes species, as recog-
nized in the phylogenetic analyses, relies on characters such as
the consistency of the basidiocarps, nature of the hymenophore
(poroid vs. lamellate, color, morphology of the pores, number
of pores per mm), surface and colors of pileus, presence of red-
orange or dark brown pigments in some parts of the basidio-
carps, spore size and shape and, to a lesser extent, presence
of specialized structures (e.g., “sword-like” binding hyphae,
cystidioles, asexual propagules).
Final overview of the Trametes-group and remaining
questions. —
With the inclusion of the generic type (Corio-
lopsis polyzona) in Trametes the name Coriolopsis becomes

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60 (6) • December 2011: 1567–1583
Benson, D.A., Karsch- Mi zrach i, I., Lipman, D. J., Ostell, J. & Say-
ers, E.W. 2011. GenBank. Nucl. Acids Res. 39: D32–D327.
Bernicchia, A. 20 05. Polyporaceae s.l. Fungi Europaei 10. Alassio:
Ed. Candusso.
Bi nder, M. , Hibb ett , D.S. , Lar s son , K.- H., Lars s on , E. & Lan ge r, E.
a synonym of Tramete s. A change of type species could be
proposed to keep the name Coriolopsis for the C. gallica group.
Alternatively, the name Funalia Pat. could be adopted for the
C. gallica group but no molecular data are available for the
type species of Funalia, viz. Polyporus mons-veneris Jungh.
(= Trametes leonina (Klotzsch) Imazeki, = Funalia leonina
(Klotzsch) Pat.). Morphological characters of F. leonina, es-
pecially the spore length above 10 µm and the strigose pileus
(Ryvarden & Johansen, 1980), suggest a relationship with the
C. gallica group. The generic name Trametella Pinto-Lopes,
typified by Trametes hispida (= C. gallica) is also available.
Molecular data for Funalia leonina are critical for making a
final decision for this group of species.
The other two species of Coriolopsis sa mpled in th is study
(Coriolopsis cf. byrsina, Coriolopsis cf. caperata) are closely
related to Ganoderma and Amauroderma and ac cor ding to the
results of the combined dataset (Fig. 4) they are not grouped
together. In both cases the species-level identification is tenta-
tive as no reliable molecular data are available for comparison.
A preliminary analysis of all ITS sequences of Coriolopsis
available in GenBank and sequences generated here indicates
that the 64 ITS sequences deposited in GenBank as “Corio-
lopsis caperata” represent at least three different, not closely
related, species and the available ITS sequence of “C. byrsin a”
represents Earliella scabrosa (Fig. S5).
The current knowledge about the phylogenetic placement
of the other genera in the Trametes-group sensu Ryvarden
(1991), including the results from the present study (Figs. 2, 4)
is summarized in Table 1. Daedaleopsis, Earliella, Hexagonia,
Megasporoporia, and Microporus appear as separate genera
from each other and from Trametes. The genus Datronia, and
also Lentinus, appear intermixed with species of Polyporus s.l.
This result was already reported by Sotome & al. (2008) and
confirmed in our five-marker dataset but a formal taxonomic
proposal involving these genera has yet to be made.
ACKNOWLEDGEMENTS
Financial suppor t from the NSF through the PolyPEET grant (DEB
0933081) is gratefully acknowledged. Dimitrios Floudas, Loreen Meyer,
Claudia Osorio, David Marcano, Kim Pivorunas, Nathan Kallen, and
Drew Minnis collected some of the specimens and helped producing
some of the molecular data presented here. Zheng Wang originally
designed the PCR protocol used for protein-coding genes. Beatriz Ortiz-
Sa nt ana , Dan Lind ner, and the pe rso n nel at For e st Pr odu ct s Lab (M a di-
son, Wisconsin) are thanked for managing the loans of specimens and
culture collections. Comments f rom two reviewers were ver y helpful
for improving the manuscript.
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Appendix.
GenBank accession numbers for the sequences used in the phylogenetic analyses. For the 117 new isolates studied here the following i nformation
is given: Geog raphic origin, collector and collection number (herbar ium, if d ifferent from the Forest Product Lab, Madison, U.S.A.). Nomenclatural authori-
ties are given after the first ment ion of a taxon. Preexist ing Gen Bank sequences are marked with an asterisk (*). Sequences are given in the following order:
nrLSU, nrITS, RPB1, RPB2, and TEF1.
Amauroderma rugosum (Blume & T. Nees) Torrend; AB368061*, AB509712*, –, AB368119*, –. Boletopsis leucomelaena (Pers.) Fayod; D Q15 411 2*,
DQ484064*, GU187494*, GU187820*, GU187763*. Cerrena aurantiopora J.S. Lee & Y.W. Lim; FJ821521*, FJ821532*, –, –, –. Cerrena consors (Berk.) K.S.
Ko & H.S. Jung; FJ821517*, FJ821528*, –, –, –. Cerrena unicolor (Bull. : Fr.) Mur rill; FJ821524*, FJ821534*, –, –, –. Climacodon septentrionalis (Fr. : Fr.)
P. Ka r st ; AY68 4165*, AY8 540 82 *, AY86 4872*, AY 78 0941*, AY 88 5151*. Coriolopsis aspera (Jungh.) Teng; AY351956*, –, –, –, –. Coriolopsis cf. byrsina
(M ont .) Ry var d en; U.S.A. (MS), R.W. Davidson, FP-105050-Sp, JN164788, JN165001, JN164838, JN164871, JN164879. Coriolopsis cf. caperata ( Ber k.) Mu r-
rill; Venezuela, C. Rodríguez, CR-22, JN164789, JN164999, JN164837, JN164870, JN164880. Coriolopsis cf. caperata; Venezuela, C. Rodrígue z, CRM-77, –,
JN165000, –, –, –. Coriolopsis gallica (Fr. : Fr.) Ryvarden; U.S.A. (AZ), R.L. Gilbertson, RLG-7630-sp, JN164814 , J N165013, J N16 4821, JN16 48 69. Coriolop-
sis gallica; U.S.A. (IL), C. T homs, FP- 66249-T, –, JN165011, –, –, –. Coriolopsis gallica; U.S.A. (MS), E.R. Toole, T328, –, JN165010, –, –, –. Coriolopsis
gallica; U.S.A. ( NM), H.H. Burdsall, Jr., HHB-1207sp, –, JN165014, –, –, –. Coriolopsis gallica; U.S.A. (OR), J. Hunt, FP-91633-T, –, JN165012, JN164845,
JN164868, –. Coriolopsis gallica; U.S.A. (OK), A. Pakula, MC-42, –, JN164998, –, –, –. Coriolopsis gallica; U.S.A. (WI), T.J. Volk & T.W. Darmono,
FP-102596A-Sp, –, J N164997, –, –, –. Coriolopsis polyzona (Pers.) Ryvarden; Ghana, B. Kumi-Woode, BKW-0 01, –, JN164980, –, –, –. Coriolopsis polyzona;
Ghana, B. Kumi-Woode, BKW-004, JN164790, JN164978, JN164844 & JN164846, JN164856, JN164881. Coriolopsis polyzona; Ghana, B. Kumi-Woode, BKW-
017, –, J N164977, –, –, –. Coriolopsis polyzona; Venezuela, O. Holmquist, OH-272-Sp, –, J N164979, –, –, –. Coriolopsis sanguinaria (Klotzsch) Teng;
AY351950*, –, –, –, –. Coriolopsis trogii (Berk.) Domanski; U.S.A. (MI), M. J. Lar se n, MJL-3721- Sp, –, J N164994, –, –, –. Coriolopsis trogii; U.S.A. (MI),
M. J. Larsen , MJL -3725- S p, –, J N164995, –, –, –. Coriolopsis trogii; U.S.A. (MN), R.L. Gilbertson, RLG-9577-R, –, JN164996, –, –, –. Coriolopsis trogii;
U.S.A. (N Y), R.L. Gilbertson, R LG-4286-Sp, JN164808, JN164993, JN164820, JN164867, JN164898. Cryptoporus volvatus (Peck) Shear; AF393050*, –, –,
–, –. Daedalea querc ina (L. : Fr.) Pers.; AF518613*, FJ403212*, –, –, –. Daedaleopsis confragosa (B ol to n : Fr.) J. Sch röt.; AB368 06 2*, GU731549*, –, AB36 812 0*,
–. Datronia mollis (Sommerf. : Fr.) Donk; U.S.A. (M I), H.H. Burdsall, Jr., HHB-3480, –, JN165005, –, –, –. Datronia mollis; U.S.A. (MT), R.L. Gilbertson,
RLG- 6304-Sp, JN164791, JN165002, JN164818, JN164872, J N164901. Datronia mollis; U.S.A. ( NY), R.L. Gilbertson, RLG-5624-Sp, –, J N165007, –, –, –.
Datronia scutellata (Schwein.) Gilb. & Ryvarden; U.S.A. (MN), D.W. French, RLG-9584-R, –, JN165003, –, –, –. Datronia scutellata; U.S.A. (M N), D. W.
Fre nch, R LG -9584-T, JN164792, JN165004, JN164817, JN164873, JN164902. Dentocorticium sulphurellum (Peck) M.J. Larsen & Gilb.; Canada (QC), J.H.
Ginns, T609, JN164815, JN165015, –, JN164875, –. Dentocorticium sulphurellum; U.S.A., collector unknown, FP-11801, –, JN165018, JN164841, JN164876,
JN1649 03. Dentocorticium sulphurellum; U.S.A. ( NY), J.L. Lowe, L14474, –, JN165016, –, –, –. Dentocorticium sulphurellum; U.S.A. (NY ), P. Rosk in,
PR52 , –, JN165017, –, –, –. Earliella scabrosa (Pers.) Gilb. & Ryvarden; Puerto Rico, C. Rodrígue z, PR-1209, JN164793, JN165009, JN164819, JN164866,
JN1648 94 . Earliella scabrosa; Venezuela, C. Rodríguez, CR-45, –, JN164992, –, –, –. Earliella scabrosa; Venezuela, C. Rodríguez, CR-95, –, JN165008, –,
–, –. Earliella scabrosa; Venezuela, C. Rodrígu ez, CR-Irel, –, JN165006, –, –, –. Fomitopsis pinicola (Sw. : Fr.) P. Karst.; AY684164*, AY854083*, AY864874*,
AY786 05 6*, AY885152*. Ganoderma lucidum (Curtis : Fr.) P. Karst.; AB368068*, EU021460*, –, AB368126*, –. Ganoderma tsugae Murrill; AY684163*,

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DQ206985*, –, DQ408116*, DQ059048*. G rifola frondosa (Dickson : Fr.) Gray; AY629318*, AY854084*, AY864876*, –, AY885153*. Grifola sordulenta
(Mont.) Singer; AY645050*, AY854085*, AY864877*, AY786058*, AY885154*. Hexagonia hir ta (P. Beauv. : Fr.) Fr.; AY351944*, –, –, –, –. Hydnel lum geo -
genium (Fr.) Banker; AY631900*, DQ218304*, –, DQ408133*, DQ059053*. Irpex lacteus (Fr. : Fr.) Fr.; EU522839*, AB079264*, –, DQ408117*, –. Lentinus
squarrosulus Mont.; AB368071*, GU001951*, –, AB368129*, –. Lentinus tigrinus (Bull. : Fr.) Fr.; AB368072*, AF516520*, –, AB368130*, –. Lenzites betu-
linus (L. : Fr.) Fr.; U.S.A. (FL), H.H. Burdsall, Jr., HHB-9942-Sp, J N164794, JN164983, JN164822, JN164860, JN164895. Lenzites betulinus; U.S.A. (MA),
A. Justo, AJ150 (CUW), –, JN164915, –, –, –. Lignosus rhinocerot is (Cooke) Ryvarden; AB368074*, FJ380871*, –, –, –. Lopharia cinera scens (Schwein.)
G. Cunn.; U.S.A. (MS), R.W. Davidson, FP-105043-sp, JN164813, JN165019, JN164840, JN164874, JN164900. Megasporoporia setulosa (Henn.) Rajchenb.;
GU566007*, –, –, –, –. Microporus vernicipes (Berk.) Imazeki; AY351929*, AB531465*, –, –, –. Microporus xanthopus (Fr. : Fr.) Kuntze, A B368 075*,
AJ698132*, –, –, –. Phanerochaete chrysosporium Burdsall; AF287883*, AY854086*, AY864880*, –, AY885155*. Phlebia radiata Fr. : Fr.; AF287885*,
AY85 4087*, AY 86 4881*, AY 21850 2*, AY 88515 6*. Polyporus arcularius (Batsch : Fr.) Fr.; A B368081*, AF516523*, –, AB368138*, –. Polyporus brumalis
(P er s. : Fr.) Fr.; AB3 68 084*, FJ59 68 83*, –, A B368141*, –. Polyporus squamosus (Hud s. : Fr.) Fr.; AY629320*, DQ 267123*, DQ 8310 23*, DQ 408120*, DQ 028601*.
Polyporus tricholoma Mont .; AB368100*, AF516555*, –, AB36 8157 *, –. Polyporus udus Jungh .; AB368108*, AF51875 6*, –, AB36 8165*, –. Polyporus umbel-
latus (Pers. : Fr.) Fr.; A B368109*, EU442276*, –, AB368166*, –. Pseudofavolus cucullatus Mont.; AB368114*, AF516601*, –, AB368170*, –. Pycnoporus
cinnabarinus (Jacq. : Fr.) P. Karst.; China, Zheng Wang, ZW 02-30 (CUW), AY684160*, DQ411525*, JN164843 & JN164847, DQ408121*, DQ02860*. Pycno-
porus puniceus (Fr.) Ryvarden; FJ372708*, FJ372686*, –, –, –. Pycnoporus sanguineus (L. : Fr.) Murrill; Puerto Rico, S.A. Cantrell, PR-SC-95, JN164795,
JN164982 , J N16484 2, J N1648 58 , J N16489 7. Pycnoporus sanguineus; Venezuela, C. Rodríg uez, CR-35, –, JN164981, –, –, –. Tr ame te s aff. maxima ( Mo nt.)
A. David & Rajchenb.; Philippines, M.A. Eusebio, FPR I-376, –, JN164918, –, –, –. Tram ete s aff. maxima; Philippines, E.M. Mendoza, FPRI-401, JN164802,
JN164933, JN164 83 6, JN1648 63, JN16 48 84. Trametes conchifer (Schwein. : Fr.) Pilát; U.S.A. (MS), P.L. L entz, FP-10 6793-Sp, J N16 4797, JN16 4924 , J N16 4823,
JN1648 49, JN1648 87. Trametes conchifer; U.S.A. (NY), H.H. Burdsall, Jr., HHB-4940-Sp, –, JN164988, –, –, –. Trametes conchifer; U.S.A. (NY), J.L. Lowe,
L- 150 6 4-Sp , –, JN164926, –, –, –. Trametes conchifer; U.S.A. (SC ), A.S . Rhoad s, FP-103312-Sp, –, JN164925, –, –, –. Trametes conchifer; U.S.A. (VA),
F. Berry, FP-86583-R, –, JN164939, –, –, –. Trametes conchifer; U.S.A. (WI), Poulle & Dorworth, FP-102251-Sp, –, JN164987, –, –, –. Trametes cubensis
(M ont .) Sac c. ; U.S.A. (FL), A. Justo, AJ177 (CUW), J N164787, J N164905, –, –, –. Trametes cubensis; U.S.A. (MS), T.J. Volk, TJV-93-213-Sp, J N164798,
JN164923, J N16 48 34 , JN16 4865, J N16 48 83 . Trametes cubensis; Venezuela, C. Rodríguez, CR-96, –, JN164922, –, –, –. Trametes cubensis; Venezuela, C. Ro-
dríguez, CRM-90, –, JN164989, –, –, –. Trametes ectypa (Berk. & M.A. Cur tis) Gilb. & Ryvarden; U.S.A. (FL), A.S . R ho ad s, FP-103976 - Sp, –, JN164961, –,
–, –. Trametes ectypa; U.S.A. (FL), G.G. Hedgcock, FP-81034-R, –, JN164930, –, –, –. Trametes ectypa; U.S.A. (MS), E .R. Toole, FP-106037-T, JN164803,
JN164929, J N16 4824, J N16 48 48, J N164 88 8. Tramete s elegans (Spreng. : Fr) Fr.; Belize, K.K. Nakasone, FP-150762, –, JN164928, –, –, –. Trametes elegans;
country unknown, collector unknown, FRI- 437-T, –, JN164985, –, –, –. Trametes elegans; Philippines, L. Lodderau, FPRI-10, –, JN164973, –, –, –. Tram e te s
elegans; Philippines, M.A. Eusebio, FPRI-390, –, J N164921, –, –, –. Trametes elegans; Puerto Rico, D.J. Lodge & L. Fish, PR-1133, –, J N164937, –, –, –.
Trametes elegans; U.S.A. (FL), H.H. Burdsall, Jr., HHB-4626-Sp, –, J N164950, –, –, –. Trametes elegans; U.S.A. (FL), H.H. Burd sall, Jr., HHB- 6551-Sp, –,
JN164938, –, –, –. Trametes elegans; U.S.A. (GA), R.W. Davidso n, FP-105679 -Sp, JN164799, JN164944, JN164833, JN164861, JN164899. Trametes elegans;
U.S.A. (MS), R.W. Davidson, FP-105038-Sp, –, JN164951, –, –, –. Trametes elegans; Venezuela, C. Rodríguez, CR M-52, –, JN164986, –, –, –. Trametes elegans;
Venezuela, O. Holmquist , OH-271-Sp, –, JN164936, –, –, –. Trametes gibbosa (Pers. : Fr.) Fr.; UK, J.L. Lowe, L-11664-Sp, JN164800, JN164943, JN164831,
JN164859, JN164896. Trametes hirsuta ( Wulfen : Fr.) Pilát; U.S.A. (MA), A. Justo, AJ196 (CUW), –, JN164916, –, –, –. Trametes hirsuta; U.S.A. (MA),
A. Justo, AJ303 (CUW), –, JN164917, –, –, –. Trametes hirsuta; Costa R ica, J.L. Lowe, L-12964-Sp, –, JN164935, –, –, –. Trametes hirsuta;, Estonia, D.R .
Richter, DR-EST-7, –, JN164934, –, –, –. Trametes hirsuta; U.S.A. (AZ), H.H. Burdsall, Jr., HHB-8591-Sp, –, JN164953, –, –, –. Trametes hirsuta; U.S.A. (IL),
H.H. McKay, FP-103959-T, –, J N164952, –, –, –. Trametes hirsuta; U.S.A. (NY), R.L. Gilbertson, RLG -5133-T, JN164801, JN164941, JN164829, JN164854,
JN1648 91. Trametes hirsuta; U.S.A. (WI), M.J. Larse n, FP-101666-Sp, –, JN164942, –, –, –. Trametes junipericola Manjón, G. Moreno & Ryvarden; AY855915*,
AY684171*, –, –, –. Trametes ljubarskii Pilát; AY855911*, AY684174*, –, –, –. Tramete s maxima (Mont.) A. David & R ajchenb.; Venezuela, O. Holmquist,
OH -17 2- S p , –, JN164932, –, –, –. Trametes maxima; Venezuela, O. Holmquist, OH-189-Sp, J N1648 0 4, J N16 49 57, JN 164 816 , J N1648 64, JN1648 85. Tra metes
maxima; Venezuela, O. Holmquist, OH-241-Sp, –, JN164940, –, –, –. Trametes membranacea (Sw. : Fr.) Kreisel; Puerto Rico, E. Horak, PR-3264, –, JN164946,
–, –, –. Trametes membranacea; Puerto Rico, R. Petersen & S.A. Cantrell, PR-SC-69, –, JN164927, –, –, –. Trametes membranacea; Puerto Rico, R. Petersen
& S.A. Cantrell, PR- SC-82, JN164805, J N164945, JN164832, J N164857, JN164893. Trametes membranacea; Venezuela, C. Rodrígu ez, CRM-125, –, JN164956,
–, –, –. Trametes ochracea (Pers.) Gilb. & Ryvarden; Canad a (ON), M. J. Larsen , MJL-2103, –, J N164948, –, –, –. Trametes ochracea; U.S.A. (AK), H.H.
Burdsall, Jr., HHB-13429-Sp, –, JN164976, –, –, –. Trametes ochracea; U.S.A. (CO), J.L. Lowe, L-6773-R, –, JN164971, –, –, –. Trametes ochracea; U.S.A .
(MI), T.J. Volk, HHB-13445-Sp, JN164812, JN164954, JN164826, JN164852, JN164904. Trametes pavonia (Hoo k.) Ry v ard en; Puerto Rico, E. Setliff, PR-2178,
–, JN164959, –, –, –. Trametes pavonia; U.S.A. ( FL), A.S. Rhoads, FP-103050-Sp, J N164806, JN164958, JN164835, JN164862, JN164886. Trametes pocas
(Berk.) Ryvarden, AY351919*, –, –, –, –. Trametes pubescens (Schumach. : Fr.) Pilát; U.S.A. ( NY), J.L. Lowe, L-15318-Sp, –, JN164949, –, –, –. Tr am e te s
pubescens; U.S.A. (NY), J.L . Lowe, L-15711-Sp, –, JN164972, –, –, –. Trametes pubescens; U.S.A. (NY), J.L. Lowe, L-16093-S p, –, JN164964, –, –, –. Tram-
etes pubescens; U.S.A. (WI), F.F. Lombard & R .G. Payne, FP-101414-Sp, JN164811, JN164963, JN164827, JN164851, JN164889. Trametes pubescens; U.S.A.
(WI), Nishijima & Evans, FP-101536-T, –, JN164960, –, –, –. Trametes pubescens; U.S.A. (WI), Burdsall & Leslie, FP-101554-T, –, JN164962, –, –, –. Tram -
etes pubescens; U.S.A. (WI), H.H. Burdsall, Jr. & T.J. Volk, HHB-13585-Sp, –, JN164947, –, –, –. Tra met es sp.; South Korea, A.M. Minnis, AJ354 (CUW), –,
JN164990, –, –, –. Trametes suaveolens (L. : Fr.) Fr.; U.S.A. (AK), H.H. B urd sall, Jr., HHB -14170-T, –, JN164968, –, –, –. Trametes suaveolens; U.S.A. (WI),
H.H. Burdsall, Jr. & T.J. Volk, FP-102529-Sp, J N164807, J N16496 6, J N164828, JN164853, J N164890. Trametes suaveolens; U.S.A. (WI), H.H. Burdsall, Jr.
& T.J. Volk, FP-102529-T, –, JN164967, –, –, –. Trametes versicolor (L : Fr.) Lloyd; U.S.A. (FL), A. Justo, AJ146 (CUW), –, JN164912, –, –, –. Tr a me tes ve r -
sicolor; U.S.A. (FL), A. Justo, AJ164 (CUW), –, JN164909, –, –, –. Trametes versicolor; U.S.A. (FL), A. Justo, AJ170 (CUW), –, JN164907, –, –, –. Tr am e te s
versicolor; U.S.A. (FL), A. Justo, AJ183 (CUW), –, JN164911, –, –, –. Trametes versicolor; U.S.A. (MA), A. Justo, AJ119 (CUW), –, JN164914, –, –, –. Tr am-
etes versicolor; U.S.A. (MA), A. Justo, AJ151 (CUW), –, J N164913, –, –, –. Trametes versicolor; U.S.A. (MA), A. Justo, AJ190 (CUW), –, JN164908, –, –, –.
Trametes versicolor; U.S.A. (MA), A. Justo, A J198 (CUW), –, JN164906, –, –, –. Trametes versicolor; U.S.A. (MA), A. Justo, AJ207 (CUW), –, JN164910, –,
–, –. Trametes versicolor; Brazil (SP), A.R. Teixeira, Braz-16, –, JN164975, –, –, –. Trametes versicolor; U.S.A. (ID), M.J. Larsen, FP-134940-Sp, –, JN164920,
–, –, –. Trametes versicolor; U.S.A. (NY), M.J. Larsen, FP-135156-S p, JN164809, JN164919, JN164825, JN164850, JN164878. Trametes versicolor; U.S. A.
(OR), M. J. Larsen, FP-133494-Sp, –, JN164931, –, –, –. Trametes versicolor; U.S.A. (WI), T.J. Volk & al., FP-102607-Sp, –, J N164955, –, –, –. Tr am ete s
versicolor; U.S.A. (WI), E.B. Dor worth, FP102316sp, –, JN164984, –, –, –. Trametes versicolor; U.S.A. (WI ), M.T. Banik, HHB-12282-Sp, –, JN164974, –, –,
–. Trametes versicolor; Argenti na, H. Lenko, BAFC-285, –, JN164965, –, –, –. Trametes villosa (Sw.) Kreisel, U.S.A. (FL), A.S. Rhoads, FP-86588-Sp, –,
JN164970, –, –, –. Tramete s villosa (Sw.) Kreisel, U.S.A. (T N), B.S. Crandall, FP-71974-R, JN164810, JN164969, JN164830, JN164855, JN164892. Tr ame -
topsis cervina (Schwein.) Tomsovský; U.S.A. (FL), A. Justo, AJ185 (CUW), –, JN164991, –, –, –. Trametopsis cervina; U.S.A. (FL), A. Justo, AJ189 (CUW),
–, JN165021, –, –, –. Trametopsis cervina; U.S.A. (MS), T. J. Volk, TJ V-93-216 -Sp , JN164796, JN165020, JN164839, JN164877, JN164882.
Appendix.
Continued.