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Fungal Diversity
41
Albatrellus piceiphilus sp. nov. on the basis of morphological and molecular
characters
Cui, B.K.1, Wang, Z.2 and Dai, Y.C.1*
1Beijing Forestry University, Beijing 100083, P.R. China
2Department of Biology, University of Iowa, Iowa City, IA 52242, USA
Cui, B.K., Wang, Z. and Dai, Y.C. (2008). Albatrellus piceiphilus sp. nov. on the basis of morphological and molecular
characters. Fungal Diversity 28: 41-48.
Albatrellus piceiphilus is described as new from Gansu Province, China based on morphological characters and
sequence data from the ITS region of nuclear ribosomal DNA. The new species is characterized by its pale yellow to
yellowish-brown basidiocarps, simple septate generative hyphae, slightly thick-walled and distinctly amyloid
basidiospores, and its habit on the ground in a Picea crassifolia forest. Parsimony analyses were applied to the ITS
dataset. Our results suggest that Albatrellus is not monophyletic, and two clades of Albatrellus species are recognized.
One clade including A. piceiphilus, A. citrinus and A. ovinus is strongly supported, which probably represents the core
of Albatrellus in the russuloid clade. Albatrellus syringae is supported as a sister group to some polypores in the
polyporoid clade, distantly away from the other Albatrellus species in the tree.
Key words: Albatrellus, ITS, phylogeny, polypore, taxonomy.
Article Information
Received 13 January 2007
Accepted 18 July 2007
Published online 15 January 2008
*Corresponding author: Y.C. Dai; e-mail: yuchengd@yahoo.com
Introduction
Albatrellus species are fairly common in
northern temperate forests, and they usually
produce medium to large fleshy fruiting bodies
of various colours. Traditionally, species of
Albatrellus has been classified as polypores
due to their poroid hymenophore on the lower
surface of basidiocarps. Most species are
terrestrial having mycorrhizal connections with
trees, while some may be wood-decaying taxa
(Pouzar, 1972; Canfield, 1981; Ryvarden and
Gilbertson, 1993). Basidiospores of Albatrellus
species are ellipsoid to subglobose, smooth,
and either amyloid or negative in Melzer’s
reagent. Recent phylogenetic studies using
molecular markers showed that Albatrellus
consists of two separate groups with affinities
to the polyporoid clade and the russuloid clade
(Gardes and Bruns, 1996; Hibbett et al., 1997;
Hibbett and Thorn, 2001; Larsson and Larsson,
2003; Greslebin et al., 2004). While most of
the species belong to the russuloid clade, the
closest relatives of Albatrellus syringae
(Parmasto) Pouzar are members of Antrodiella
Ryvarden & I. Johans., Junghuhnia Corda,
Steccherinum Gray and Ceriporiopsis
Domański, of the polyporoid clade (Binder et
al., 2005).
During a survey of polypores in
Northwest China, two stipitate poroid speci-
mens of Aphyllophorales were collected on the
ground in a Picea crassifolia forest. Their
macroscopic and microscopic features fit well
with Albatrellus but do not match with the
descriptions of any existing species in the
genus. Members of both the polyporoid and the
russuloid clades are highly diverse in their
fruiting body morphology and microscopic
characters, such as characters of spores and
hyphae (Binder et al., 2005). To confirm the
affinity of the taxon and to infer the
relationships within the genus Albatrellus,
phylogenetic analyses were carried out based
on the internal transcribed spacer sequences of
ribosomal DNA (ITS) sequences data, in
42
addition to comparative studies of morpholo-
gical characters. ITS sequences have been
shown particularly valuable for the classifi-
cation of fungi at the species level (e.g., Yao et
al., 1999; Liu et al., 2001; Ryman et al., 2003).
With accumulating ITS data of Albatrellus
species in the GenBank, a phylogenetic
reclassification of the genus becomes possible.
The results of both the morphological and
DNA studies supported the recognition of the
newly found species and its placement in
Albatrellus, and the taxon is therefore
described as new.
Materials and methods
Morphological studies
The studied specimens are deposited at
the Herbarium of the Institute of Applied
Ecology, Chinese Academy of Sciences (IFP).
Collections were examined in the microscope,
extensive measurements in particular of spore
dimensions were made, and pictures were
drawn from slide preparations stained with
Cotton Blue and Melzer’s reagent. Spores were
measured from sections cut from the tubes. IKI
= Melzer's reagent, KOH = 5% potassium
hydroxide, and CB = Cotton Blue. CB+ =
cyanophilous and CB– = acyanophilous, and
IKI– = both inamyloid and indextrinoid. In
presenting the variation in the size of the
spores, 5% of the measurements were excluded
from each end of the range, and are given in
parentheses. In the text the following
abbreviations are used: L = mean spore length
(arithmetical mean of all spores), W = mean
spore width (arithmetical mean of all spores),
Q = variation in the L/W ratios between the
specimens studied (quotient of the mean spore
length and the mean spore width of each
specimen), n = number of spores measured
from given number of specimens. The width of
a basidium was measured at the thickest part,
and the length was measured from the apex
(sterigmata excluded) to the basal septum.
Sections were studied at magnification up to
×1000 by using a Nikon Eclipse E600
microscope and phase contrast illumination.
Drawings were made with the aid of a drawing
tube. Colour descriptions follow Petersen
(1996) and Anonymous (1969).
Molecular techniques
DNA was extracted from dried herbarium
materials following the methods of Grades and
Bruns (1993), Yao et al. (1999) and Liu et al.
(2001), with some modifications. About 10-30
mg dried specimen material was ground in
liquid nitrogen, and then was mixed well with
600 µl pre-warmed 2.5% CTAB extraction
buff, containing 100 mM Tris (pH 8.0), 20 mM
EDTA (pH 8.0), and 1.4 mol/L NaCl, followed
by a water bath at 65ºC for 50 to 80 minutes.
600µl chloroform-isoamyl alcohol (24:1 in
volume) was added to precipitate unwanted cell
components and proteins. 30 µl 3 M NaAc was
added to the aqueous phase followed by 200µl
cold (–20ºC) isopropanol to precipitate the
DNA. The precipitated DNA was collected by
centrifuge, washed with 500 µl ice-cold 70%
ethanol, and dried, then dissolved in 50 µl
sterile distilled water and stored at –20ºC.
PCR amplifications follow the methods
of Yao et al. (1999) and Liu et al. (2001).
Primers for the PCR amplification were ITS1
and ITS4 described by White et al. (1990). 50
µl reaction mixtures for the PCR were:
genomic DNA in optimum concentration, 0.5
µM of each primer, 10 mM Tris-HCl (pH 8.0),
50 mM KCl, 0.08% Nonidet P40, 1.5 mM
magnesium chloride (MgCl2), 200µM each of
dATP, dCTP, dGTP and dTTP (Promega Co.,
USA), and 2 units Taq DNA polymerase
(Sangon Ltd., Canada). Mineral oil (about 40
µl) was overlaid on the reaction mixture. The
PCR was performed in a MiniCyclerTM (MJ
Research Inc., USA) for 35 cycles, including
94ºC for 35 s, 50ºC for 45 s, and 72ºC for 45 s.
PCR products used for sequencing were
cleaned using the QIAquick PCR purification
columns (Qiagen Ltd., UK). The DNA
sequences were determined through direct PCR
sequencing by GeneCore Biotechnologies
(Shanghai, China). Sequences were edited with
Sequencher version 3.1 (geneCodes Corpo-
ration, Ann Arbor, Michigan) and deposited in
GenBank (Table 1).
Phylogenetic analyses
Taxa used in the phylogenetic analyses
are listed in Table 1. One data set was prepared
based on ITS sequences of 25 taxa, among
which 18 taxa represent nine Albatrellus
Fungal Diversity
43
Table 1. Species and sequences database accession numbers used in this study.
Species name Collection no. Locality GenBank no.
Albatrellus caeruleoporus (Peck) Pouzar
Albatrellus citrinus Ryman
Albatrellus citrinus Ryman
Albatrellus citrinus Ryman
Albatrellus ellisii (Berk.) Pouzar
Albatrellus flettii Morse ex Pouzar
Albatrellus ovinus (Schaeff.:Fr.) Kotl. & Pouzar
Albatrellus ovinus (Schaeff.:Fr.) Kotl. & Pouzar
Albatrellus ovinus (Schaeff.:Fr.) Kotl. & Pouzar
Albatrellus ovinus (Schaeff.:Fr.) Kotl. & Pouzar
Albatrellus piceiphilus B.K. Cui & Y.C. Dai
Albatrellus piceiphilus B.K. Cui & Y.C. Dai
Albatrellus similis Pouzar
Albatrellus subrubescens (Murrill) Pouzar
Albatrellus syringae (Parmasto) Pouzar
Albatrellus syringae (Parmasto) Pouzar
Albatrellus syringae (Parmasto) Pouzar
Albatrellus syringae (Parmasto) Pouzar
Antrodiella americana Ryvarden & Gilb.
Antrodiella pallasii Renvall et al.
Antrodiella romellii (Donk) Niemelä
Auricularia sp.
Hericium alpestre Pers.
Junghuhnia collabens (Fr.) Ryvarden
Leucophleps spinispora Fogel
K.A. Harrison 8825
Muskos 850928
Ryman 9077
Strid 16319A
Fransson 2
Fransson 3
Ryman 9132
Danell 11/8 00
Cui 2220
Cui 2221
Ryman 6085
Cui 2183
Cui 2177
Ryman 6388
K.A. Harrison 6224
Haikonen 14727
T. Renvall 89a
Renvall 3501
F3082
Sweden
Sweden
Switzerland
Sweden
Sweden
Sweden
Sweden
China
China
Sweden
China
China
Sweden
Finland
Finland
AY963565
AY198190
AY198192
AY198194
AY621803
AY061738
AY198198
AY198201
AY198202
AY198203
DQ789396
DQ789397
AY963566
AY198208
DQ789394
DQ789395
AY198209
AY621804
AF126877
AF126896
AF126902
DQ200918
AY534580
AF533965
AY621749
species. ITS sequences of all the Albatrellus
species included in this study were used as
references to blast against GenBank, and
sequences of Leucophleps spinispora Fogel,
Hericium alpestre Pers., and four polypores
including Junghuhnia collabens (Fr.) Ryvarden
and three species of Antrodiella, were
downloaded from GenBank for high sequence
similarity with Albatrellus species. DNA
sequences were aligned with ClustalX using
default setting (Thompson et al., 1997) and
further adjusted by eye in the data editor of
PAUP* 4.0b (Swofford, 1999). ITS sequences
were highly variable across sampled taxa, but
sequences of two groups, one including
Albatrellus syringae and four polypores, and
the other comprising the other Albatrellus
species, Leucophleps Harkn., and Hericium
Pers., are alignable. Since our goal in this study
was to investigate groups of closely related
species within the genus Albatrellus under
current concept, gaps were introduced for an
arbitrary alignment with these two classes and
contributed partially to the long branch length
observed in our tree. A proper outgroup for
inferring relationships among these groups
using ITS data is not possible. The data set in
this study was arbitrarily rooted with Hericium
alpestre. Data set was analyzed in PAUP* 4.0b
(Swofford, 1999), with gaps treated as missing
data.
Parsimony analyses were performed
using equally weighting of characters and
transformations. Heuristic searches were
performed with one thousand replicate
searches, each with one random taxon addition
sequences. MAXTREES set to autoincrease,
and TBR branch swapping. Robustness of
individual branches was estimated by
maximum parsimony bootstrap proportions
(BP), using 500 replicates, each consisting of a
single heuristic search with 50 random taxon
44
Fig. 1. Microscopic structures of Albatrellus piceiphilus B.K. Cui & Y.C. Dai (drawn from the holotype). a.
Basidiospores. b. Basidia and basidioles. c. Hyphae from trama. d. Hyphae from context. e. Hyphae from stipe.
addition sequences, MAXTREES set to
autoincrease, and TBR branch swapping.
Results
Albatrellus piceiphilus B.K. Cui & Y.C. Dai,
sp. nov. (Fig. 1)
MycoBank: 510979.
Etymology: Piceiphilus (Lat.), growing on Picea.
Carpophorum annuum, stipitatum. Facies
pororum pallide luteola vel luteola, pori 2-4 per mm.
Systema hypharum monomiticum, hyphae generatoriae
sine fibulis, hyphae contexti 5-10 µm in diam. Sporae
ellipsoideae vel perlate ellipsoideae, amyloideae, 4.4-5 ×
3.5-4.1 µm.
Fruitbody - Basidiocarps annual, terres-
trial, centrally or laterally stipitate, solitary, or
several pilei fused to form more compound
cluster, fleshy and watery, without odour or
taste when fresh, becoming fragile with an
unpleasant odour upon drying. Pileus more or
less circular, sometimes depressed and infundi-
buliform, 7-12 cm in diam., and up to 6 mm
thick at centre. Pileal surface yellowish to
yellowish brown, smooth when fresh,
becoming clay buff to pale brown, wrinkled
Fungal Diversity
45
Fig. 2. Phylogenetic analyses of Albatrellus piceiphilus and the related species based on ITS region sequence data.
Bootstrap less than 50% are not shown.
upon drying, azonate; margin sharp, sometimes
lobed, yellowish when fresh, becoming pale
brown upon drying. Pore surface pale yellow
to yellow when fresh, becoming pale yellowish
brown to olivaceous buff or cinnamon buff
when dry; pores angular, 2-4 per mm, tube
mouths thin, entire to slightly lacerate. Context
yellowish, fleshy and watery when fresh,
becoming yellowish to olivaceous buff, with a
dark line next to the tubes, fragile to brittle
upon drying, up to 3 mm thick. Tubes
concolorous with pore surface, soft when fresh,
become brittle when dry, up to 3 mm long.
Stipe pale brownish vinaceous, watery and
fleshy when fresh, becoming pale greyish
brown, wrinkled and brittle upon drying, up to
4 cm long and 1 cm in diam.
Hyphal structure — Hyphal system
monomitic; generative hyphae simple septate,
IKI–, CB–; tissue more or less darkening but
otherwise unchanged in KOH.
Context — Contextual hyphae hyaline,
thin- to slightly thick-walled, simple septate,
occasionally branched, interwoven, 5-10 µm in
diam., sometimes inflated, up to 23 µm in
diam.; gloeoplerous hyphae absent; hyphae at
stipe similar to contextual hyphae.
Tubes — Tramal hyphae hyaline, thin-
walled, frequently simple septate, rarely
branched, more or less straight and parallel
46
along the tubes, 2.6-5 µm in diam. Cystidia and
cystidioles absent; basidia clavate, with four
sterigmata and a basal simple septum, 15-25 ×
5-7 µm; basidioles in shape similar to basidia,
but slightly smaller.
Spores — Basidiospores ellipsoid to
broadly ellipsoid, hyaline, slightly thick-
walled, smooth, distinctly amyloid, CB-, (4.2-)
4.4-5(-5.2) × (3.4-)3.5-4.1 (-4.4) µm, L = 4.78
µm, W = 3.85 µm, Q = 1.24-1.25 (n=60/2).
Habitat: On ground in Picea crassifolia
forest.
Known distribution: Gansu Province of
China.
Material examined: CHINA, Gansu Province,
Yuzhong County, Xinglongshan Nature Reserve, on
ground in Picea crassifolia forest, 26.VIII.2005 Cui
2221 (holotype in IFP, isotypes in H and HMAS);
Gansu Province, Yuzhong County, Xinglongshan Nature
Reserve, on ground in Picea crassifolia forest,
26.VIII.2005 Cui 2220 (paratype in IFP).
Relationships among the Albatrellus
species were investigated using ITS region
from 24 taxa. The data set had an aligned
length of 772 base pair with 300 parsimony-
informative positions.
Equally weighted parsimony analysis
yielded 24 equally parsimonious trees of 900
steps with a consistency index CI = 0.718 (Fig.
2). Two strongly supported clades were
recognized within targeted Albatrellus species:
One consisted of A. citrinus Ryman, A.
piceiphilus, A. ovinus (Schaeff.:Fr.) Kotl. &
Pouzar, A. subrubescens (Murrill) Pouzar and
A. similis Pouzar (BP = 100%), and within the
clade, the new species A. piceiphilus shared a
clade with A. citrinus (BP = 81%) with A.
ovinus as weakly supported sister groups (BP =
70%). Relationships of A. flettii Morse ex
Pouzar, A. caeruleoporus (Peck) Pouzar, and A.
ellisii (Berk.) Pouzar to the clade and
Leucophleps species were not resolved, though
they were supported as clade with BP = 100%.
Another clade (BP = 100%) consisted of four
isolates of A. syringae with almost no
difference in sequences, and this clade was
strongly supported as the sister group to four
investigated polypores (BP = 100%).
Discussion
Albatrellus piceiphilus is characterized
by its simple septate generative hyphae,
slightly thick-walled and distinctly amyloid
basidiospores, and by its occurrence on ground
in Picea crassifolia forests. Its pileal surface is
yellowish to yellowish brown when fresh,
becoming clay buff to pale brown upon drying;
pore surface pale yellow to yellow when fresh,
becoming pale yellowish brown to olivaceous
buff or cinnamon buff when dry. ITS
phylogeny in this study strongly supports a
close relationship among A. citrinus, A.
piceiphilus, A. ovinus, A. subrubescens and A.
similis, some of which have already been
shown to belong to the russuloid clade
(Larsson and Larsson, 2003; Binder et al.,
2005).
Phylogenetically, Albatrellus piceiphilus
appeared to be closely related to A. citrinus,
and both taxa having yellowish basidiocarps,
simple septate generative hyphae, distinctly
amyloid basidiospores and link with Picea.
Basidiocarps of A. citrinus are white when
fresh, and its context has no special odour
when dry (Ryman et al., 2003). On the
contrary, Albatrellus piceiphilus has yellowish
basidiocarps when fresh, and it has strongly
unpleasant smell upon drying. In addition,
basidiospores in Albatrellus piceiphilus are
slightly wider than those in A. citrinus (4.4-5 ×
3.-4.1 µm, L = 4.78 µm, W = 3.85 µm, Q =
1.24-1.25 vs. 4-4.7 × 3-3.6 µm, L = 4.3 µm, W
= 3.2 µm, Q = 1.3-1.4, Niemelä, 2005).
Albatrellus citrinus was reported from China
by Zheng et al. (2004), which has thin-walled
basidiospores, while the spore size (4.5-5.5 ×
3-4.5 µm) given by them is almost identical to
A. piceiphilus.
Albatrellus piceiphilus is easily confused
with A. ovinus, in particular because of the
pale brown pileal colour, simple septate
generative hyphae, and slightly thick-walled
basidiospores (Gilbertson and Ryvarden,
1986; Ryvarden and Gilbertson, 1993).
However, the latter species has gloeoplerous
hyphae, and inamyloid and slightly smaller
basidiospores (3.7-4.2 × 3.1-3.5 µm, Niemelä,
2005).
Albatrellus piceiphilus resembles A.
subrubescens by having simple septate
generative hyphae, and distinctly amyloid
basidiospores (Gilbertson and Ryvarden,
1986; Ryvarden and Gilbertson, 1993),
Fungal Diversity
47
however, A. subrubescens has gloeoplerous
hyphae and smaller basidiospores (3.8-4.6 ×
2.8-3.3 µm, Niemelä, 2005).
Another Asian species, Albatrellus
dispansus (Lloyd) Canf. & Gilb., has yellow
basidiocarps; it was reported in Japan (Imazeki
et al., 1988), and we found it from central
China, too. However, its pore surface is white
and with a fragrant odour when fresh, and its
basidiospores are negative in Melzer’s reagent.
Macroscopically Albatrellus piceiphilus
is similar to A. syringae, but the latter species
has concentrically zonate pileal surface,
clamped generative hyphae, and smaller,
apically tapering basidiospores (3.8-4.7 × 2-3.5
µm, L = 4.24 µm, W = 3.15 µm, Dai et al.,
2004; Niemelä, 1970). In addition, Albatrellus
syringae was reported as a terrestrial species,
mostly on lawn or in angiosperm forests, but it
was recorded in spruce forest in Italy
(Ryvarden and Gilbertson, 1993). We found
this species on ground in Salix forest of
northeast China, and on ground in spruce forest
in northwest China. Also the ITS sequences of
Albatrellus syringae are distinctly different
from the other species of Albatrellus, our
analysis of ITS regions placed Albatrellus
syringae in the polyporoid clade, which
congruent with previous studies (Bruns et al.,
1998; Binder and Hibbett, 2002; Ryman et al.,
2003).
Acknowledgements
Special thanks are due to Mr. Mingyun Huang
(Shenyang, China) for companion on field trips. Mr.
Xiaoyong Liu, Mr. Lei Jiao (Beijing, China), Miss Juan
Li and Mr. Changjun Yu (Shenyang, China) also are
thanked for help in obtaining DNA sequences. We
express our gratitude to Prof. Teuvo Ahti and Dr. Tuomo
Niemelä (H, Finland) for revising the Latin description
and improving the manuscript. The research was
financed by the Ministry of Science and Technology of
China (Project No. 2005DFA30280).
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(Aphyllophorales, Albatrellaceae) from China.
Mycotaxon 90: 291-299.
