Mycol Progress (2014) 13:211–217

DOI 10.1007/s11557-013-0906-6

ORIGINAL ARTICLE

A new species of Amanita section Lepidella from South China

Wang-Qiu Deng & Tai-Hui Li & Peng Li & Zhu L. Yang

Received: 21 March 2013 / Revised: 7 May 2013 / Accepted: 10 May 2013 / Published online: 11 June 2013
# German Mycological Society and Springer-Verlag Berlin Heidelberg 2013

Abstract Amanita macrocarpa from Guangdong Province More than 80 taxa of Amanita have been reported from
of China is described. The new species is a large, brownish China (Yang 2005; Zhang et al. 2010), and about 50
orange to light brown mushroom, characterized by numer- Amanita species have been reported in Guangdong, a prov-
ous pyramidal warts on the pileus, yellowish lamellae, a ince in South China with rich mushroom resources.
large annulus at the middle of the stipe, and amyloid ellip- Recently, a species, morphologically different from any
soidal basidiospores. Phylogenetic analyses of the new spe- known Amanita taxon, was discovered in Guangzhou. To
cies and related taxa based on the large subunit of nuclear confirm the affinity of the new taxon and infer its evolu-
ribosomal DNA (LSU) and the internal transcribed spacer tionary relationships to similar species, phylogenetic analy-
(ITS) sequences are provided. Morphological and molecular ses were performed based on the large subunit of nuclear
data show that A. macrocarpa is a member of Amanita, ribosomal DNA (LSU) and internal transcribed spacer (ITS)
section Lepidella, and clearly different from any known sequences.
taxon of the section.

Keywords Agaricales . Guangzhou . Taxonomy Materials and methods

Morphological studies
Introduction
Specimens were annotated and photographed in the field,
Amanita Pers. is a widespread basidiomycetous genus com- dried in an electric oven, and deposited in the Fungal
prised of about 500 species (Yang 2005; Kirk et al. 2008). Herbarium of Guangdong Institute of Microbiology
The genus is divided into two subgenera, Amanita and (GDGM). Color description was according to Kornerup
Lepidella (E.-J. Gilbert) Veselý using traditional classifica- and Wanscher (1978). Spores, basidia and veil remnants
tion methods based on morphological and anatomical char- were observed in 50 g/L KOH, Melzer’s reagent or 10 g/L
acters (Corner and Bas 1962; Bas 1969). Subgenus Amanita Congo Red, and the pileipellis was examined in 50 g/L
includes sections Amanita, Caesareae Singer, and KOH or H2O. In the description of basidiospores, the ab-
Vaginatae (Fr.) Quél., while subgenus Lepidella includes breviations [n/m/p] denote n basidiospores measured from
sections Amidella (E.-J. Gilbert) Konrad & Maubl., m basidiomata of p collections. The notation (a–)b–c(−d) is
Lepidella, Phalloideae (Fr.) Quél., and Validae (Fr.) Quél. used to describe basidiospore dimensions where the range
(Yang 1997, 2005). b–c represents 90% or more of the measured values, and ‘a’
and ‘d’ are the extreme values. Q refers to the length/width
W.<Q. Deng : T.<H. Li (*) : P. Li ratio of an individual basidiospore; Qm refers to the average
Guangdong Provincial Public Laboratory of New Microbial
Application Technology, Guangdong Key Laboratory of Microbial Q of all basidiospores±sample standard deviation.
Culture Collection and Application, Guangdong State Key
Laboratory of Applied Microbiology (Ministry-Guangdong
DNA extraction, PCR amplification and DNA sequencing
Province Jointly Breeding Base) South China, Guangdong
Institute of Microbiology, Guangdong Guangzhou 510070, China
e-mail: [email protected] Genomic DNA was extracted from specimens using the
modified CTAB procedure of Doyle and Doyle (1987).
Z. L. Yang
ITS rDNA was amplified using primers ITS1 and ITS4,
Key Laboratory of Biodiversity and Biogeography, Kunming
Institute of Botany, Chinese Academy of Sciences, Heilongtan, and the nuclear LSU region was amplified with primers
Kunming 650201, China LR0R and LR5 (White et al. 1990). Reaction mixtures
212 Mycol Progress (2014) 13:211–217

(20 μL) contained 0.5 μL template DNA, 8.5 μL distilled macrocarpa to be distinctive in the section Lepidella with
water, 0.5 μL of each primer, and 10 μL PCR mix a high bootstrap value, and to be closely related to A.
[DreamTaq™ Green PCR Master Mix (2×), Fermentas]. japonica among the selected taxa (Fig. 2).
Amplication conditions were 35 cycles of 95°C for 30 s,
52°C for 30 s, and 72°C for 1 min, followed by a final Taxonomy
extension at 72°C for 10 min. Amplified PCR products were
purified, sequenced, and edited by Invitrogen Biotechnology Amanita macrocarpa W. Q. Deng, T.H. Li & Zhu L. Yang,
(Beijing). Sequences generated in this study have been depos- sp. nov. Figs. 3 and 4
ited in GenBank. Other sequence data used for phylogenetic MycoBank no.: MB 803263
analyses were download from GenBank (Table 1). Pileus 150–240 mm broad, covered with numerous pale
dirty brown to brownish orange pyramidal warts; yellowish
Alignment and phylogenetic analyses lamellae; subclavate stipe without distinct boundary be-
tween stipe and the bulb; large annulus at the middle stipe;
Sequences were aligned and edited using ClustalX (Tompson presence of clamps common; amyloid ellipsoidal basidio-
et al. 1997) and Bioedit (Hall 1999). Phylogenetic analyses spores (6–)7–9×(4–)5–6 μm; Remnants of volva consisting
were performed with MEGA v.5.0 Beta (Tamura et al. 2011). of moderately abundant to abundant hyphae intermixed with
A phylogenetic tree was constructed using the Maximum abundant inflated cells.
Likelihood method. Bootstrap values were calculated from Type: China, Guangzhou City, Baiyun Mountain, in
1,000 replicates. Branches corresponding to partitions broad-leaved forests, at 113°17′E, 23°10′N, alt. 269 m, 11.
reproduced in less than 50% bootstrap replicates are col- V. 2012, X. L. He, P. Li, C. Y. Deng and C. S. Qiu (GDGM
lapsed. All positions containing gaps and missing data were 31939).
used from the dataset. Etymology: macrocarpa (Lat.): referring to the large
basidiomata of the new species.
Pileus 150–240 mm broad, globose when young, convex
Results to planar with age, dry, pale orange (5A3–7A3) when
young, dirty white to brownish orange or light brown
Molecular phylogeny (6C3–6D5) with age, covered with numerous pale dirty
brown to brownish orange (7A3–7C5) pyramidal warts (2–
The nrLSU dataset included 33 taxa with 988 nucleotide sites, 3 mm in length); margin smooth, appendiculate. Lamellae
and Limacella glioderma (Fr.) Maire served as the outgroup. free, pale yellow to light yellow (4A3–4), crowded, 7–11
The ITS dataset included 10 taxa with 708 nucleotide sites, per cm, thick, 12–18 mm wide; lamellulae attenuate, plen-
and A. altipes Zhu L. Yang, M. Weiss & Oberw. in subgenus tiful, evenly distributed. Stipe 180–240 mm long, 20–45
Amanita served as the outgroup. Both datasets are available at mm thick at apex, subclavate, enlarged downwards to ven-
www.treebase (http://purl.org/phylo/treebase/phylows/study/ tricose bulbous base (57–62 mm diam.), generally pale
TB2:S13861) yellow to pale orange (4A3–5A3), but usually with pinkish
A phylogenetic tree constructed using LSU sequences flush to pinkish white when young, solid, covered with pale
(Fig. 1) revealed that the Amanita sequences were distribut- red (7A3) to orange white (5A2) squamules; ventricose
ed across seven monophyletic clades supported by high basal bulb, 57–62 mm diam., pale yellow to pale orange
clade credibility values, which corresponded to the different (4A3–5A3), volval remnants as pale orange floccose
sections described for the genus. Amanita afrospinosa patches or warts on the upper half of bulb, no distinct
Pegler & Shah-Smith, A. japonica Hongo ex Bas, A. boundary between stipe and bulb. Annulus persistent, pen-
kotohiraensis Nagas. & Mitani, A. longipes Bas ex Tulloss dant from attachment 60–80 mm below apex of stipe,
& D.T. Jenkins, A. macrocarpa, A. oberwinklerana Zhu L. floccose-membranous, thick, white to yellowish white and
Yang & Yoshim. Doi, A. rhoadsii (Murrill) Murrill, and A. striate above, orange white (5A2) to pale brown (6D5) and
virgineoides Bas formed a single clade within the section verrucose below. Context of pileus white, 20–30 mm thick
Lepidella of the subgenus Lepidella with a 91% bootstrap in center, becoming yellowish to light yellow (4A3–5) when
value. Therefore, we propose that the new species, A. exposed. Strong unpleasant odour.
macrocarpa, should be in section Lepidella along with A. Lamellar trama bilateral. Mediostratum 50–75 μm wide,
oberwinklerana which, as suggested by Zhang et al. (2010), composed of abundant ellipsoidal to cylindrical cells (30–
is also a member of this section. 50×10–15 μm), mixed with branching, interwoven, fila-
To further investigate the relationship between the new mentous hyphae, 3–8 μm wide; lateral stratum made up of
species and other species in the section Lepidella, an anal- abundant clavate to subellipsoidal inflated cells (30–70×
ysis of ITS sequences was undertaken. This revealed A. 10–15 μm), mixed with abundant, filamentous hyphae, 3–
Mycol Progress (2014) 13:211–217 213

Table 1 Taxa included in analyses

Taxon Voucher Locality GenBank accession no.

LSU ITS

Amanita afrospinosa Pegler & Shah-Smith RET 347-1 Zimbabwe HQ539666 –

A. altipes Zhu L. Yang, M. Weiss & Oberw. HKAS 30669 China AY436487 –
A. altipes HKAS 36609 China – AY436445
A. avellaneosquamosa (S. Imai) S. Imai HKAS 29500 China AF024441 –
A. chepangiana Tulloss & Bhandary HKAS 25772 China AF024445 –
A. citrina (Schaeff.) Pers. HKAS 31449 Germany AF024446 –
A. curtipes E.-J. Gilbert AH31718 Spain EF653961 –
A. eijii Zhu L. Yang A9 China – FJ441039
A. excelsa (Fr.) Bertill. Yang D 97 Germany AF024449 –
A. exitialis Zhu L. Yang & T. H. Li HKAS 38162 China AY436492 –
A. flavipes S. Imai HKAS 32505 China AF024451 –
A. fritillaria Sacc. HKAS 29511 China AF024452 –
A. fuliginea Hongo HKAS 32521 China AF024454 –
A. gilbertii Beauseign. RET 97016 – – AY325838
A. hemibapha var. ochracea Zhu L. Yang HKAS 29522 China AF024458 –
A. japonica Hongo ex Bas LEM960167 Japan – AB015684
A. japonica HMAS 59778 China AF024460 –
A. kotohiraensis Nagas. & Mitani MHHNU 6998 China FJ011681 FJ176722
A. kotohiraensis MHHNU 7112 China – FJ176723
A. longipes Bas ex Tulloss & D.T. Jenkins RET 360-1 USA HQ539704 –
A. longipes TENN61554 USA – FJ596834
A. longistriata S. Imai Bas 9040 Japan AF024462 –
A. macrocarpa W. Q. Deng, T.H. Li & Zhu L. Yang GDGM 31939 China KC408378a KC408379a
A. macrocarpa GDGM 32531 China – KC408380a
A. muscaria (L.) Lam. B.S-I Switzerland AF097368 –
A. nivalis Grev. Watling 17489 United Kingdom AF024466 –
A. oberwinklerana Zhu L. Yang & Yoshim. Doi MHHNU 7113 China FJ011683 FJ176725
A. oberwinklerana MHHNU 6826 China – FJ176727
A. phalloides (Vaill. ex Fr.) Link TUB 011556 Germany DQ071721 –
A. ponderosa Malençon & R. Heim AH19752 Spain EF653958 –
A. pseudovaginata Hongo HKAS 29524 China AF024472 –
A. rhoadsii (Murrill) Murrill DD97/13 USA AF097391 –
A. rubrovolvata S. Imai HKAS 32511 China AF024473 –
A. sinensis Zhu L. Yang HKAS 25761 China AF024474 –
A. solitaria (Bull.) Fr. HKAS 31459 Germany – AY436475
A. subjunquillea S. Imai HKAS 24169 China AF024479 –
A. umbrinolutea (Secr. ex Gillet) Bataille HKAS 31451 Germany AF024481 –
A. vaginata (Bull.) Lam. RV4Aug96 USA AF097375 –
A. virgineoides Bas LEM960205 Japan – AB015686
A. virgineoides HKAS 18394 China AF024484 –
A. virgineoides A2 China – FJ441032
A. volvata (Peck) Lloyd RV97/24 USA AF097388 –
A. yuaniana Zhu L. Yang HKAS 29516 China AF024488 –
Limacella glioderma (Fr.) Maire HKAS 31561 Germany AF024489 –
a
Sequences obtained in this study. Sequences for the other Amanita taxa and Limacella glioderma were from the literature (Weiß et al. 1998; Oda et
al. 1999; Zhang et al. 2004, 2010; Moreno et al. 2008)
214 Mycol Progress (2014) 13:211–217

Fig. 1 Phylogenetic tree of

Amanita macrocarpa and
related species within the genus
Amanita based on nuc-LSU
rDNA sequences generated by
Maximum Likelihood method
in MEGA5. Limacella
glioderma was chosen as the
outgroup. Bootstrap values (>
50%) are shown above or
beneath individual branches

8 μm wide; septa often with clamps. Subhymenium 20–40 thin-walled, colorless to subcolorless hyphae. Volval rem-
μm thick, with 2–3 layers of subglobose, ovoid to irregu- nants on pileus composed of±vertically arranged elements:
larly shaped cells, 9–30×7–18 μm. Basidia 30–45×6–9 μm, inflated cells abundant to very abundant, globose to
clavate, 4-spored; sterigmata 3–5 μm long; basal septa often subglobose or ovoid (15–45×15–40 μm), occasionally el-
with clamps. Basidiospores (100/5/5) (6–)7–9×(4–)5–6 μm lipsoidal (20–45×10–25 μm), often in chains of 2–4 and
(Q=1.3–1.7, Qm =1.5±0.05), ellipsoidal, amyloid, colorless, terminal, thin-walled to slightly thick-walled (up to 0.5 μm),
hyaline, thin-walled, smooth. Lamellar edge sterile, com- almost colorless, occasionally with yellowish vacuolar pig-
posed of subglobose or broadly clavate (15–25×9–15 μm) ments; filamentous hyphae scattered to abundant, 3–
to elongated clavate, thin-walled, colorless, hyaline cells 10(−15) μm wide, colorless, occasionally with yellowish
(25–30×5–8 μm); filamentous hyphae abundant, 3–7 μm vacuolar pigments, clamped; vascular hyphae rare; base of
wide, thin-walled, hyaline or with yellowish vacuolar pig- volval remnants with dominant hyphae; top of volval rem-
ments. Pileipellis 160–300 μm thick, two-layered; upper nants with dominant inflated cells. Volval remnants on base
layer (120–200 μm thick) gelatinized, composed of of stipe composed of more or less irregularly arranged
subradially arranged, (4–)8–16(−25) μm wide, filamentous elements: inflated cells abundant, subglobose or ovoid
to cylindrical hyphae with yellowish vacuolar pigments; (15–35×10–26 μm), occasionally ellipsoidal to clavate
lower layer (50–80 μm thick) non-gelatinized, composed (20–90×10–25 μm), often in chains of 2–4 and terminal,
of radially and compactly arranged, 6–15(−22) μm wide, thin-walled, colorless, hyaline; filamentous hyphae fairly
Mycol Progress (2014) 13:211–217 215

Fig. 2 Phylogenetic tree of Amanita macrocarpa and related species altipes was chosen as the outgroup. Bootstrap values (> 50%) are
within Amanita sect. Lepidella based on nuc-ITS rDNA sequences shown above or beneath individual branches
generated by Maximum Likelihood method in MEGA5. Amanita

abundant, 2–10(−18) μm wide, colorless, infrequently Habitat: Scattered on soil, in subtropical broad-leaved
clamped; vascular hyphae rare. Stipe trama composed of forests dominated by Castanopsis fissa, at an altitude of
longitudinally arranged, long clavate, terminal cells 200–300 m.
(80–)100–200×10–20 μm; filamentous hyphae scattered to Collections examined: CHINA, Guangdong Province:
fairly abundant, 2–8(−15) μm wide; vascular hyphae 6–16 Guangzhou City, Baiyun Mountain, in broad-leaved forests,
μm wide, rare. Annulus composed of loosely and subradially at 113°17′E, 23°10′N, alt. 269 m, 12 Jun 2009, X. L. Chen
arranged, 3–9 μm wide, thin-walled, filamentous hyphae; and H. Huang (GDGM 27129); the same location, 18 May
inflated cells scattered to locally (fairly) abundant, subglobose 2012, W. Q. Deng and S. H. Zhou (GDGM 31069); the
to broadly clavate (16–40×12–30 μm), thin-walled, colorless same location, 22 May 2012, Z. D. Xiao and S. H. Zhou
or yellowish; vascular hyphae 6–18 μm wide, rare. (GDGM 31932); Guangzhou City, Tianluhu Forest Park, in

Fig. 3 Basidiomata of Amanita

macrocarpa. a, b (GDGM
27129), c, d (GDGM 31939,
holotype) (bars 5 cm)
216 Mycol Progress (2014) 13:211–217

Fig. 4 Microscopic structures

of Amanita macrocarpa
(GDGM 31939, drawn by Zhu
L. Yang). a Basidia and
subhymenium, b basidiospores,
c longitudinal section of volval
remnant from pileus

broad-leaved forests, 06 Sep 2012, M. Zhang and S. H. However, A. japonica differs from A. macrocarpa in pro-
Zhou (GDGM 32531). ducing a distinctly dark gray to pale buffy gray pileus and
longer basidiospores (8.5–10.5×5.5–7 μm) (Bas 1969;
Yang 1997, 2005). Amanita rhoadsii differs from A.
Discussion macrocarpa in producing smaller and white basidiomata
with white and unchanging context, and much longer basid-
Amanita macrocarpa is characterized by a large pileus with iospores [10.5–13.5(−15.5)×3.5–4.5 μm] with a larger Q
numerous brownish orange to pale dirty brown pyramidal (Bas 1969).
warts on the dirty white or brownish pileal surface, a Five species assigned to section Lepidella from eastern and
subclavate stipe with no distinct boundary between the stipe southeastern Asia, namely A. hongoi Bas, A. perpasta Corner
and the bulb, the common presence of clamps, and amyloid & Bas, A. timida Corner & Bas, A. virginea Massee, and A.
ellipsoidal basidiospores. Initial systematic placement in the yenii Zhu L. Yang & C. M. Chen, resemble A. macrocarpa in
Amanita subgenus Lepidella section Lepidella (Bas 1969; having pyramidal warts on the pileus. Amanita hongoi, orig-
Yang 1997, 2005) was confirmed by molecular analyses inally described from Japan, also has similarly colored
(Figs. 1 and 2). basidiomata but differs from A. macrocarpa in having smaller
Amanita macrocarpa may be keyed out in the Amanita pilei white to cream gills, the absence of clamps, and globose
subsect. Solitariae Bas stirps Virgineoides (Bas 1969) but to broadly ellipsoidal basidiospores (7.5–9.5×6.5–8.0 μm)
differs from other species of the stirps. Among the species in with a lower Q value (Bas 1969; Yang 2005). Amanita
stirps Virgineoides, the Asian A. virgineoides exhibits closest perpasta, originally described from Singapore, can be distin-
similarity to the new species, but differs from A. macrocarpa guished from A. macrocarpa by its whitish to pallid cream or
in having smaller and white basidiomata with white and pale brown pileus, cream-white gills, the absence of clamps,
unchanging context, and larger basidiospores (8.0–10.5× and globose to subglobose basidiospores [7–8.5(−10.5)×6.5–
6.0–7.5 μm) with a lower Q value (Bas 1969; Yang 1997, 8(−9.5) μm] (Corner and Bas 1962). Amanita timida, also
2005). Amanita gracilior Bas & Honrubia, originally de- described originally from Singapore, differs from A.
scribed from Spain and France, differs from A. macrocarpa macrocarpa in having a pale pink to flesh-colored pileus,
in having smaller basidiomata, white stem and gills, longer fleshy white then cream gills, very slowly becoming pinkish
basidiospores [(9.5–)10–12.5(−13.4)×(5.4–)5.5–6.3 (−7.8)], in places on breaking, and broader basidiospores [(6.5–)7–
and its association with oak (Quercus) and pine (Pinus) in 9(−10)×5.5– 7(–8.5) μm]. Furthermore, the stipe often has a
autumn (Bas and Honrubia 1982). Amanita miculifera Bas & strobiliform base (Corner and Bas 1962; Bas 1969). Amanita
Hatan., originally described from Japan, differs from A. virginea, originally described from Singapore, differs from A.
macrocarpa in having a gray pileus and stipe, white gills, macrocarpa by its white pileus, warts and gills, and
and larger basidiospores [(9.0–)10.0–12.5(−13.5)×(6.0–)6.5– subglobose to broadly ellipsoidal basidiospores (7–9×6.5–
8.0 μm] (Bas and Hatanaka 1984). 7.5 μm) (Bas 1969). Amanita yenii, originally described from
Phylogenetically, the designation of A. macrocarpa as a China, has whitish warts on the pileus, yellowish lamellae,
distinct species was supported by the LSU and ITS se- and ellipsoidal basidiospores that are similar to those of A.
quences, and the ITS and LSU datasets revealed a close macrocarpa. However, differing features include a smaller
relationship with A. japonica and A. rhoadsii, respectively. and often white fruit body, the absence of clamps, less
Mycol Progress (2014) 13:211–217 217

vascular hyphae, unchanging flesh when exposed, and an Molecular phylogenetic analysis shows that Amanita ponderosa
and A. curtipes are distinct species. Mycol Progr 7:41–47
indistinct smell (Yang and Chen 2003; Yang 2005).
Oda T, Tanaka C, Tsuda M (1999) Molecular phylogeny of Japanese
Amanita species based on nucleotide sequences of the internal
Acknowledgments This research was supported by the National transcribed spacer region of nuclear ribosomal DNA.
Natural Science Foundation of China (Project Nos. 31101592, Mycoscience 40:57–64
31070024 and 31170026). We thank Ya-Heng Shen and Dr. Lei Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011)
Wang, Guangdong Institute of Microbiology, for their valuable assis- MEGA5: Molecular evolutionary genetics analysis using maxi-
tance, and Dr John Buswell, Shanghai Academy of Agricultural mum likelihood, evolutionary distance, and maximum parsimony
Sciences, for linguistic revision of the manuscript. methods. Mol Biol Evol 28(10):2731–2739. doi:10.1093/molbev/
msr121
Tompson JD, Gibson TJ, Plewnlak F, Jianmougin F, Higgins DG
(1997) The Clustal X windows interfaces: flexible strategies for
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