Bull. Natn . Sci. Mus., Tokyo, Ser. B, 25(3), pp. I07- 130, September 22, 1999

A Contribution to the Knowledge of Amanita(Amanitaceae, Agaricales) in Japan
Zhu-Liang Yang 1 and Yoshimichi Doi 2
1

Kunming Institute of Botany, th e Chinese Academy of Sciences,
Kunming 650204, P. R. China
2
Department of Botany, National Science Museum,4- 1- 1 Amak ubo. Tsukuba, lbarak i, 305- 0005 Japa n

Abstract Specimens of the genus Amanita collected from Japan and deposited inth e herbar ium of th e Department of Botany, Nationa l Science Museum, Tsukuba,lbara ki , Japan (TNS) were studied . Eighteen taxa, includ ing two species new to science and two taxa new to Japan are reported here. Brief notes on a few type specimens of the Amanitae described by T. Hongo are also included in the present account. The new spec ies are Amanita orientogemmara and A. oberwinklerana, whileth e new records are A. manginiana sensu Chiu and A. subjunquillea var. alba .
Amanita sphaerobulbosa is treated as a distinct species, independent of A. abrupta .
Key words:

Amanita , A. orienrogemmara , A. oberwinklerana , Japan, East Asia

The Amanitae in Japan are relatively well known thanks to the researches ofseveral active mycologists (Bas, 1969; Bas & Hatanaka, 1984; Hongo, 1953- 1983;
lmai , 1933 , 193 8, 1939; Imazeki & Hongo, 1970, 1975, 198 7; Imazeki & Toki , 1955 ;
lmazeki et al., 1988; !to, I 959; Matsuda & Hongo 1955, 1956; Nagasawa & Hon go,
1984, 1985 ; Tulloss et al., 1992 a, and many others) . It is interested to note that T.
Hongo alone has desc ribed thirteen new taxa of Amanita from Japan (Doi , 1991 ),many of wh ich are a lso found in China (Mao, 1991 ; Yang, 1994, 1997). During a cooperative research between China and Japan , we studied some specimens of the
Amanitae collected from Japan by Y. Doi and hi s colleagues. In order to understandbetter the concept of some spec ies described by T. Hongo, his type collections werealso exam ined. Having studied type material as well as newly collected and welldried specimens, we add some new data to the Japanese Amanita-flora. Color codesof the form e.g., 3 84 are from Kornerup & Wanscher ( 198 1); col or names with first
letters cap itali zed (e.g., Cream Co lor) are from Ridgway ( 1912).
Subgenus Amanita Section Amanita
I. Amanita orientogemmata Z. L. Yang & Y. Doi , sp. nov.
Figs. 1-4
Name misapplied to the present spec ies: Amanita gemmata sensu Nagasawa &Hongo in Mem. Natn. Sci. M us. , Tokyo 18: 73 ( 1985); non Amanita gemma/a

108

Zhu-Liang Yang and Yoshimichi Doi

003

Figs. 1- 3. Amanita orien togemmata(TNS-F-198537 , Holotype). I. Basidiocarp ; 2. Hymenium and subhymenium ; 3. Basidiospores.

I~

20 /1111

I

Fig. 4. Longitudinal section of volval remnant from pileusof Amanita orientogemmata (TNS-F-198537, Holotype).

(Fr.) Bertillon in DeChambre, Diet. Encycl. Sci . Medic. 1(3): 496 (1866).
Selected Illustr.: Nagasawa & Hongo in Mem. Natn. Sci . Mus., Tokyo 18: 74, fig.
1: 87, fig. 9 (1985).
Pileus 5- 10 cm latus, initio semiglobosus, deinde convexus vel plano-convexus,
giseo-flavus vel pallide flavus, reliquiis valvae coactis, flocculosis, applanatis,
tenuibus, sordide albidis ornatus, margine brevite striata, non appendiculata. Lamellae liberae, albae vel cremeae, confertae, lamellulis truncatis. Stipes 5- 11 X 1- 1.5 cm,
sursum attenuatus, a/bus vel albidus, annulatus. Bulbus volvatus, subglobosus, sub-

Amanitae in Japan109

radicatus, 2- 3 cm latus. Valva breviter limbata. Annulus membranaceus, a/bus, inferior. Caro alba. Basidia 4-sporigera. Sporae (7.5- ) 8.0- /0.0 (- II.O) X (6.0- ) 6.5- 7.5(- 8.0) J..lm, lato-ellipsoideae vel ellipsoideae, non-amyloideae. Fibulae praesentes.
Holotypus: Ochidani, Tottori City, Japan, /2. IX 1979, E. & F Nagasawa (TNS-F19853 7).
Etymology: "orientogemmata" is named because of the similarity to Amanita
gemmata from Europe.
Pileus 5- 10 cm in diameter, at first hemispherical, then becoming convex topiano-convex, occasionally slightly depressed at the center, moderate greyish yellow(3-48- C4-5) to light yellow (Apricot Yellow, Celonial Buff, Straw Yellow, 3A43A5), somewhat darker on disc, viscid when moist; volval remnants as felty, floccose
patches or small subconical warts, 1-4 mm wide, up to 1.5 mm high (thick), whitish,
becoming dirty white , yellowish or greyish when dried, randomly arranged, frequently densely placed over disc; margin short tuberculate-striate (0. 15- 0.25 R; R=radius) , non-appendiculate; trama white. Lamellae free, whitish to pale creamy, crowded, with edge finely fimbriate and floccose; lamellulae truncate, plentiful, in 2- 3
ranks. Stipe 5- 11 X 1- 1.5 cm, usually tapering upward, with apex slightly expanded,
white to whitish , with cracked to appressed scaly belts above annulus and fibrillose
below it or entirely more or less fibrillose ; context white, loosely stuffed to hollow;
basal bulb ovate to subglobose, subradicating, 2- 3 cm wide, 3-4 cm high, white topallid; volva short and appressed-limbate or as floccose patches or warts near apex ofbulb, white to yellowish. Annulus present or torn from stipe during expansion of
pileus, usually inferior when present, rarely median, white with occasional yellowish
tinge. Odor and taste indistinct. Color of spore print not known.
Lamella trama bilateral. Mediostratum 30- 50 J..lm wide , consisting of long ellipsoid to fusiform cells, I 0- 30 J..lm wide, mixed with abundant, branching hyphae,
2- 5 (- 8) J..lm wide; vascular hyphae rare. Lateral stratum consisting of long ellipsoid
to fusiform cells, I 0- 20 J..lm wide, mixed with abundant hyphae, 3- 7 J..l.m wide, diverging at an angle of 30°-60° to the central stratum; septa often with clamps. Subhymenium 30-45 J..lm thick, with 2- 3 (-4) layers of subglobose, ovoid to short ellipsoid, sometimes dolioform cells, I0- 25 X 10- 18 ,urn. Basidia 45- 60 X 10- 14 J..lm ,clavate, 4-spored; sterigmata 4- 6 ,urn long; basal septa often with clamps. Basidiospores [I 00/4/3] (=I 00 spores from 4 basidiocarps of 3 collections measured in
5% KOH solution; Q is used to mean "length/width ratio" of a spore; Q means average Q of all specimens:tsample standard deviation): (7.5- ) 8.0-10.0 (- ll.O) X(6.0- )
6.5- 7.5 (- 8.0)f..lm [Q = (l.07- ) 1.15- 1.46 (- 1.59), Q = 1.30:t0.01], broadly ellipsoid
to ellipsoid, rarely subglobose, inamyloid, colorless, hyaline, thin-walled, smooth,
apiculus small. Lamella edge as sterile, somewhat gelatinized strip, 100- 200 ,urnwide in side view, made up of clavate to broadly clavate (45- 60 X 15- 20 J..lm), to ovoid
to subglobose (25-45 X20- 35 J..lm) or ellipsoid (30-40 X 15- 25 ,urn) cells, 2-4 inchains, thin-walled, colorless or sometimes with yellowish contents, surface often

110

Zhu-Liang Yang and Yoshimi chi Doi

with fine granular substance, mixed with scattered to fairly abundant, filamentous hyphae, 2- 5 pm wide, often clamped; elements embedded in yellowish amorphous matrix which gradually disappears in KOH. Pileipellis 80- 150 pm thick ; upper layer(60-90 J.lm thick) strongly gelatinized, made up of subradially arranged to somewhat
interwoven hyphae , 1- 5 pm wide, colorless to subcolorless, often with fine granular
incrustations; lower layer (40- 60 pm thick) made up of radially, subradially and compactly arranged hyphae, 3- 8 J.lm wide, subcolorless or with yellowish to brownish
vacuolar pigment; vascular hyphae rare, 2- 7 J.lm wide . Volval remnants on pi le uscomprising±vertically, sometimes irregularly arranged elements: inflated cells fairly
abundant to abundant, ellipsoid to subglobose cells, 25- 90 X 25-40 pm, often in shortchains, thin-walled, colorless, or sometimes with brownish content; fairly abundant tolocally abundant, filamentous hyphae, 3- 5 (- 8) f.1m wide, thin-walled, colorless orsometimes with yellowish to brownish content; septa often with clamps; vascular hyphae rare or locally abundant, 2- 10 (- 15)J.1m wide. Volval remnants on the stipe baseand the upper bulb similar to those on the pileus, but elements irregularly arranged.
Stipe trama dominantly consisting of longitudinally arranged, long clavate, terminal
cells, 200- 350 X 18-40 pm, mixed with 2- 8 pm wide hyphae scattered in interior,
fairly abundant on stipe surface; vascular hyphae rare to locally fairly abundant,
2- 8 J.lm wide. Annulus of abundant to locally very abundant, branching and interwoven hyphae, 3- 5 (- 8)f.1m wide, mixed with fairly abundant to abundant, subglobose
to ellipsoid cells, 35- 60X20- 35 J.lm.
Specimens examined: Japan: Ochidani, Tottori City, Tottori Pref. , 12. IX. 1979,
E. & F. Nagasawa s.n. (=sine numero) (TNS-F-198537 , Holotype!); ditto, 8.1X.
1979, E. & F. Nagasawa s.n. (TNS-F-198540) ; ditto, 9. IX. 1979, E. & F. Nagasawa
s.n. (TNS-F-198546).
Habitat and distribution: Solitary or gregarious on ground in forest of AbiesCastanopsis. Only known from Japan .
Observations: In the above description the color and size of the basidiocarps
are mainly based on the descriptions of Nagasawa and Hongo ( 1985). Other data arefrom observations on the collections by the present authors.
Amanita orientogemmata is characterized by its greyish yellow to light yellow
pileus with whitish , felted volval patches or warts, inferior and fugacious annulus,
limbate volva, inamyloid, broadly ellipsoid to ellipsoid spores, and presence ofclamps. It differs from A. gemmata (Fr.) Bertillon in the rather common presence ofclamps, smaller and proportionately narrower spores, and different structure of volval
remnants on pileus. The volval remnants on pileus of Amanita gemmata from Europe[Netherlands: Friesland, 23. X. 1994, C . Bas 8942 (L); Schoorl , 26. XI. 1994, C. Bas
8949 (L)] dominantly consist of irregularly to subradially arranged, filamentous
hypae, mixed with scattered to locally fairly abundant inflated cells (Yang, 1997: 51) .
The spores of A. gemmata from the two European collections listed above are(120/6/2] (9.0- ) 9.5- 12.0 (13.0) X (6.0- ) 6.5- 8.5 (- 9.0)J.1m (Q=(l.l7- ) 1.25- 1.62

Amanitae in JapanIll(- 1.77), Q= 1.42 ±0 .13].
Nagasawa & Hongo ( 1985) indicated that this species may be closely related to
Amanita pantherina var. ve/atipes (Atk.) Jenkins (=A. velatipes Atk.) from North
America. However, A. velatipes has a persistent annulus, an ocreate valva on thebulb, and narrower spores (Jenkins, 1977, 1982, 1986; Tulloss et al., 1995).
Species in the section Amanita having yellow to yellowish pilei and exhibitingsome morphological sim ilarities to A. orientogemmata are A. pantherina var. lutea
C hiu from China., A. elata (Mass.) Corner & Bas from Southeast Asia, A. crenulata
Peck from North America, A. xylinovolva Tulloss, Overbo & Hailing from Colombia,
and A. diemii Singer from Argentina. Amanita pantherina var. lutea differs from thepresent species by its bulbs with an ocreate collar, a superior annulus, and absence ofclamps (Yang, 1997). Amanita elata differs by its more detachable volval remnantson pileus, smaller, globose to subglobose spores, and absence of clamps (Corner &Bas, 1962). Amanita crenulata differs from this species in its sordid yellowish tan tosomewhat sordid pale tan pileus with pulverulent, champagne-colored volval remnants, superior annulus with few inflated cells, absence of clamps, and rounder spores(Tulloss, 1990). Amanita xylin ivolva has longer marginal striations on the pileus, noclamps, and wider spores (Tulloss et al. , 1992 b). Amanita diemii, occurring under
Nothofagus, differs from A. orientogemmata by its golden yellow to orange ochre yel low pileus becoming brown with age, lacking annulus, and rounder spores (Singer,
1954).
2. Amanita melleiceps Hongo in J. Jpn . Bot. 41: 165 ( 1966).
Fig. 5Basidia 32-40 X 10- 13 J..lm , clavate, 4-spored; basal septa without clamps. Basidiospores [50/211] (7.5- ) 8.5- 11.0 (- 12.0) X 6.0- 7.0 (- 8.0),um [Q=(l.25- ) 1.291.64 (- 1.83), Q = 1.4 7±0. 11 ], mostly ellipsoid, inamyloid, colorless, hyaline, thinwalled, smooth; apiculus sma ll. Volval remnants on pileus made up of ±verticallyarranged elements; inflated cells abundant to very abundant, subglobose to ovoid(25- 35X20- 25,um), to ellipsoid (30-45Xl5- 30,um), probably in short chains, thinwalled, colourless, hyaline to subhyaline; filamentous hyphae fairly abundant, 2- 5 ,umwide, branching, thin-walled, hyaline to subhyaline; septa without clamps; vascularhyphae rare.
Specimen examined: Japan: Senjo, Otsu, 22. VI. 1965, T. Hongo s.n. (Hongo
Herb. 3070, TNS-F-2372 79, Holotype).
Habitat and distribution: In forest of Pinus densif/.ora, mixed with young
Quercus serrata. Also known from South Korea and China (Kim, et al., 1993 ; Mao,
1991 ).
Observations: Having studied the holotype , we know that this species has noclamps and the volval remnants on pileus are made up of vertically arranged elements.

112

Zhu-Liang Yang and Yoshimichi Doi

00
00
001

10 fllll

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OOs

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o0010~mI()

Figs. 5- 6. Bas idiospores. 5. Amanita melleiceps(TNS-F-237279, Holotype); 6. Amanita pseudogemma/a (TNS-F-237280, Holotype) .

3. Amanita pseudogemmata Hongo in Travaux mycologiques dedies a R. Kuhner,
Bull. Soc. Linn. Lyon, no. spec.: 189 ( 1974).
Fig. 6
Basidiospores [301111] (7 .0- ) 7.5- 9.5 (- 10.0) X(6.0- ) 6.5- 7.5 (- 8.0),um [Q=(1.07- ) 1.1 - 1.28 (- 1.31 ), Q= 1.20±0.06], subglobose to broadly ellipsoid, inamyloid,
colorless, hyaline, thin-walled, smooth, apiculus small. Volval remnants on pileus
made up of ±vertical elements: inflated cells fairly abundant to abundant, subglobose
to ovoid to ellipsoid (50- 75 X25- 50 ,urn), thin-walled, hyaline or often with yellowish
vacuolar pigment; filamentous hyphae fairly abundant to scattered, 3- 5 ,urn wide; vascular hyphae plentiful, 2.5- 8 ,urn wide.
Specimen examined: Japan: Ginkakuji, Sakyo-Ku, Kyoto, 9. VIII. 1970, H.
Noro s.n. (Hongo Herb. 4201 , TNS-F-237280, Holotype).
Habitat and Distribution: In forest of Castanopsis cuspidata. Only known
from Japan.
Observations: The holotype is poorly rehydrating. Basal septa of basidia probably have no clamps. Vascular hyphae in the volval remnants on pileus are quite conspicuous.
4. Amanita cf. pantherina.
Figs. 7- 9
Pileus 5.5- 8.5 cm in diameter, convex to planate, yellowish-greyish, darker at
the center; volval remnants felt-like to subconical, 1-4 mm wide, up to 3 mm high,dirty white, randomly arranged, frequently denser at the center; margin short tuberculate-striate (0.1 - 0.2 R), often with remnants of annulus; trama white. Lamellae free ,white to whitish, crowded; lamellulae truncate, plentiful. Stipe 9- 11 X0.5- l cm,
slightly attenuate upward, white to whitish, lower part with somewhat reftexed scales;
basal bulb ovate to subglobose, ea. 1.5- 2.5 cm wide, 2- 3 cm high, white to pallid;
volva an ocreate ring. Annulus present but often torn from stipe during expansion of
pileus, and then hanging on margin of pileus. Odor and taste unknown. Color of
spore print unknown.
Basidia 45- 5 5 X 11 - 12 pm, clavate, 4-spored; sterigmata 3- 5 ,urn long; basal
septa often with clamps. Basidiospores [301111] (8.5- ) 9.0- ll.O X(6.5- ) 7.0- 8.5(- 9.0),um [Q=(l.lO- ) 1.19- 1.38 (- 1.40), Q= 1.29±0.07], broadly ellipsoid to ellipsoid, inamyloid, colorless, hyaline, thin-walled, smooth, apiculus small. Volval rem-

Amanitae in Japan113

Figs. 7- 8. Amanita cf. pantherina (TNS-F-182971 ). 7. Bas idiocarp; 8. Basidiospores.

9Fig. 9. Longitudinal section o f volval remnant from pileus of Amanita cf. pantherina(TNS -F-1 8297 1).

nants on pileus made up of ±vertically arranged elements: inflated cells abundant tovery abundant, subglobose, ovoid to short ellipsoid, 20--45 X l5- 35p.m, sometimes
long ellipsoid to subfusiform, 55- 60X20- 25 J.lm, single and terminal, or 2- 3 (--4) inchains, thin-walled, colorless, hyaline, sometimes with brownish vacuolar pigment;
fairly abundant to abundant hyphae, 2- 7 (- 10) J.lm wide, thin-walled, colorless, hyaline; septa often with clamps; many hyphae vascular, i.e. with yellowish-brownish,reflactive, vacuolar pigment (content).

114

Zhu-Liang Yang and Yoshimi chi Doi

Specimen examined: Japan: Tsukuba Botanical Garden , Tsukuba , lbaraki Pref. ,I. X. 1997, Y. Doi s.n. (TNS-F-182971).
Habitat and Distribution: On ground under trees of Castanea.
Observations: This species probably has been regarded as Amanita pantherina(DC.: Fr.) Krombh. in the past. It differs, however, from the latter in , among otherfeatures , its presence of clamps. lt is probably an undescribed taxon. Since we do notunderstand this species well, it will need to be studied further when more collections
are available.
Subgenus Amanita Section Caesareae Singer ex Singer
5. Amanita longistriata lmai in 1. Facul. Agr. Hokkaido Imp. Univ. 43: 11 ( 1938).
Fig. 10
Pileus 3- 9 cm in diameter, at first hemispherical , then convex to piano-convex ,with or without small umbo, slightly greyish buff, sometimes with very slightly pinkish tinge, greyish, sometimes greyish brown to si ightly brownish at disc, subviscid,
glabrous and usually without volval remnants; margin usually long sulcate [0 .30.5 R] , occasionally short striate [0.2- 0.25 R], non-appendiculate; context white.
Lamellae free, or even remote from stipe, (sub) distant, rather broad (up to !!mmbroad), already when very young beautifully pale but clear pink, later paler pink, withedge minutely white flocculose, subdenticulate; lamellulae obliquely truncate. Stipe
9- 18 X 0.5- 1.5 cm , subcylilndrical or si ightly tapering upward, at first pinkish white ,later white, with proportionately broad central cylinder, base lacking bulb; volva saccate, 2- 3 (-4) X 1- 2 cm, ea. 0.5- 1.5mm thick, membranous, white to whitish , withlimbus internus placed on inside of volval limb, slightly above point of juncture ofstipe and limb. Annulus present, superior to almost median , membranous, thin,
whitish . Taste mild; Odor indistinct. Color of spore print not known .
Subhymenium 30-40 pm thick , with 2- 3 layers of subglobose, ovoid to ellipsoid cells, I 0- 23 X 8- 15 pm . Basidia 34-48 X 11 - 13 pm , clavate, 4-spored; sterigmata 3- 5 pm long ; basal septa often with clamps. Basidiospores [175/8/5] (8.0- ) 9.012.0 (- 13 .5) X(7.5- ) 8.0- 10.0 (- li.O)pm [Q=(I.02- ) 1.06- 1.33 (- 1.53), Q = 1.19 ::!:::
0.08] , dominantly broadly ellipsoid, sometimes ellipsoid or subglobose, inamyloid,
colorless, hyaline, thin-walled, smooth, apiculus small. Volval remnants on stipe
base made up of irregularly to somewhat longitudinally arranged elements: filamentous hyphae dominant, 2- 7 (- 12)pm wide, hyaline, thin-walled, branching, interwoven, with clamps; inflated cells scarce to scattered to locally fairly abundant, subglobose to ovoid, 50- 70 X 35- 50 pm, sometimes ellipsoid to fusiform , 50-120 X 2050pm , colourless hyaline, thin-walled ; vascular hyphae rare. Outer surface of volva
consisting of ::!::: irregularly arranged filamentous hyphae, 2- 5 (- 7) pm wide , hyaline,
thin-walled ; inflated cells rare. Inner surface of volva gelatinized, with filamentous
hyphae, 1- 3 pm wide. Volval remnants on pileus not seen. Annulus dominantly con-

115

Amanitae in Japan

Figs. 10- 11. Basidiospores. 10. Amanita long istriata (C. Bas 9040, in L);
11. Amanita pseudovaginata (TNS-F237282, Ho1otype)si sting of subradially to irregularly arranged, filamentous hyphae, 2~6 J..Lm wide,
mixed with usually rare but locally abundant, clavate to broadly clavate, inflated cells,
20~ 30 X 8~ 12 J..l.m, colorless, hyaline, thin-walled; clamps very common; vascular hyphae rare.
Specimens examined: Japan: Along hiking course, left-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 16. VII. 1997, Y. Doi & K.
Miyazaki s.n. (TNS-F-182475 ; ditto, TNS-F- 182478); Along hiking course, left-side
ridge of Kurosawa-gawa River, Kurosawa 3-Chome, Oume City, Tokyo, 5. X. 1997, Y.
Doi s.n. (TNS-F-237593); In garden of Fukiage-Gosho Imperial Palace; Chiyoda-ku,
Tokyo, 12. VII. 1995, T. Tobishima s.n. (TNS-F-180370); Mt. Fuji , Yoshida-guchi ,Yamanashi Pref. , 6. IX. 1983 , C. Bas 9040 (L).
Habitat and distribution: Solitary or gregarious on ground in forests (e.g., ofAbies). Also known from South Korea, and China (Kim et al. , 1993; Mao, 199 I;
Yang, I 997).
Observations: The colour of the fruitbody is mainly based on the field notes of
C. Bas 9040. This species is somewhat similar to A. spreta (Peck) Sacc., originallydescribed from North America. The latter has, however, considerably shorter marginal striations, smaller volva on stipe base, and significantly narrower spores. Amanita
longistriata is close to A. incarnatifolia Z . L.Yang (a new name for A. rhodophyllalmazeki & Toki , see Yang I 997: 52). lt differs from the latter in its somewhat larger
basidiocarps, longer striations on the pileus margin, and wider basidiospores (Yang,1997).
Subgenus Amanita Section Vaginatae (Fr.) Quel.
6. Amanita pseudovaginata Hongo in M em. Shiga. Univ., Nat. Sci., 33: I 08 (I 983).
Fig. I I
Basidiospores [50/211] (9 . 5~) 10 . 5~13 . 0 X 8.0~9.0 (~9.5)J..Lm [Q=(l.20~) 1.25 ~1.51 (~ 1.54) , Q= 1.37 : :1: : 0.08] , broadly ellipsoid to ellipsoid, inamyloid, colorless,hyaline, thin-walled, smooth , apiculus small. Volval remnants on pileus made up of: :1: : irregularly or sometimes subvertically arranged elements: inflated cells very abun-

116

Zhu-Liang Yang and Yoshimichi Doi

dant, subglobose to ovoid to ellipsoid (20- 50 X 15--40 ,urn), thin-walled, hyaline oroften with brownish vacuolar pigment, single and terminal or in short chains; filamentous hyphae fairly abundant to locally abundant, 2- 7 (- 10),um wide, often withbrown vacuolar pigment; vascular hyphae rare. On the outer surface of the volval
remnants there are more abundant±radially arranged, filamentous hyphae.
Specimen examined: Japan: Along road-cuts, Ikeno-o, Uji-City, Aug. 4, 1981 ,T Hongos.n. (Hongo Herb. 6134, TNS-F-237282, Holotype).
Habitat and distribution: In Pinus-Quercus forest. Also known from China(Yang, 1997).
Observations: The holotype specimen is poorly rehydrating. From our observations of bases of basidia, their basal septa probably have no clamps.
Subgenus Lepidella Section Amidella (E. J. Gilb.) Konr. & Maubl.
7. Amanita avellaneosquamosa (lmai) Imai in lto, Mycol. Fl. Jpn. 2(5) : 250 (1959).
Pileus ea. 5 cm in diameter, convex to piano-convex to planate, dirty white tobrownish, subviscid, occasionally with whitish volval remnants; margin short striate(0.15- 0.2 R). Lamellae free , crowded; lamellulae 0- 2, truncate.
Basidiospores [20/1/1] 8.5- 10.5 (- 11.0) X(4.5- ) 5.0- 5.5 (- 6.0),um [Q=(1.55- )1.66- 2.09 (-2.20), Q= 1.87 ±0. 16], long elongate to subcylindrical , amyloid, colorless, hyaline, thin-walled, smooth, apiculus small.
Specimen examined: Japan: Along right-side ridge of Kurosawa-gawa River,
Kurosawa 3-Chome, Oume City, Tokyo, 25 . IX. 1997, Y. Doi s.n. (TNS-F-237935).
Habitat and distribution: Solitary on ground in Quercus-forests. Also known
from China (Yang, 1997).
Observations: Amanita avellaneosquamosa was usually regarded as a synonymof A. volvata (Peck) Lloyd, originally named and described for a North Americanmushroom. Recent studies show that they should be treated as distinct taxa (Yang,
1997; Weil3 et al., 1998).
Subgenus Lepidella Section Lepidella
8. Amanita albojlavescens Hongo in Mem. Shiga Univ. , Nat. Sci. 20: 50 (1970).
Fig. 12Basidia 36--40 X 8- 11 ,urn, dominantly 4-spored, sometimes 2-spored. Basidiospores [30/1/1] 8.0- 11.0 (- 13.0)X(4.0- ) 4.5- 5.5 (- 6.5),um [Q=(1.63- ) 1.67- 2.09(- 2.12), Q= 1.88±0.16], elongate, colorless, hyaline, thin-walled, smooth, apiculus
small.
Specimen examined: Japan: Do, 6tsu, Shiga Pref. , 18. IX. 1969, T. Hongo s.n.
(Hongo Herb. 4014, TNS-F-237274, Holotype).
Habitat and Distribution: In forest under Quercus acutissima , Q. glauca , etc.
Also known from South Korea and China (Kim et al., 1993 ; Yang, 1997).

Am anitae in Japan

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ODQd'~'"'l 00 00

CJ C)12IOI'mOO 13

117

Figs. 12- 13. Basidiospores. 12. Amanita a/bo.fla vescens (TNS-F-23 7274, Holotype); 13. Amanita castanopsis (TNS-F-2372 75, Holotype) .

Observations: Yang ( 1997) noted that it is unknown whether A. albojiavescens
possesses clamps or not. Unfortunately, the holotype is not well rehydrating and,hence , the question is still open.
9. Amanita castanopsis Hongo in Travaux mycologiques dedies a R. Kiihner, Bull.
Soc. Linn. Soc. Lyon , no. spec.: 192 (1974).
Fig. 13Basidia 40- 55 X 9.0- 11 f.1m clavate, 4-spored; sterigmata 3- 5 f.1m long; basal
septa with clamps. Basidiospores [50/2/1] (8.5- ) 9.0 (- ll.O) X 5.0- 6.5 f.1m [Q=(1.55- ) 1.58- 1.91 (- 2.06), Q= 1.75 ± 0. 11], dominantly elongate, rarely ellipsoid orsubcylindrical , amyloid, colorless, hyaline, thin-walled, smooth, apiculus small.
Volval remnants on pileus made up of vertically arranged elements: inflated cells
abundant to dominant, subglobose, ovoid to ellipsoid, 35- 75 X 25- 50 f.1m, single andterminal , or 2- 3 (-4) in chains, thin-walled, colorless, hyaline to subhyaline; mixed
with scattered to locally fairly abundant, filamentous hyphae, 3- 5 f.1m wide, thinwa lled, colorless, hyaline; septa often with clamps; vascular hyphae rare .
Specimen examined: Japan: Kokubu, Otsu, 18. IX. I 972, T. Hongo s.n. (Hongo
Herb. 4753 , TNS-F-237275 , Holotype).
Habitat and distribution: In forest of Castanopsis. Also known from South
Korea and China (Kim et al. , 1993 ; Yang, 1997).
Observations: Amanita castanopsis is close to A. virgineoides Bas. The latter
differs, however, from the present species in the broadly ellipsoid to ellipsoid spores(Hongo, 1974 b). In addition , the volval remnants on the pileus of A. virgineoides areusually smaller, and made up of smaller inflated cells mixed with usually scattered
but conspicuous vascular hyphae (cf. Bas, 1969; Yang, 1997).
I 0. Amanita griseofarinosa Hongo in M em . Shiga Univ. , Nat. Sci . 11: 39 ( 1961 ).
Fig. 14
Pileus ea. 2- 5 cm in diameter, convex to applanate, greyish to paler, covered
with greyish, grey to dark grey, tomentose to farinose , volval remnants, sometimes
with subconical paler volval warts; margin appendiculate. Stipe ea. 6-7 X0.5- l cm,
attenuate upwards to subcylindrical ; bulbus clavate, subglobose, covered with grey,
farinose volval remnants. Annulus fugacious .
Basidia 45- 75 X 12- 15 f.1m , 4-spored, probably without clamps. Basidiospores

118

Zhu-Liang Yang and Yoshimichi Doi

OQO 000000

ooo
ooo ooo
ooooDO ooo
~14

~15

~16

Fi gs. 14- 16. Basidiospores. 14. Amanita griseojarinosa (TNS-F-2372 77,
Holotype); 15 . Amanita hongoi, (TNSF-237946); 16. Amanita lutescens(TNS-F -23 7278, Holotype).

[101 /4/2] (8.5- ) 9.0-11.0 (- 12.0) X(6.0- ) 6.5- 8.5 (- lO.O)pm [Q=(l.l - ) 1.18- 1.47(- 1.54), Q= 1.31 ::!:::0.09] , broadly ellipsoid to ellipsoid, amyloid, colorless, hyaline ,thin-walled, smooth, apiculus small.
Specimens examined: Japan : Kurozu, Otsu, !.IX. 1959, T. Hongo s.n. (Hongo
Herb. 2011, TNS-F-237277, Holotype); ditto, 16. VII. 1964, T. Hongo s.n. (Hongo
Herb. 2887, TNS-F).
Habitat and distribution: Gregarious among mosses in woods. Also known
from China (Mao, 1991 ; Yang, 1997).
Observations: Spores from the holotype of A. griseofarinosa are [71 /311] (8.0- )9.0-11.0 (- I2 .0)X(6.0- ) 6.5- 8.5 (- IO.O)pm [Q=(l.l - ) 1.18- 1.47 (- 1.54), Q =1.29::!:::0.09]. Both studied collections are poorly rehydrating. Based on observable basidia, there are probably no clamps.
11. Amanita hongoi Bas in Persoonia 5: 410 ( 1969).
Fig. 15
Pileus ea. 8 cm in diameter, convex to planate, whitish, covered with conical to
subconical, brownish to whitish volval remnants, 1- 1.5 mm high. Stipe ea. 12 X 11.5 cm, base subfusiform; annulus fugacious.
Basidiospores [301111] (7 .5- ) 8.0- IO.O X(6.5- ) 7.0- 8.5 (9.0),um [Q=l.l0- 1.21(- 1.29), Q= 1.16::!:::0.05], subglobose to broadly ellipsoid, amyloid, colorless, hyaline,
thin-walled, smooth, apiculus small. Clamps absent.
Specimen examined: Japan: Right-side ridge of Kurosawa-gawa River, Kurosawa 3-Chome, Oume City, Tokyo, 25 . IX. 1997, Y. Doi s.n. (TNS-F-237946).
Habitat and distribution: Solitary on ground in Quercus-forests. Also known
from South Korea (Kim et al., 1993).
Observations: Amanita hongoi is closely related to A. perpasta Corner & Bas.
A detailed comparison of these two species is given in Bas ( 1969: 412).
12. Amanita lutescens Hongo in J. Jpn. Bot. 33 : 347 (1958).
Fig. 16
Basidiospores [60/211] (7.5- ) 8.0- 10.0(10.5) X(5.0- ) 5.5- 7.0 (7.5)pm [Q =( 1.25- ) 1.31 - 1.73 (- 1.87), Q= 1.48 ::!:::0. 14] , ellipsoid to elongate, amyloid, colorless,hyaline, thin-walled, smooth, apiculus small.
Specimen examined: Japan: Hie-Shrine, Otsu, 20. IX. 1956, T. Hongo s.n.

Amanitae in Japan

019

10

20J11llI~

119

Figs. 17- 19. Amanita sphaerobulbosa.
17. Basidiocarps (a. TNS-F-237770; b.
TNS-F-237569); 18. Hymenium andsubhymenium (TNS-F-237569); 19. Basidiospores (TNS-F-237569).

(Hongo Herb. 1529, TNS-F-237278, Holotype).
Habitat and distribution: Gregarious on the ground under frondose trees. Only
known from Japan.
Observations: The holotype specimen is poorly rehydrating. We do not knowwhether there are any clamps on septa of hyphae in this mushroom.
13. Amanita sphaerobulbosa Hongo in J. Jpn. Bot. 44: 230 ( 1969).
Figs. 17- 19
Pileus ea. 2--4 cm in diameter, convex to planate, white to cream-colored, covered with white, conical to subconical , small volval remnants, ea. I mm high; margin
appendiculate. Stipe 5- 8 X0.5- 0.8 cm, attenuate upwards to subcylindrical; bulb subglobose to napiform, often submarginate, upper part of bulb and stipe base covered
with whitish, small volval remnants. Annulus pendant from ea. 1- 1.5 cm below apexof stipe, finely radially striate above.
Basidia 45- 60 X 12- 15 ,urn, clavate, 4-spored, rarely 2-spored; sterigmata 35 ,urn long; basal septa often with clamps. Basidiospores [I 05/5/5] (7 .0- ) 8.0- 10.5(- 11.5) X(6.5- ) 7.0- 9.5 (- IO.O),um [Q=(I.O- ) 1.02- 1.18 (- 1.25), Q=1.1±0.05] ,globose, subglobose to broadly ellipsoid, amyloid, colorless, hyaline, thin-walled,smooth, apiculus small. Volval remnants on pileus made up of ±vertically arranged
elements: inflated cells very abundant to dominant, subglobose to ovoid to ellipsoid,
15--40 (- 60) X 12- 25 (- 35),um, usually longer in base ofwart, thin-walled, sometimes
slightly thick-walled ( ~ 0 . 5 ,urn thick), often hyaline, 2- 5 in chains, sometimes anastomosing; filamentous hyphae scattered, in base of wart fairly abundant, 2- 7(- IO),um wide, branching, anastomosing and interwoven, thin-walled, hyaline to subhyaline, vascular hyphae rarely scattered, 2- 8 ,urn wide but in upper part of volval
wart often with conspicuous vascular hyphae and inflated cells; clamps rare. Annulus
made up of abundant to very abundant inflated cells, broadly clavate, ellipsoid, sometimes fusiform, 35- 80 X20- 30 ,urn, mixed with fairly abundant filamentous hyphae,

120

Zhu-Liang Yang and Yoshimichi Doi

2- 5 Jlm wide, hyaline, thin-walled; vascular hyphae rare.
Specimens examined: Japan: Mt. Watamuki, Shiga Pref., 17. IX. 1967, T.
Hongo s.n. (Hongo Herb. 3504, TNS-F-244377, Holotype); Along hiking course,
Nagayama-Kyury6 Hills, Kurosawa 3-Chome, Oume City, Tokyo, 5. X. 1997, Y. Doi
s.n. (TNS-F-237770); In Quercus forests, right side of Kurosawa-gawa River, Kurosawa 3-Chome, Oume City, Tokyo, 25. IX. 1997, Y. Doi s.n. (TNS-F-237949); ditto,
2. VIII. 1997, Y. Doi & K. Miyazaki s.n. (TNS-F-182502); Along hiking course, leftside of Kurosawa-gawa River, Nagayama-Kyury6 Hills, Kurosawa 3-Chome, Oume
City, Tokyo, 5. X. 1997, Y. Doi & K. Miyazaki s.n . (TNS-F-237569).
Habitat and distribution: In forests, Only known from Japan.
Observations: Spores of the holotype are [30/1/1] (7.0- ) 8.0-10.0X(7.0- ) 7.59.0 (- 9.5)J1m [Q=(l.O- ) 1.03- 1.18 (- 1.20), Q=l.l0 ::!:::0.05]. Although the holotype
is poorly rehydrating, we have observed clamps on bases of basidia in preparations
from the holotype. Amanita sphaerobulbosa has recently been regarded as a synonymof A. abrupta Peck from North America (e.g., Hongo, 1982; lmazeki & Hongo,
1987). Undoubtedly they are very closely related to each other. However, the sporesof A. abrupta are narrower (Bas, 1969; Jenkins, 1986; Tulloss et al. , 1995). Further,in A. sphaerobulbosa many hyphae and inflated cells, especially in the upper part ofvolval remnants on pileus, are vascular (oleiferous); this is not the case in A. abrupta(Bas, 1969: 434). For the time being, we prefer to treat them separately.
Subgenus Lepidella Section Phalloideae (Fr.) Que!.
14. Amanita oberwinklerana Z. L. Yang & Y. Doi, sp. nov.

Figs. 20- 25
Pileus 3- 8 cm latus, initio semiglobosus, deinde convex us, plano-convexus vel
planus, a/bus, albidus, cremeus vel subjlavidus, reliquiis valvae magnis, coactis,
membranaceis, applanatis, tenuibus, albis ornatus, margine nonstriata, non appendiculata. Lamellae liberae, albae vel cremeae, confertae, lamellulis attenuatis. Stipes
5- 12 X0.5- 1 cm, sursum attenuatus, a/bus vel albidus, annulatus; bulbus volvatus,
napiformis, subglobosus vel clavatus, 1-2 cm latus. Volva breviter limbata, sed saepe
marginata vel subnulla. Annulus membranaceus, a/bus, superior. Caro alba. Basidia
4-sporigera, interdum 2- ve/3-sporigera. Sporae (7.5-) 8.0-10.5 (- 12.5) X(5.5- ) 6.58.0 (- 8.5) Jlm, lato-ellipsoideae vel ellipsoideae, amyloideae. Fibulae absentes.
Holotypus: Y Doi s.n. (TNS-F-237722), 5. X.1997, Nagayama-Kyury6, Kurosawa,
Oume, Tokyo, Japan .
Etymology: "oberwinklerana" is named in honour of Prof. Dr. F. Oberwinkler
on the occasion of his 60th birthday.
Pileus 3- 8 cm in diameter, at first hemisphaerical, then becoming convex topiano-convex to almost applanate, white, whitish, sometimes creamy to pale yellowish [Cream Col or, Pale Chalcedony Yellow, Light Chalcedony Yellow, I A I, 2A2 ,2A3, 3A2, 3A3], slightly deeper on disc, viscid when moist; volval remnants white toAmanitae in Japan121

whitish [!AI, IA2], often in 1- 3 large pieces (I - 2.5 X 1- 2.5 cm) on pileus, felt-like ,submembranous, up to I (- 2) mm thick, randomly arranged, sometimes glabrous andwithout volval remnants, then the volval remnants usually retained at the base of stipe
as limbate volva; margin smooth, occasionally finely striate [0.1 - 0.2 (- 0.3) R], nonappendiculate; trama white. Lamellae free , white, whitish to pale creamy, crowded,edge finely fimbriate and floccose ; lamellulae attenuate to subattenuate, plentiful, in
2- 3 ranks. Stipe 5- 12 X0.5- I cm, usually tapering upward, with apex slightly expanded, white to whitish, subglabrous, finely fibrillose or sometimes with reflexed
scales; context white; basal bulb napiform, subglobose to clavate, 1- 2 cm wide,
1.5- 2 cm high, white to pallid; volva short limbate to limbate, membranous, white towhitish, but often with few volval remnants on stipe base, and then bulb marginate.
Annulus present, superior, pendant from ea. 1- 1.5 cm below apex of stipe, membranous, white to whitish, above fine radially striate. Odor and taste not known. Colorof spore print not known. Dried basidiocarps not giving yellow reaction to KOH solution (5 %- 10%).
Lamella trama bilateral. Mediostratum ea. 30 f..lm wide, distinct, consisting oflong ellipsoid, subfusiform to sometimes subcylindrical, inflated cells, 50- 90 X I 030 f..lm wide, mixed with abundant, branching hyphae, 2.5- 7 f..lm wide; vascular hyphae rare. Lateral stratum consisting of long ellipsoid to fusiform, inflated cells,
40- 70 X 15- 20 f..lm, mixed with fairly abundant filamentous hyphae, 3- 5 f..lm wide, diverging at an angle of 30°-45 ° to the central stratum ; septa without clamps. Subhymenium 25-40 (- 50)f..lm thick, with 2- 3 layers of subglobose, ovoid to short ellipsoid cells, 10- 20 (- 28) X9- I8f..lm. Basidia 33- 55 X9- 12 (- 14)f..lm, clavate, 4-spored,sometimes 2- or 3-spored, rarely 1-spored; sterigmata 3- 5 (- 7) f..lm long; basal septa
without clamps. Basidiospores [155/7/6] (7.5- ) 8.0- 10.5 (- 12.5) X(5.5- ) 6.5- 8.0(- 8.5)f..lm [Q=(I.I3- ) 1.19- 1.50 (- 1.71), Q=l.34±0.10] , broadly ellipsoid to ellipsoid, amyloid, colorless, hyaline, thin-walled, smooth, apiculus small. Lamella edge asterile, somewhat gelatinized strip, 20- 80 f..lm wide in side view, made up of abundant, subglobose, ovoid, clavate to sphaeropedunculate cells (20-40 X 12- 25 f..lm), single and terminal or 2- 3 in chains, thin-walled, colorless, hyaline, often somewhat
gelatinized, mixed with fairly abundant (to abundant), filamentous hyphae, 2- 5 f..lm
wide, irregularly arranged or ±running pararell to the lamella edge. Pileipellis
90- 160 f..lm thick: the upper layer (60- 100 f..lm thick) strongly gelatinized, made up ofradially arranged, filamentous hyphae, 1- 5 f..lm wide, colorless, hyaline; vascular hyphae rare; the lower layer 30- 50 f..lm thick, made up of radially, subradially and compactly arranged, filamentous hyphae, 3- 7 (- 12) f..lm wide, colorless, usually hyaline,
sometimes subhyaline. Volval remnants on pileus made up of irregularly arranged
elements: filamentous hyphae abundant to locally (towards the surface of pileus) very
abundant, 3- 7 (- 12)f..lm wide, branching, interwoven, loosely arranged, hyaline, thinwalled; inflated cells abundant, sub globose, ovoid, 25- 75 X25- 55 f..lm, single and terminal , sometimes 2- 3 in chains, colorless, thin-walled, sometimes slightly thick-122

Z hu-Liang Yang and Yoshimichi Doi

walled ( :s:0.5 ,urn); vascular hyphae rare . Volval remnants on the stipe base and upper
bulb similar to those on the pileus, but filamentous hyphae somewhat more abundant,
especially towards the somewhat gelatinized inner surface. Stipe trama dominantly
consisting of longitudinally arranged, long clavate, terminal cells, 180350X 17-40,um, mixed with 2- 7 (- 12),um wide hyphae scattered interior, fairly
abundant on stipe surface; vascular hyphae rare . Annulus consisting of : :!: : radiallyarranged, abundant to very abundant, filamentous hyphae, 2- 5 (- 8),um wide, thinwalled, branching and anastomosing, mixed with fairly abundant (on upper surface)to very abundant (on edge of annulus), clavate, broadly clavate, subglobose tosphaeropedunculate, inflated cells, 20-45 X I 0- 20 ,urn, colorless hyaline, thin-walled,usually single and terminal; vascular hyphae rare.
Specimens examined: Japan: Along hiking course, left-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 16. VII . 1997, Y. Doi & K .
Miyazaki s.n . (TNS-F- 182594); ditto, 25.IX.l997, Y. Doi s.n. (TNS-F-237780);
Along hiking course, Nagayama-Kyuryo Hills, Kurosawa 3-Chome, Oume City,
Tokyo, 5. X. 1997, Y. Doi s.n. (TNS-F-237572; TNS-F-237722=Holotype! ; TNS-F237727; & TNS-F-237750).
Habitat and distribution: Solitary or gregarious on ground in forests. Presently
only known from Japan.
Observations: Amanita oberwinklerana was probably regarded in East Asia as
A. verna (Bull. : Fr.) Lamarck or A. virosa Bertillon for its white , off-white fruitbodies
with a membranous annulus and white volva, and for its amyloid spores. It differs,
however, from A. verna in its weak volval remnants with abundant inflated cells, andin its smaller spores. Two specimens of A. verna collected from North Africa [Marocco: Maroliko, Tanger, 20. IV 1971 , C. Bas 5523 (L); Larache, 22. IV 1971 , C. Bas
5536 (L)] have been studied. The spores are [170/7/2] (8.5- ) 9.5- 12.0 (- 14.5) X(6.0- ) 6.5- 8.0 (- 10.5),um [Q=(l.19- ) 1.30- 1.67 (-1.79), Q = 1.47 : :!: : 0.11]. The volva
is dominantly made up of filamentous hyphae 2- 8 ,urn wide , mixed with scattered to
locally fairy abundant, inflated cells, 70-80 X25-60 ,urn, often single. Amanita virosa
possesses an irregularly formed, conical to campanulate pileus, and globose to subglobose spores. We have studied a collection of A. virosa from Europe [Poland: Biolowiesa virgin forest, 5. IX. 1966, C. Bas 4697 (L)]. The spores are [30/1 11 /] (8.0- )9.0- 11.0 (- 12.5) X 8.0- 10.0 (- ll.O),um [Q=(l.01 - ) 1.03- 1.16 (- 1.21), Q = l.09 2::
0.05]. The volva is dominantly made up of filamentous hyphae 3- 10 ,urn wide, mixed
with rare to scattered to locally fairly abundant, inflated cells, I 00-175 X40- 70 ,urn,
often single.
Amanita oberwinklerana is somewhat similar to A. bisporigera Atk. and A. magnivelaris Peck (A. elliptosperma Atk.) from North America, and A. eburnea Tulloss
from Belize and Honduras. However, A. bisporigera usually has a limbate volva withdominantly filamentous hyphae, predominantly 2-spored basidia, and broader spores(Jenkins, 1986; Tulloss et al., 1995). A man ita magnivelaris possesses larger basidio-

123

Amanitae in Japan

oooo
C)2
0

0
0

20

Figs. 20- 22. Amanita oberwinklerana(TNS-F-237722, Ho1otype). 20. Basidiocarp; 21. Hymenium and subhymenium; 22. Basidiospores.

Fig. 23. Crushed volval remnant from pileus of Amanita oberwinklerana (TNS -F237722, Holotype).

0

0

0
IOftm

24

Figs. 24- 27. Basidiocarps and basidiaspores. 24- 25. Amanita oberwinklerana(TNS -F-237750); 26. Amanita manginiana
sensu Chiu (TNS-F-182476); 27. Amanita
pseudoporphyria (TNS -F-237281, Ha lotype).

124

Zhu-Liang Yang and Yoshimichi Doi

carps with a limbate volva, and pinkish lamellae (Coker, 1917; Tulloss et al, 1995).
In addtion, A. magnivelaris has no or fewer inflated cells in its annulus (cf. Jenkins,
1978, 1982). Amanita eburnea differs from A. oberwinklerana by its often darker colored pileus, more abundant inflated cells in the annulus, and narrower spores (cf. Tulloss, 1989). The volva of Amanita eburnea seems usually to be retained at the stipe
base.
15 . Amanita manginiana sensu Chiu in Sci. Rept. Natl. Tsing Hua Univ. , ser. B.
Bioi. & Psychol. Sci. 3(3): 166 (1948).
Fig. 26?=Amanita manginiana Hariot & Pat. in Bull. Mus. Hist. Nat. (Paris) 20(3): 155(1914).
Pileus 7- 9 cm in diameter, convex to piano-convex to aplanate, brownish-gray,with darker, radial, innate fibrils , subviscid, usually glabrous, occasionally with white,whitish membranous volval remnants; margin smooth, often with remnants of annulus. Lamellae free, crowded; lamellulae 0- 2, attenuate. Stipe I 0- 15 X 1- 2.5 cm, subcylindrical or slightly tapering upward, white to whitish, smooth or with white fibrils
or scales; volva limbate, 3- 5 X 1.5- 3 cm, membranous, white , upper 1/3 free, lower
part adnate to indistinctly enlarged base. Annulus present, subapical , white, membranous, fragile.
Basidiospores [80/4/3] (5 .5- ) 6.0- 7.5 (- 8.0) X(4.5- ) 5.0- 6.0.um [Q=(I.O- )1.08-1.40 (- 1.46), Q= 1.24±0. 10] , subglobose to broadly ellipsoid, sometimes ellipsoid, amyloid, colourless, hyaline , thin-walled, smooth, apiculus small. Volval remnants on stipe base made up of ±irregularly arranged elements; filamentous hyphae
dominant, 2- 7 (- I 0)
wide, hyaline, thin- to slightly thick-walled ( < 0.5 .urn),branching, interwoven, without clamps; inflated cells scattered to locally fairly abuncolorless, hyadant, subglobose, ovoid, ellipsoid to long ellipsoid, 40- 90 X25-40
line, thin- to slightly thick-walled ( < 0.5 .urn), single and terminal or intercalary; vascular hyphae rare. Outer surface of volva consisting of filamentous hyphae. Inner surface ofvolva strongly gelatinized.
Specimens examined: Japan: Along hiking course, left-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 16. VII. 1997, Y. Doi & K.
Miyazaki s.n. (TNS-F-182476, TNS-F-182482 & TNS-F-182492).
Habitat and distribution: Solitary or gregarious on the ground in forests.
Amanita manginiana sensu Chiu is quite common in China, and is new to Japan.
Observations: Amanita manginiana Hariot & Pat. is not a well known species.
The type specimen was bought in a market in Hue, VietNam, and is now not in goodcondition (Yang, 1997: 167). According to the protologue (Hariot & Patouillard,1914), and the description and the coloured illustration published by Gilbert (1940;
1941 ), there are some differences between Amanita manginiana Hariot & Pat. and A.
manginiana sensu Chiu (Yang, 1997), the latter is an edible mushroom often sold atfree market in summer in Southwest China. In Japan it was probably regarded as A.

.urn

.urn,

Amanitae in Japan125

pseudoporphyria Hongo, the edibility of which is not kown. For comparison of these
two species, see the observations under A. pseudoporphyria.
It is interesting to note that the outward appearance of A. manginiana sensu
Chiu and A. pseudoporphyria is somewhat similar to that of the famous, fatally poisonous mushroom A. phalloides (Fr.) Link, originally described from Europe. However, the former two seem to have no close phylogenetic relationships with the latteraccording to molecular data (Weif3 et al. , 1998).
16. Amanita pseudoporphyria Hongo in J. Jpn. Bot. 32: 141 ( 1957).
Fig. 27
Pileus 3- 10 cm in diameter, convex to piano-convex to applanate, greyishbrown, brownish, darker at center, with darker innate fibrils, subviscid, glabrous or
bearing white , whitish, creamy or grayish, membranous volval remnants; marginsmooth, often with white remnants of annulus, context white. Lamellae free, crowded; lamellulae 0- 2, attenuate. Stipe 3- 11 X 0.3- 2 cm, subcylindrical or slightly tapering upward, white to whitish, smooth or with white fibrils or scales; volva limbate, 13 X0.5- 2.5 cm, membranous, white to whitish, sometimes with greyish tinge especially towards upper rim of limb, upper 1/3 free, lower part adnate to indistinctly enlarged or almost uninflated, sometimes rooting base. Annulus present, apical to subapical, white, membranous, fragile.
Basidiospores [110/6/3] (6.0- ) 6.5- 8.5 (- 11.5) X(4.0- ) 4.5- 5.5 (- 6.5),um [Q=( 1.18- ) 1.30- 1.78 (- 2.17), Q= 1.53 ±0.17] , ellipsoid to elongate, rarely subcylindrical , amyloid, colorless, hyaline, thin-walled, smooth, apiculus small. Volval remnants on stipe base made up of ±irregularly arranged elements: filamentous hyphaeabundant to very abundant, 2- 7 (- 12) ,urn wide, hyaline, thin- to slightly thick-walled( < 0.5 ,urn), branching, interwoven, without clamps; inflated cells fairly abundant to(locally) abundant, ovoid, ellipsoid to long ellipsoid, sometimes subglobose, 40120 X 25- 60 ,urn, colourless hyaline, thin-walled, single and terminal, sometimes intercalary, occasionally in short chains, vascular hyphae rare. Outer surface of volva
consisting of more abundant filamentous hyphae. Inner surface of volva strongly
gelatinized.
Specimens examined: Japan: lshiyama-Hiratsu-cho, Otsu, 23. VII. 1956, T.
Hongo s.n. (Hongo Herb. 1506, TNS-F-237281, Holotype); Right-side ridge of Kurosawa-gawa River, Kurosawa 3-Chome, Oume City, Tokyo, 2. VIII. 1997, Y. Doi & K.
Miyazaki s.n. (TNS-F-182564); Along hiking course, left-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 16. VII. 1997, Y. Doi & K.
Miyazaki s.n. (TNS-F-182479).
Habitat and distribution: Gregarious on ground in forests (e.g., of Pinus).
Also known from South Korea and China (Kim et al., 1993; Yang, 1997).
Observations: Spores of the holotype of A. pseudoporphyria are [60/3/1 /] 7.09.0 (- 11.5) X(4.0-) 4.5- 5.5 (- 6.5),um [Q=(l.36- ) 1.43- 1.83 (- 2.17), Q= 1.63±
0.14], ellipsoid to long ellipsoid, rarely subcylindrical , amyloid, colorless, hyaline,

126

Zhu-Liang Yang and Yoshimichi Doi

thin-walled, smooth, apiculus small. Amanita manginiana sensu Chiu is very similar
to this species, but differs from the latter in its shorter and wider spores, and in thevolva with fewer inflated cells. Furthermore, the stipe base of A. pseudoporphyria isoften rooting (Yang, 1997: 168).
17. Amanita subjunquillea lmai in Bot. Mag. (Tokyo) 47: 424 (1933). Figs. 28- 29
Pileus 3- 5 cm in diameter, at first hemisphaerical , then convex to piano-convex,
brownish yellow, citrin-yellow at disc, yellowish towards margin, sometimes yellowish with carneous tinge, subviscid, glabrous and usually without volval remnants;
margin smooth or finely striate, non-appendiculate; context white or whitish. Lamellae free, crowded, lamellulae 0- 2, attenuate. Stipe 5- 1 I X 0.5- 1 cm, subcylindrical orslightly tapering upward, whitish to yellowish, with yellow to yellowish fibrils or
scales; base with a bulb; volva limbate, 2- 3 X 1- 2 cm, ea. 0.5- 1 cm thick, membranous, white to whitish. Annulus present, pendant from ea. 1.5 cm below apex ofstipe, membranous, white.
Subhymenium 20- 30 pm thick, with 2- 3 layers of subglobose, ovoid to ellipsoid cells, 8- 18 X 6- 15 pm . Basidia 28-40 X 8- 12 pm , clavate, 4-spored; sterigmata
3-4 pm long; basal septa without clamps. Basidiospores [50/3/3] (6.0- ) 6.5- 8.0(- 8.5)X(5.0- ) 6.0- 7.5 (8.0)pm [Q= 1.0- 1.17 (- 1.2), Q= 1.09~0.05] , globose to
subglobose, rarely broadly ellipsoid, amyloid, colorless, hyaline , thin-walled, smooth,
apiculus small. Annulus dominantly consisting of subradially arranged, filamentous
hyphae, 2- 7 pm wide, mixed with scattered to fairly abundant, broadly clavate, sometimes ovoid to subglobose, inflated cells, 20- 50 X 15- 30 pm, colorless, hyaline, thinwalled, usually single and terminal, rarely 2- 3 in chains; vascular hyphae rare.
Specimens examined: Japan: Along hiking course, left-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 16. VI I. 1997, Y. Doi & K.
Miyazaki s.n. (TNS-F-182483); Right-side ridge ofKurosawa-gawa River, Kurosawa
3-Chome, Oume City, Tokyo, 25. IX . 1997, Y. Doi s.n. (TNS-F-237829); Along hiking course, Nagayama-Kyury6 Hills, Kurosawa 3-Chome, Oume City, Tokyo,
5. X. 1997, Y. Doi s.n. (TNS-F-237728).
Habitat and distribution: Solitary or gregarious on ground in forests. Also
known from China (Mao, 1991 ).
Observations: This species is characterized by its small, globose to subglobose,amyloid spores. A white variety of this taxon also occurs in Japan (see below).
18. Amanita subjunquillea var. alba Z. L. Yang in Biblioth. Mycol. 170: 174 ( 1997).
Figs. 30- 31
Pileus 3- 5 cm in diameter, at first hemispherical , then convex to piano-convex,
white to whitish, sometimes creamy to pale yellowish at disc, subviscid, glabrous andusually without volval remnants; margin smooth or finely striate, non-appendiculate;context white, whitish. Lamellae free , crowded; lamellulae 0- 2, attenuate. StipeAmanitae in Japan

127

Figs. 28- 31. Basidiocarps, hymenium , subhymenium and basidiospores. 28- 29. Amanita subjunquiella (TNS-F-182483); 3031. Amanita subjunquiella var. alba (Basidiocarp: TNS-F-237599; Basidiospores:
TNS-F-194890).

5- 11 X 0.3- 1 cm, subcylindrical or slightly tapering upward, white to whitish, smoothor with white fibrils or scales; base bulbous; valva limbate, 2- 3 X 1- 2 cm, membranous, white to whitish. Annulus present, 1- 2 cm below apex of stipe, white, membranous. Pilei of dried basidiocarps often give yellow reaction to KOH solution (5%) .
Basidia 30-45 X 9- 11 J..Lm , clavate, 4-spored, rarely 1-spored, sterigmata 3-5 J..Lm
long, basal septa without clamps. Basidiospores [124/10/6] (6.0- ) 6.5- 8.0 (- 9.0) X(5.0- ) 6.0- 7.5 (-8.5),um [Q = 1.0- 1.15 (- 1.3), Q= 1.07::!::0.05], globose to subglobose, rarely broadly ellipsoid, amyloid, colorless, hyaline, thin-walled, smooth, apiculus small.
Specimens examined: Japan : Along hiking course, left-side ridge of Nagayama-Kyuryo Hills, Kurosawa 3-Chome, Oume City, Tokyo, 5. X. 1997, Y. Doi s.n.
(TNS-F-237599, TNS-F-237594, & TNS-F-237748); Right-side ridge of Kurosawagawa River, Kurosawa 3-Chome, Oume City, Tokyo, 25. IX. 1997, Y. Doi s.n. (TNSF-237846); Do-zawa, Fudagou, Kiyosumi , Chiba Pref., 14. VII. 1983, Y. Doi s.n.
(TNS-F-174870); Kyoto Univ. Forest in Ahiu, Kyoto, 24. VII. 1964, Y. Doi s.n. (TNSF-194890).
Habitat and distribution: Solitary or gregarious on ground in forests. Amanita
subjunquillea var. alba was described from China, and is new to Japan.
Observations: Amanita subjunquillea var. alba differs from A. subjunquillea
var. subjunquillea in its white pileus and stipe. This taxon was probably often treatedas A. virosa, a species originally described from Europe. The latter possesses, however, irregularly formed, conical to campanulate pileus, thin and fragile annulus, anddistinctly larger basidiospores (Yang, 1997: 177).

128

Zhu-Li ang Yang and Yoshimichi Doi

Acknowledgements
The first author is very grateful to Dr. C. Bas, Leiden, for discussions with himon the systematics of Amanita, and for providing him working space for the study ofsome collection of Amanita deposited there and to Dr. R. E. Tulloss, New Jersey, for
his valuable published and unpublished data of Amanita, and for his many informative discussions and correspondence, and for criti cally reviewing the manuscript. He
expresses his sincere thanks to Prof. Dr. F. Oberwinkler, Ti.ibingen, for hi s guidance.
He thanks his colleagues in Japan for their organizations, hospitality and support during his visit to Japan. The authors extend thanks to Dr. T. Hongo for his kind help.
This study was supported partially by the Grant-in-Aid for Scientific Research ,Ministry of Education, Science, Sports and Culture, Japan (no. I 0041186 in 1998),
and by the National Natural Science Foundation of China under the project no.
39800007 for the first author. The field works by the second author in the forest areaof Oume City, Tokyo, Japan in 1997 were financially supported by the Grant-in-Aid
from the World Wide Fund for Nature, Japan (WWF Japan ).

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