全 文 ：鬼笔腹菌在东亚首次发现?
杨祝良1 , 葛再伟1 , 2
(1 中国科学院昆明植物研究所生物多样性与生物地理学院重点实验室 , 云南 昆明 650204 ;
2 中国科学院研究生院 , 北京 100049 )
摘要 : 根据采自我国四川的材料和对模式标本的研究 , 对鬼笔腹菌 ( Phallogaster saccatus) 的特征进行了详
细描述 , 并附有插图。该种所隶属的鬼笔腹菌属 ( Phallogaster) 为一个单型属 , 在东亚为首次报道。其主
要特征是包被单层 , 表面有不规则的下陷 , 成熟时下陷处发育成不规则小孔 ; 孢体由不规则的小腔组成 ,
成熟时与中柱一并全部胶化溶解 , 仅剩下深绿色至橄榄色的孢子成堆附着在包被的内表面 ; 担子果无菌
托 , 成熟时袋状、中空。
关键词 : 鬼笔腹菌科 ; 幅片包目 ; 分类学
中图分类号 : Q 949 文献标识码 : A 文章编号 : 0253 - 2700 (2008) 02 - 147 - 04
Phallogaster saccatus (Basidiomycetes, Fungi) ,
First Record from East Asia *
YANG Zhu-Liang1 , GE Zai-Wei1 , 2
( 1 Key Laboratory of Biodiversity and Biogeography, Kunming Instituteof Botany, Chinese Academy of Sciences,
Kunming 650204 , China; 2 Graduate University of Chinese Academy of Sciences, Beijing 100049 , China)
Abstract: Phallogaster saccatus is reported, described and illustrated fromEast Asia for thefirst time . It is characterized
by its sac-like, single-layered peridiumwith irregularly shaped thin depressed areas which becomes perforated, the colu-
mella that deliquesces and entirely disappears by maturity, the olivaceousspore mass (adheringto the inner surface of the
otherwise hollow basidioma) that breaks irregularly whenmature, the absence of a volva and the bacilliformto cylindric,
smooth basidiosporeswith a nearly truncate base and a short sterigmal appendage . The genus Phallogaster is currently
viewed as monotypic .
Key words: Phallogastraceae; Hysterangiales; Taxonomy

Montane southwestern China is one of the world′s
34 hotspots of biodiversity ( Boufford and van Dijk,
2000; Mittermeier et al. , 2005 ) . The fauna and flora
there are relatively better known than the mycota . In
the last ten years, mycological expeditions to the re-
gions have been taken annually, and many fungal col-
lections have been made . A taxonomic study revealed
that the genus Phallogaster occurs there . This genus
has not been reported fromAsia before .
Materials and methods
Mature and developing basidiomatawere collected in aforest
dominated by Picea and Abies in Sichuan Province, China .
Specimenswereannotated and?or photographed in the field . Co-
lour codes of the form“2B2”indicatea colour sample, row, and
colour block in Kornerup and Wanscher (1981) . Specimenswere
云 南 植 物 研 究 2008 , 30 (2) : 147～150
Acta Botanica Yunnanica

? ?Foundation items: This study was supported by the Knowledge Innovation Program of the Chinese Academy of Sciences ( KSCX2-YW-G-025 ) and
the National Natural Science Foundation of China ( Nos . 30525002 , 30420120049 ) . The field work was supported by the National Science
Foundation of the USA ( DEB-0321846)
Received date: 2007 - 07 - 06 , Accepted date: 2007 - 11 - 25
作者简介 : 杨祝良 (1963 - ) 男 , 研究员 , 主要从事高等真菌的分类及分子系统学研究。
dried in an electric drier, and then deposited in herbaria . Her-
bariumabbreviations follow Holmgren et al. ( 1990 ) with one ex-
ception: HKAS-the Herbarium of Cryptogams, Kunming Institute
of Botany, Chinese Academy of Sciences .
Tissueswere mounted in 3 - 5% KOH and Melzer′s reagent
for microscopic examination . Q refers to the length?width ratio of
basidiospores; Q refers to the average Q of all basidiospores±
sample standard deviation .
Taxonomy
Phallogaster saccatusMorgan, Journ . Cincinnati
Soc . Nat . Hist . 15 : 171 , 1893 . Fig . 1 : 1 - 7
Basidiomata ( figs . 1 - 3) scattered to gregarious,
3 - 5 cm×2 - 3 (4) cm, ovoid tosubglobose to piriform
when young, becomingovoid to piriformor subellipsoid
when mature, hollow to very base at maturity; stipe
short to nearly sessile, up to 1 .5×1 cm, whitish, at-
tenuate towards base with white rhizomorphs; volva ab-
sent . Peridium single-layered, 1 - 2 mm thick, with
surface glabrous, pale olivaceous ( 2B2-2C2 , 3B2 ,
29B2-29B3) , but often with pinkish tinge ( 6A2-6B2 )
especially in lower parts, with irregularly shaped, thin,
lighter colored, depressed areas becoming perforate at
mature, then with holes 0 .3 - 0 .7 (1 .2) cmwide, fi-
nally often breaking irregularly at apex . Gleba at first
irregularly lobed, olivaceous green ( 29D4-29D5) , vis-
cid, with gelatinous tramal plates and with irregular to
angular, minute locules; at maturity entirely del-
iquesced and leavingslimy sporemass adhering to inner
surfaceof peridium . Columella extending from base of
basidioma to center of gleba, branching and penetrating
gleba and separating its lobes, gelatinous and translu-
cent, entirely deliquesced and absent at maturity .
Spore mass mucilaginous, dark green to olivaceous
(30F4-30F6) , adhering in irregular sizes and shapes to
inner surfaceof matureperidium . Odor not repugnant,
weakly chive ( Alliumtuberosum)-like .
Basidiospores ( Fig . 1: 5 , 7 ) [ 40?2?1 ] ( 4 .5 )
5 .0 - 6 .5 ( 7 .0) ×1.8 - 2 .3 (2 .5 ) μm, Q = (1 .8 )
2 .27 - 3 .33 ( 3 .61 ) , Q = 2.68 ± 0 .35 , statismos-
poric, orthotropic, oblong to bacilliform to cylindric,
greenish to nearly colorless and hyaline in KOH,
smooth, thin-walled, non-amyloid, non-dextrinoid;
base nearly truncate, with very short sterigmal append-
age . Basidia ( Fig . 1 : 4 , 6 ) 20 - 32×4 - 6μm, clav-
ate to narrowly clavate, 6 - 8-spored; sterigmata< 0 .5
μmlong . Cystidia absent . Tramal plates (Fig . 1 : 4)
80 - 200μm thick, with subparallel to loosely interwo-
ven, multibranched hyphae 2 - 4 ( 6 ) μm broad, hya-
line, thin-walled, strongly gelatinized . Peridiopellis
well differentiated, dominantly consistingof subfusiform
or ellipsoid, sometimes ovoid, thin-walled, colorless
and hyaline, non-gelatinous inflated cells 10 - 20 (30)
μm broad and in chains, and mixed with scattered to
locally abundant ( on inner surface of peridiopellis,
abundant) , non-gelatinized filamentous hyphae 2 - 7
μm broad; outer surface of peridiopellis bearing rela-
tively common±peripherally arranged, 2 - 6μm broad
filamentous hyphae with yellowish to ochreous contents
in KOH; thin, depressed areas of peridium dominantly
consisting of gelatinized filamentous hyphae 2 - 5 μm
wide . Clamp connections common in all parts of ba-
sidioma .
Habit, habitat, distribution and season—Fruit-
ing in summer; scattered to gregarious, among fallen
wood in forest dominated by Picea and Abies; North
America ( Canada, Mexico and USA) , Central America
(Costa Rica) , Europe ( Austria, France, Germany,
Hungary, Italy, Poland, Slovakia, Slovenia, Spain,
Switzerland, and Turkey) , new to East Asia .
COLLECTIONS EXAMINED—China , SI-
CHUAN PROVINCE , Xiangcheng County, Reda, alt .
3 700 m, 15 .VII .2004, Z . L . Yang 4127 ( HKAS
45512) . USA , OHIO STATE , Preston, 18 . IV .
1892 , A . P . Morgan s . n . (FH 1142-holotype) .
Discussion
The above description is based on a collection
made in southwestern China . In comparison with other
descriptions of this species ( e . g . Fitzpatrick, 1913;
Lohwag, 1936; Thaxter, 1893; Aron et al. , 2006 ) ,
theChinese collection has a slightly longer and broader
basidiospores . The holotype, consistingof ahalf basid-
ioma, was probably not fully mature when dried . The
basidiospores fromthe holotype are [15?1?1] 4 .5 - 5 .5
(6 .0) × ( 1 .5 ) 1 .8 - 2 .0 (2 .2 ) μm, Q= (2 .4) 2 .5
- 3.0 (3 .06) , Q = 2 .70±0 .19 (Fig . 1: 7) . Phal-
841 云 南 植 物 研 究 30 卷
Fig . 1 : 1 - 5 : Phallogaster saccatus ( HKAS 45512) : 1 . Basidiomata at different stagesof development . Noteyoung basidiomatawith irregular-
ly shaped, thin, lighter colored, depressed areas, which perforate as holes in mature basidiomata; 2 . Basidiomata in longitudinal sections
showing peridium, gleba and columella . Note the right basidioma with two thin and depressed areas which become holes in later development;
3 . Gleba and columella entirely deliquesced and left slimy sporemass adhering to the inner faceof theperidium, and basidiomabecomeshollow
to its very base; 4 . Tramal plate, subhymenium and hymeniumfrom the right basidioma in fig . 1 : 2 ; 5 . Basidiospores from a spore mass in
fig . 1 : 3 . 6 - 7 : Basidia and basidiosporesof P . saccatus ( FH 1142-holotype) . Theholotype consistsof half of a longitudinally split basidioma
dried at a stage of development comparable to the left basidiomaof fig . 1 : 2
logaster saccatus was originally described from eastern
North America and later found in Europe and Central
America ( e . g . Morgan, 1893; Lloyd, 1907; Lo-
hwag, 1936; Furia and Bernicchia, 1983; Gross,
1978; Krieglsteiner, 1991; Calonge et al. , 2005;
Aron et al. , 2006; Doˇgan, 2006) . Thepresent article
reports this taxon occurringoutsideAmerica and Europe
for the first time .
The systematic arrangement of Phallogaster was
long under dispute . Morgan ( 1893 ) and Thaxter
(1893 ) regarded the genus as a member of the Phal-
loideae, oneof thefour subdivisions (Phalloideae, Hy-
menogastreae, Nidularieae, and Lycoperdaceae) of
Gasteromycetes at that time . Fischer (1899 ) placed it
in the Hysterangiaceae of Hymenogastrineae near Pro-
tubera, which assignment was followed by Fitzpatrick
( 1913 ) . Coker and Couch ( 1928 ) and Johnson
(1929 ) placed Phallogaster in the Hysterangiaceae of
Gasteromycetes and Clathraceaeof Phallales respective-
ly, whileG?umann ( 1964 ) andDring (1973) placed it
in Hysterangiaceae of Phallales . Jülich ( 1981) placed
it in Hysterangiaceaeof Hysterangiales . In their molec-
ular phylogenetic analysis, Hosaka et al. (2007 ) pro-
vided evidence supporting placement of Phallogaster
and its relatives ( some Protubera-like and Trappea-
like taxa) in a newly named, independent family,
namely Phallogastraceae, in theHysterangialesof Phal-
lomycetidae .
In the literature two additional species have been
described in Phallogaster, i . e ., P . whitei Peck
(1907) and P . globosus Lloyd (1917 ) . Phallogaster
whitei is adepauperate formof P . saccatus accordingto
Llyod (1909 ) . Phallogaster globosuswas originally de-
scribed from New Zealand . Cunningham ( 1944 ) sug-
9412 期 YANG and GE: Phallogaster saccatus ( Basidiomycetes, Fungi) , First Record fromEast Asia
gested that the species was based on an“egg”of some
phalloid,“probably thatof Clathrus cibarius”. Howev-
er, Castellano and Beever ( 1994 ) suspected that it
represents a species of Protubera . To date the genus
Phallogaster is still monotypic . Research indicating
molecular differentiation between populations of differ-
ent continents is not known to us .
Kobayasia nipponica (Kobayasi ) S . Imai & A .
Kawam . (≡ Protubera nipponica Kobayasi ) alsooccurs
in southwestern China and is somewhat similar to
P . saccatus . However, the former can easily bedistin-
guished from the latter by its concentrically arranged,
non-deliquesced gleba and a complete peridiumwithout
holes (Liu 1994; Liu 2005) .
Acknowledgements: The authors are very grateful to Prof . Dr .
D . H . Pfister, Farlow Herbarium at Harvard University for al-
lowingthemaccess to thetypeof P . saccatus there . They express
their sinceregratitudetoDr . R . E . Tulloss (New Jersey, USA)
for improving the manuscript .
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