全 文 ：狭义干蘑属 (蘑菇目 ) 概要及新的系统学处理 ?
王 岚1 , 杨祝良1
??
, 张丽芳1 , MUELLER G . M .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重点实验室 , 云南 昆明 650204;
2 美国芝加哥费尔德自然历史博物馆植物研究室 , 芝加哥 IL 60605 - 2496 , 美国 )
摘要 : 对狭义干蘑属 ( Xerula s . str .) 的范围进行了重新界定 , 该属仅包括模式种干蘑 ( X. pudens) 及其
周围的几个种。属的特征是担子果金钱菌状 , 有假根 ; 菌盖及菌柄表面干燥 , 被有黄色至褐色直立、披针
形的刚毛 ; 菌褶边缘无不育带 ; 子实层有近梭形的小担子。这些特征可以将干蘑属的物种从干蘑属 - 小奥
德蘑属复合群中分开。现知干蘑属包含六种 , 本文提供了这些种的检索表 , 并对各种进行了描述、图示或
讨论。其中 , 硬毛干蘑 ( X. strigosa) 是一新种 , 它与干蘑 ( X. pudens) 和中华干蘑 ( X. sinopudens) 相似 ,
但与干蘑不同在于其担孢子较长 , 锁状联合稀少 , 侧生囊状体顶端壁薄 ; 与中华干蘑的区别在于其侧生囊
状体壁厚 , 顶端有头状至近头状膨大并覆盖有结晶 , 担孢子宽椭圆形至椭圆形。中华干蘑和小刺干蘑
( X. setulosa) 分别为我国和哥斯达黎加的新记录种。干蘑属虽然广布北半球 , 但未见东亚 - 美洲或欧亚广
布的物种分布类型。
关键词 : 生物地理学 ; 命名 ; 物种多样性 ; 分类
中图分类号 : Q 949 文献标识码 : A 文章编号 : 0253 - 2700 (2008) 06 - 631 - 14
Synopsis and Systematic Reconsideration of
Xerula s . str . ( Agaricales) *
WANG Lan1 , YANG Zhu-Liang1 * * , ZHANG Li-Fang1 , MUELLER G . M .2
( 1 Key Laboratory of Biodiversity and Biogeography, Kunming Instituteof Botany, Chinese Academy
of Sciences, Kunming 650204 , China; 2 Department of Botany, Field Museumof
Natural History, Chicago, IL 60605 - 2496 , USA )
Abstract : Thegenus Xerula s . str . has been revised and is circumscribed as anassemblageof taxa strictly aroundthetype
of thegenus, X. pudens . The collybioid basidiomawith a pseudorhiza; the dry surfaces of the pileus and the stipecovered
with erect, lanceolate, yellowto brown, thick-walled setae; the subacerose basidioles in the hymeniumand the fertile la-
mellar edge delimit Xerula species from other taxa of the Xerula-Oudemansiella complex . Currently, Xerula contains six
species . A key to the species, descriptions, illustrations or discussions of thetaxaareprovided . Xerula strigosa represents
a new species resembling X. pudens and X. sinopudens . Xerula sinopudens and X. setulosa are records new to China and
Costa Rica respectively . Although Xerula is widely distributed in the Northern Hemisphere, no species of it has an East
Asian-American or European-Asian distribution pattern .
Key words: Biogeography; Nomenclature; Species diversity; Taxonomy
云 南 植 物 研 究 2008 , 30 (6) : 631～644
Acta Botanica Yunnanica DOI : 10 .3724?SP. J . 1143 .2008.08156
?
?? ?Author for correspondence; E-mail : fungi@ mail. kib. ac. cn; fungiamanita@ gmail . com
Received date: 2008 - 08 - 11 , Accepted date: 2008 - 09 - 18
作者简介 : 王岚 ( 1981 - ) 女 , 硕士 , 主要从事高等真菌的分类学和生物地理学研究。 ?
Foun ?dation items: The National Natural Science Foundation of China ( Nos . 30470010 and 30525002 ) , the Knowledge Innovation Program of the
Chinese Academy of Sciences ( No . KSCX2-YW-G-025) , and theKey Laboratory of Biodiversity and Biogeography, ChineseAcademy of Sci-
ences ( No . 0806371111 ) , and the Ministry of Science and Technology of China ( No . 2005DKA21006 )
The genus Xerula was initially proposed byMaire
(1933 ) to accommodate Agaricus longipes Bull . [ ≡
X. pudens ( Pers .) Singer] . He emphasized the pu-
bescent basidioma and the absence of a gelatinized
layer on the pileal surface in A. longipes, that are not
found in A. radicatus ( Relhan: Fr .) Fr . [ ≡ Oude-
mansiella radicata ( Relhan: Fr .) Singer ] . Moser
(1955) treated Xerula and Mucidula Pat . ( 1887 ) as
synonyms of Oudemansiella Speg . (1881) , which was
based on the neotropical O. platensis ( Speg .) Speg .
Singer ( 1962a, b, 1964 , 1986 ) adapted Moser′s
treatment, yet regarded Xerula as a subgenus within
thegenus Oudemansiella .
D?rfelt ( 1979 , 1980a, b, 1981 , 1983 , 1984 )
accepted and greatly enlarged the genus Xerula to in-
clude species both with a dry or viscid pileus . He
transferred the sections Albotomentosae Clémen?on,
Protoxerula Clémen?on and Radicatae Clémen?on from
Oudemansiella to Xerula based on the similarities of
the absence of an annulus, the presence of a pseudo-
rhiza, a hymeniform pileipellis, and gymnocarpic de-
velopment of basidiomata, and, thus, restricted Oude-
mansiella to the species with an annulus on the stipe
without a pseudorhiza and hemiangiocarpic or bivelan-
giocarpic development of basidiomata [ e.g ., O. ca-
narii ( Jungh .) H?hn . and O. mucida ( Schrad .)
H?hn .] . His concept was accepted by many authors,
and many species have been transferred from Oude-
mansiella to, or newly described in, Xerula . Several
section names, e.g . Albotomentosae ( Clémen?on)
D?rfelt, Hyalosetae D?rfelt, Radicatae and Xerula
have been proposed in the genus Xerula that has been
divided into three subgenera, viz . Xerula, Radicatae
D?rfelt and Ixoflammula Contu ( Boekhout and Bas,
1986; Redhead et al. , 1987; Petersen and Methven,
1994; Boekhout, 1999; Halling and Mueller, 1999;
Contu, 2000; Petersen, 2000; Horak, 2005; Petersen
and Nagasawa, 2006; Petersen and Baroni , 2007; Pe-
tersen, 2008a, b, c) .
Meanwhile, many authors did not accept D?rfelt′s
revised concept of Xerula and Oudemansiella, and
treated Xerula as a subgenus of Oudemansiella
( Clémen?on, 1979; Singer, 1986; Pegler and Young,
1987; Rexer and Kost, 1989a, b; Yang and Zang,
1993; Corner, 1994; Yang, 2000; Mizuta, 2006) .
A recent analysisof the internal transcribed spac-
ers region ( ITS) data set showed that X. pudens and a
related species, X. hispida Halling et G . M . Mueller
formed a separated clade from the complexes of“ O. -
canarii” ( in fact O. platensis) and X. furfuracea
( Peck ) Redhead [ ≡ O. radicata var. furfuracea
(Peck) Pegler] ( Binder et al. , 2006) . Our own mo-
lecular phylogenetic reconstruction with increased taxon
sampling and using sequences of the ITS and the large
subunit of nuclear ribosomal DNA ( nLSU) also reveals
that Xerula s . str . and Oudemansiella are indepen-
dent, monophyletic lineages and should be treated as
separategenera (Zhang et al. , 2003; unpublished data
of Zhang) . In this paper, we reported our studies of
Xerula in its strict sense .
Materials and Methods
Macromorphological characters were described based on
fresh material , colored photos, and?or field-notes . For micro-
morphological studies, sections were cut with a razor blade from
basidiomata and mounted on slides in 5 % KOH , and then ob-
served andmeasured under acompoundmicroscope .Color desig-
nations ( e.g ., 5E7 ) are from Kornerup and Wanscher ( 1981 )
while color nameswith first letters capitalized ( e.g ., Cinnamon
Brown) are from Ridgway (1912) . In the basidiospore descrip-
tions, the abbreviation [ n?m?p] indicates n basidiospores mea-
sured from mbasidiomata of pcollections in5 % KOH solution;
Q means“length?width ratio”of a spore in sideview; Q is aver-
age Q of all basidospores±sample standard deviation . Herbaria
are abbreviated according to Holmgren et al. ( 1990 ) with one
exception: HKAS= Herbariumof Cryptogams, Kunming Institute
of Botany of theChinese Academyof Sciences, which is not list-
ed in the index or relative publications .
Taxonomic Description
Xerula Maire, Treb . Mus . Ciènc . Nat . Barcelo-
na 15 , Sèr . Bot . 2 : 66 . 1933 .
Basidiomata collybioid, growingon theground but
connected to buried wood by a pseudorhiza . Pileus
convex to plane, umbonate or slightly depressed, yel-
lowish, brownish to dark brown, unpolished and dry,
neither viscid nor gelatinized, covered with yellowish
and brownish setae . Lamellae sinuate to adnexed,
236 云 南 植 物 研 究 30 卷
thick, subdistant, whitish . Stipe non-annulate, radi-
cate, surface covered with yellowish to brownish setae;
pseudorhiza tapering . Basidiospores globose, subglo-
bose, broadly ellipsoid or ellipsoid, thin-walled,
smooth, inamyloid . Basidia clavate, mostly 4-spored,
occasionally 2-spored; basidioles fusiformtosubacerose .
Pleurocystidia fusiform to ventricose, thin- to thick-
walled, prominent .Lamellar edge fertile; cheilocystidia
scattered, often similar to pleurocystidia . Hymenophoral
tramaregular to subregular . Pileipellis hymeniformcom-
posedof clavate, pyriform, sphaeropedunculateor near-
ly globose cells . Pileocystidia ( pileosetae) and caulo-
cystidia (caulosetae) setaceous, ventricoseat thebase,
tapering towards apex, thick-walled, yellowish to
brownish . Clamp connections common, rareor absent .
Type of genus: Xerula pudens (Pers .) Singer
Key to the species of Xerula
1 . Clamp connections common to very common; Europe
2 . dPileus with relatively short ( < 1 mm) setae; pleurocystidia thick-walled (up to 6μm) , often with a capitate to subcapitate and
yellowish crystalline deposited apex ( usually ca . 2μmthick-walled) X . pudens⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
2 . cPileuswith long ( upto3 mmlong) setae; pleurocystidia thin-walled (mostly < 1μmthick) with an occasionally subcapitate apex
rarely with crystalline deposits X . melanotricha⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
1 . Clamp connections rare or absent; East Asiaor America
3 . Basidia 4-spored; pleurocystidia fusiform, capitate to subcapitate; Asia
4 . ?Basidiospores subglobose to broadly ellipsoid (Q= 1.05 - 1 .20) ; pleurocystidia thin-walled (mostly ≤0 .5μm thick) , usually
with non-capitateapex without crystalline deposits; mostly in subtropical and tropical areas X . sinopudens⋯⋯⋯⋯⋯⋯⋯
4 . ?Basidiospores broadly ellipsoid to ellipsoid (Q= 1 .10 - 1.40) ; pleurocystidia thick-walled (mostly 1 - 3μmthick) , oftenwith a
capitate to subcapitate apex covered with crystalline deposits; mostly in temperate and subtropical areas X . strigosa⋯⋯⋯
3 . Basidia 2 or 4-spored; pleurocystidia slender fusiform, usually with a non-capitate apex; Central and South America
5 . Basidia 2-spored; basidiospores subglobose to subovoid; Central and northern South America X. hispida⋯⋯⋯⋯⋯⋯⋯⋯
5 . ?Basidia 4- or amixtureof 2- and4-spored; basidiospores subgloboseor broadly ellipsoid to ellipsoid; Central and SouthAmerica
X. setulosa⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
Xerula pudens (Pers .) Singer, Lilloa 22: 289 .
1951 . ——— Agaricus radicatus var. pudens Pers .,
Syn . Meth . Fung . 2 : 313 .1801 . ——— Gymnopuspu-
dens (Pers .) Gray, Nat . Arr . Brit . Pl . ( London) :
605 . 1821 .——— Agaricuspudens (Pers .) Pers ., My-
col . Eur . 3 : 140 . 1828 . ——— Collybia pudens
(Pers .) S . Lundell , Fung . Exs . Suec . no . 1717 .
1949 . ——— Oudemansiella pudens ( Pers .) Pegler et
Young, Trans . Br . Mycol . Soc . 87 : 590 . 1987 .
(Figs. 1 - 5)
Pileus yellowish brown to dark brown, covered
with yellowish brown setae on the surface . Basidio-
spores (Fig. 1) [80?4?4] ( 8.0 ) 9 .0 - 12 .5 ( 15 .0 ) ×
(7 . 5) 8 .0 - 11 .0 ( 13.0 ) μm [ Q = ( 1 .0 ) 1.05 -
1 .25 ( 1 .29 ) , Q = 1 .14 ± 0.07 ] , subglobose to
broadly ellipsoid, occasionally globose . Pleurocystidia
(Fig. 2) 140 - 180×25 - 35μm, fusiform, often capi-
tate to subcapitate, thick-walled ( up to 6 μm) , the
apex always slightly thick-walled (ca . 2μmthick) and
with yellowish crystalline deposits . Pileipellis ( Fig. 3)
hymeniform, composed of broadly clavate, pyriform or
sphaeropedunculate cells . Pileocystidia ( Fig. 3 ) 70 -
270×9 - 17μm, erect, lanceolate with ventricose base
and tapering tips, thick-walled ( up to 4μm thick) .
Stipe trama (Fig. 4 ) monomitic, composed of parallel ,
sometimes branching, hyaline, colorless, thin-walled
hyphae connected with slender, colorless, thin-walled
and often short celled-hyphae 2 - 4μm diam . Stipiti-
pellis ( Fig. 5) composed of appressed, parallel , light
yellowish, thin-walled hyphae 3 - 8μm diam . Caulo-
cystidia ( Fig. 5) 60 - 240×12 - 20μm . Clamp con-
nections common .
Habitat—Mostly on buried rotten wood in broad-
leaved forest of Fagus and Quercus, etc .
3366 期 WANG Lan et al. : Synopsis and Systematic Reconsideration of Xerula s . str . (Agaricales)
Figs. 1 - 5 . Xerula pudens . 1 . Basidiospores; 2 . Hymeniumwith basidia and pleurocystidia, subhymenium and lamellar trama;
3 . Pileipellis and pileocystidia; 4 . Stipe strama; 5 . Stipitipellis and caulocystidia . All figures are from TENN 59330
Distribution—EUROPE: Austria, Belgium, Croat-
ia, Denmark, England, Estonia, Finland, France,
Germany, Greece, Hungary, Italy, the Netherlands,
Poland, Romania, Slovakia, southwestern Russia (Cau-
casus) , Sweden, and Switzerland ( Ronikier, 2005a) .
Materials examined—AUSTRIA: 28- IX-2001 ,
R . H . Petersen 11469 ( TENN 59330 ) . DENMARK :
28-VII-2002 , P.B . Hansen & S . Vesterholt 11469
(HKAS 45057) . GERMANY : 4-IX-1990 , R .E . Hal-
ling 6507 (NY 184 , HKAS 43811 ) . SWEDEN: 7-X-
436 云 南 植 物 研 究 30 卷
2005 , S . Lundell & J .A . Nannfeldt 1717 (NY 183) .
Notes: The name X. longipes ( Bull .) Maire has
frequently been used for this taxon, but according to
D?rfelt (1982 ) , the basionym of X. longipes, Agari-
cus longipes Bull . (1785 ) , is a homonym of Agaricus
longipes Scopoli (1772) , a nomen dubium of a fungus
froma mine . Thus, X. pudens is the correct name of
this species . Two varieties are recognized, X. pudens
var. pudens and X. pudens var. fusca ( Lucan ex
Quél .) D?rfelt . Thecollections cited above fit the con-
cept of X. pudens var. pudens (D?rfelt, 1980a) .
Xerula melanotricha D?rfelt, Feddes Repert . 90:
367 .1979 . ——— Oudemansiella melanotricha (D?rfelt)
M .M . Moser in Gams, Kryptogamenfl . 11b?2 , rev .
ed . 5 : 156 . 1983 . (Figs. 6 - 10)
Pileus dark brownor blackishbrown, withup to 3
mmlong and dark brown setae . Basidiospores ( Fig. 7)
[100?5?5] (8 .0) 9 .0 - 11 .0 ( 11 .5 ) × ( 7 . 0) 7 .5 -
10 .0 ( 10 .5 ) μm [ Q = ( 1 .00 ) 1 .05 - 1 .29 , Q =
1 .13±0 .07 ] , subglobose to broadly ellipsoid . Pleuro-
cystidia ( Fig. 6 ) fusiform, thin-walled, occasionally
subcapitate, the apex usuallywithout crystalline depo-
sits . Pileipellis ( Fig. 8 ) hymeniform, composed of
broadly clavate to sphaeropedunculate cells . Pileocys-
tidia (Fig. 8 ) up to 3 mm long ( but mostly 100 - 350
long) , 10 - 20 μm diam, thick-walled ( up to 5 μm
thick) . Stipe trama (Fig. 9 ) composedof parallel , oc-
casionally branching, hyaline, colorless, thin-walled
hyphae 2 - 15μm diam . Stipitipellis ( Fig. 10 ) com-
posed of appressed, parallel hyphae 3 - 8 μm diam .
Caulocystidia ( Fig. 10 ) 40 - 300×8 - 20μm, thick-
walled . Clamp connections common .
Habitat—Mostly on buried rotten wood in conifer
forest of Abies .
Distribution—EUROPE : Austria, Croatia, the
Czech Republic, France, Germany, Greece, Italy,
Poland, Romania, Slovakia, southwestern Russia
(Caucasus) , and Switzerland . NORTH AFRICA : Al-
geria (Contu, 2000; Ronikier, 2005b) .
Figs. 6 - 10 . Xerula melanotricha . 6 . Basidia and pleurocystidia; 7 . Basidiospores; 8 . Pileipellis and pileocystidia;
9 . Stipe strama; 10 . Stipitipellis and caulocystidia . All figures are from GMM 6994 ( F)
5366 期 WANG Lan et al. : Synopsis and Systematic Reconsideration of Xerula s . str . (Agaricales)
Materials examined—GERMANY : 1- IX-1986 ,
K . H . Rexer 1605 ( HKAS 49782 ) . RUSSIA : Cauca-
sus, 17- IX-2003 , G. M . Mueller 6994 , 7015 , 7030
and 71031 (F ) .
Notes: Xerula melanotricha can be confusedwith
X. pudens . For the separation of them see the key
above .
Xerula strigosa Zhu L . Yang, L . Wang et G .
M . Mueller, sp . nov . (Figs. 11 - 20)
MycoBank: MB 512372
Etymology: Named because of the thick-walled
setae on the surfaceof the pileus and the stipe .
Pileus 20 - 50 mm latus, convexus vel plano-con-
vexus, interdum sub-umbonatus vel depressus, flavo-
brunneus vel atro-brunneus, non-viscidus, pubescens;
pubes flavo-brunneae, acerosae . Caro tenuis, albida,
inodora . Lamellae sinuatae vel adnatae, subdistantes,
albidae; lamellulae praesenites . Stipes 50 - 80×3 - 6
mm, subcylindricus, ad basim incrassatus, ochra-
ceous, apicemversus pallidior, pubescens; pubes fla-
vo-brunneae; pseudorhiza praesentes . Basidiosporae
11 .0 - 15 .0×9 .0 - 11 .5μm, lato-ellipsoideae vel el-
lipsoideae, inamyloideae . Basidia 45 - 80× 10 - 18
μm, clavata, 4-sporigera . Pleurocystidia 120 - 150×
25 - 40μm, fusoidea, parietibus crassis ( 1 - 3μm) ,
capitata vel subcapitata . Cheilocystidia pleurocystidis
similis, subfusoidea, saepe capitata . Epicutis pilei ex
hyphis latoclavatis, subfusoideis, pyriformis sphaerope-
dunculatis composita . Pileocystidia lanceolata, 8 - 20
μm lata, usque ad 1 mm longa, sursum attenuata,
setacea, erecta, parietibus crassis ( 1 - 3μm) , fulvo-
brunnea . Caulocystidia pileocystidis similis . Fibulae
absentes vel raro . Holotypus: J . F . Liang 162 (HKAS
48778 ) , 1 August 2005 , alt . 3 200 m, Yulong Coun-
ty, Yunnan, China .
Basidiomata (Fig. 11) medium-sized . Pileus20 -
50 mm in diam, convex to plano-convex, sometimes
slightly umbonate or depressed over disc, yellowish
brown ( 5E7 + 5E8; Cinnamon Brown) to dark brown
(6F6 + 6F8; Natal Brown) to palegrayish brown (4B3
+ 5B3; Pale Drab-Gray) , unpolished, not viscid,
covered with erect and yellowish brown ( 5D7 + 5E7;
Cinnamon Brown) setae, the margin smooth to faintly
striate; context whitish ( 7A2 + 8A2; Pale Flesh Col-
or) , up to 4 mmthick at disc . Lamellae sinuate to ad-
nexed or adnate, subdistant, whitish ( 7A2 + 8A2;
Pale FleshColor) , up to 5 mmin height, with lamellu-
lae . Stipe 50 - 80×3 - 6 mm ( pseudorhiza excluded) ,
subcylindrical , slightly broader below, yellowish brown
(5D7 + 5E7; Cinnamon Brown) , paler towards the
apex, with yellowish brown ( 5D7 + 5E7; Cinnamon
Brown) setae; pseudorhiza up to 60 mm long, taper-
ing . Spore print white . Smell mild .
Basidiospores ( Figs. 16 , 19) [280?24?20] (9 .5)
11 .0 - 15 .0 (16 .5 ) × (8 . 5 ) 9 .0 - 11 .5 (15 .5 ) μm
[Q = ( 1 .03 ) 1 .10 - 1 .40 ( 1 .47 ) , Q = 1 .24 ±
0 .10] , broadly ellipsoid to ellipsoid, rarely subglo-
bose, colorless, hyaline, inamyloid, thin-walled . Ba-
sidia (Figs.15 , 18) 45 - 80×10 - 18μm, clavate, 4-
spored, rarely 2-spored; sterigmata5 - 7μmlong; bas-
al septa rarely clamped; basidioles fusiformtosubacer-
ose . Pleurocystidia (Figs. 15 , 17 , 20 ) scattered, 120
- 150 × 25 - 40 μm, fusiform, hyaline, thick-walled
( mostly 1 - 3 μm thick, occasionally up to 5 μm
thick) , often capitate to subcapitate; the apex always
thin-walled and with yellowish crystallinedeposits . La-
mellar edge fertile, with scattered cheilocystidia which
are similar to pleurocystidia in shape but somewhat
smaller in size . Subhymenium ( Fig. 15 ) tightly inter-
woven, composed of cylindrical cells 3 - 5μm diam .
Lamellar trama regular, composed of thin-walled fila-
mentous hyphae4 - 8μmdiam . Pileipellis (Fig. 14) 55
- 150 μm thick, hymeniform, composed of broadly
clavate, subfusiform, pyriform or sphaeropedunculate
cells (30 - 60×10 - 30μm) , thin-walled, often with
dark brown vacuolar pigment . Pileocystidia ( Fig. 14 )
up to 1 mm long, 8 - 20μm diam, erect, lanceolate
with ventricose base and tapering tips, thick-walled
(mostly 1 - 3μmthick, sometimes up to 5μmthick) ,
withyellowish brown cell wall . Pileal trama composed
of tightly interwoven, hyaline, colorless, thin-walled,
filamentous hyphae 6 - 15 μm diam . Stipe trama
(Fig. 12) monomitic, composed of parallel , sometimes
branching, hyaline, colorless, thin-walled hyphae ( 12
- 18μmdiam) connected with slender, colorless, thin-
636 云 南 植 物 研 究 30 卷
walled and often short celled-hyphae ca . 4μm diam .
Stipitipellis ( Fig. 13 ) composed of appressed, par-
allel , hyaline, light yellowish, slightly thin-walled
(ca . 2 μm) hyphae 4 - 10 μm diam . Caulocystidia
( Fig.13 ) similar to pileocystidia . Clamp connec-
tions rare .
Habitat—On buried rotten wood in conifer forest
of Pinus and subtropical broad-leaved forests of Litho-
carpus and Quercus, etc .
Known distribution—Central and northern Yun-
nan Province and southern Sichuan Province, south-
western China at 1 500 - 3 200 melevation .
Figs. 11 - 20 . Xerula strigosa . 11 . Basidioma; 12 . Stipe strama; 13 . Stipitipellis and caulocystidia; 14 . Pileipellis and pileocystidia;
15 . Hymeniumwith basidia and a pleurocystidium; 16 . Basidiospores; 17 . Pleurocystidia; 18 . Basidia at different stages of development;
19 . Basidiospores; 20 . Pleurocystidia . Figures 11 - 17 are from the holotype, while figures 18 - 20 from HKAS 42657
7366 期 WANG Lan et al. : Synopsis and Systematic Reconsideration of Xerula s . str . (Agaricales)
Materials examined—CHINA : Yunnan Prov .:
Binchuan County, Jizu Mountain, 8-VIII-1985 , G. P .
Xiao 424 ( HKAS 15402 , as X. pudens in Yang and
Zang, 1993; Ying and Zang, 1994 ) ; 16-VII-2003 ,
L . F . Zhang 23 and 25 ( HKAS 38229 and 38231 re-
spectively) . Chuxiong, Wuding County, Shizi Moun-
tain, 14-VIII-2003 , L . F . Zhang280 (HKAS 42557) .
Kunming, VIII-2006 , K . Hosaka CH04-274b (HKAS
52590 ) ; 26-VIII-2004 , L . Wang 398 and 399 (HKAS
46276 and 43467 respectively ) ; 27-VIII-2007 , L .
Wang 521 ( HKAS 52157 ) ; 25-VII-2007 , L . Wang
522 (HKAS 52357 ) ; 12-VIII-1990 , Z.L . Yang 1047
( HKAS 22758 , as X. pudens in Yang and Zang,
1993; Ying and Zang, 1994 ) ; 17-VII-2005 , Z.L .
Yang 4474 ( HKAS 48326 ) ; 8-IX-2007 , Z.L . Yang
4954 ( HKAS 52271 ) ; X-1980 , S . Zeng ( HKAS
7015 , as X. pudens in Yang and Zang, 1993; Ying
and Zang, 1994 ) ; 30-VI-2003 , L . F . Zhang 178
( HKAS 42440 ) ; 31-VIII-2003 , L . F . Zhang 331
( HKAS 42657 ) . Lijiang, IX-X-1916 , anonym s.n .
(WU 12869 , as Collybia longipes in Lohwag, 1937 ) .
Qujing, 14-VIII-1999 , Z.L . Yang 2618 ( HKAS
34061 ) . Shangri-La County, 17-VIII-2000 , Z.L .
Yang 4524 (HKAS 36615 ) . Songming County, Baiyi
Town, 22-VII-1998 , X . H . Wang 411 ( HKAS
32069a) . Yulong County, Laojun Mountain, 1-VIII-
2005 , J . F . Liang 162 ( HKAS 48778 , holotype) . Si-
chuan Prov ., Xichang, 19-IX-1999 , Z.L . Yang 2670
(HKAS 34136) .
Notes: Xerula strigosa is characterized by its
broadly ellipsoid to ellipsoid basidiospores, fusiform
and thick-walled pleurocystidiawith capitateor subcap-
itate apex oftenwith yellowish crystalline deposits, and
the rarity of clamp connections .
Xerula strigosa closely resembles the European
X. pudens, and, thus, was incorrectly regarded as the
latter in the last century ( Lohwag, 1937; Yang and
Zang, 1993; Ying and Zang, 1994 ) , although Horak
(1987 ) indicated that the material WU 12869 differs
from European X. pudens by“much larger spores and
cystidia of different shape”. Our study revealed that
unlike X. pudens, X. strigosa exhibits longer basidio-
spores, a rare presence of clamp connections on the
septaof basidia and hyphae in lamellar trama, and a
thin-walled apex of pleurocystidia .
Sharing yellowish brown setae on yellowish brown
to dark brown pileus and rarity of clamp connections
with X. sinopudens, which has been found in Japan,
Indonesia, Papua NewGuinea, and China, X. strigosa
also strongly resembles the latter . Due to their morpho-
logical similarity and overlap in distribution range
( e. g . in Kunming) , they can be easily confusedwith
each other . When immature, the basidiospores of
X. strigosa are shorter and smaller than in mature
specimens, and the pleurocystidiamay only be slightly
thick-walled, which reflect the characters of X. si-
nopudens ( see the key of this paper) . Our molecular
phylogenetic analyses (datanot shown) support treating
both a distinct species . Ecologically X. strigosa mainly
occurs in temperate regions or occasionally in subtropi-
cal areas, while X. sinopudens prefers warmer habi-
tats, and is distributed in the south at lower altitudes
(Yang and Zang, 2003) .
Xerula strigosa may also be confused with X. his-
pida, originally described from Central America . One
example is that Mueller et al. ( 2001 ) reported a
“ X. hispida”collection ( F 1129046 ) from Yunnan
Provinceof China . Our molecular phylogenetic studies
(data not shown) revealed that this is a collection of
X. strigosa . Morphologically, X. hispida differs from
X. strigosa by its 2-sterigmate ( rarely 4 ) basidia and
slender fusiformpleurocystidiawithout a capitate apex .
Xerula sinopudens R .H . Petersen et Nagasa .,
Rep . Tottori Mycol . Inst . 43 : 41 . 2006 . Figs.21 - 30
Pileus yellowish brown or brownish, covered with
yellowish brown setae . Basidiospores ( Figs. 21 , 27 ,
30) [ 250?25?25] (10 .5) 11 .0 - 13 .0 × (9 . 0) 9 .5 -
11 .5μm [Q = (1 .00) 1 .05 - 1 .20 (1 .28 ) , Q = 1 .12
±0 .07 ] , subglobose to broadly ellipsoid, sometimes
globose . Pleurocystidia ( Figs. 22 , 26 , 29 ) broadly
fusiform, thin-walled ( up to 0 .5 μm) , occasionally
subcapitate, the apex without crystalline deposits .
Stipe trama ( Fig. 23 ) monomitic, composed of paral-
lel , branching, hyaline, colorless, thin-walled hyphae
connected with slender, colorless, thin-walled and of-
836 云 南 植 物 研 究 30 卷
ten short celled-hyphae 2 - 5 μm diam . Stipitipellis
(Fig. 24 ) composed of appressed, parallel , light yel-
lowish hyphae 3 - 10μmdiam . Caulocystidia (Fig. 24)
similar with pileocystidia . Clamp connections rare .
Habitat—Mostly on buried rotten wood in tropical
and subtropical broad-leaved forests of Lithocarpus,
occasionally in conifer forests .
Distribution—ASIA : Indonesia, Japan, and
Papua New Guinea ( Petersen and Nagasawa, 2006 ) .
New to China .
Materials examined—CHINA : Hunan Prov .,
Mangshan, 2-IX-2007 , P . Zhang 644 (HKAS 52715) .
Yunnan Prov ., Baoshan, 22-VII-2003 , L .F . Zhang
219 (HKAS 42502) ; 25-VII-2003 , L .F . Zhang 255
Figs. 21 - 30 . Xerula sinopudens . 21 . Basidiospores; 22 . Hymeniumwith basidia and apleurocystidium; 23 . Stipe strama;
24 . Stipitipellis and caulocystidia; 25 . Pileipellis and pileocystidia; 26 . Pleurocystidia; 27 . Basidiospores; 28 . Basidia at
different stages of development; 29 . Pleurocystidia; 30 . Basidiospores . Figures 21 - 25 are from HKAS 43304 ,
while figures 26 - 27 and figures 28 - 30 are from HKAS 42101 and E 186523 respectively
9366 期 WANG Lan et al. : Synopsis and Systematic Reconsideration of Xerula s . str . (Agaricales)
(HKAS 42583) . Gaoligong Mountain, 11-VIII-1973 ,
X . J . Li 615A ( HKAS 3505A , as X. pudens in Yang
and Zang, 1993; Ying and Zang, 1994 ) . Jingdong
County, Ailao Mountain, 20-VII-2006 , Y .C . Li 607
(HKAS 50361 ) . J inghong, Dadugang, 30-VIII-2004 ,
X . H .Wang1795 (HKAS 5899) . Kunming, VIII-2006 ,
K . Hosaka CH04-274a ( HKAS 52589 ) ; 1- IX-2007 ,
Z.L . Yang 4948 ( HKAS 52265 ) . Longling County,
5-IX-2002 , Z.L . Yang 3459 ( HKAS 41528 ) ; 9-IX-
2002 , Z.L . Yang 3514 ( HKAS 42098) ; 10- IX-2002 ,
Z.L . Yang 3542 ( HKAS 42101) ; 11- IX-2002 , Z.L .
Yang 3560 ( HKAS 42099) ; 12-IX-2002 , Z.L . Yang
3585 ( HKAS 42093 ) . Luxi County, 11-VIII-1980 ,
M . Zang 6552 ( HKAS 6552 , as X. pudens in Yang
and Zang, 1993; Ying and Zang, 1994 ) . Tengchong
County, 20-VII-2003 , L . Wang 209 ( HKAS 43304) ;
27-VIII-2007 , Z.L . Yang 3835 ( HKAS 42988 ) .
Yingjiang County, 13-VII-2003 , Z.L . Yang 3641
( HKAS 42782 ) ; 14-VII-2003 , Z.L . Yang 3669
(HKAS 42805 ) . INDONESIA : North Borneo, Kin-
abalu, 19-I-1964 , E. J . H . Corner RSNB 5014A ( E
186519 , as X. longipes in Corner, 1996 ) ; 25- I-1964 ,
E . J . H . Corner RSNB 5014B ( E 186520 , as X. lon-
gipes in Corner, 1996) ; 10- III-1964 , E. J . H . Corner
RSNB 5014C ( E 186521 , as X. longipes in Corner,
1996) ; 9-IV-1964 , E. J . H . Corner RSNB 5014D ( E
186522 , as X. longipes in Corner, 1996 ) ; 20-IV-
1964 , E. J . H . Corner RSNB 5014E ( E 186523, as
X. longipes inCorner 1996 ) . JAPAN: Tottori , 26-VI-
II-1987 , E . Nagasawa 87 - 178 (TMI 13445 ) ; 22-VI-
1995 , E . Nagasawa (TMI 19467) .
Notes: X. sinopudens was regarded X. pudens in
China as well as in other regions of East Asia ( Yang
and Zang, 1993; Ying and Zang, 1994 ) . However,
the European X. pudens has common clamp conne-
ctions, smaller basidiospores measuring 9 .0 - 12 .5×
8 .0 - 11 .0μm, and much thicker-walled ( up to 6μm)
pleurocystidia . We agree with Petersen and Nagasawa
(2006) that Corner′s collections of X. longipes should
be regarded as X. sinopudens because of the rarity of
clamp connections, thin-walled pleurocystidia with a
non-capitate apex and without crystalline deposits
(Figs.28 - 30) .
Bi et al. ( 1997) cited two collections under the
name Oudemansiella longipes . Reexamination revealed
that both of them are neither X. strigosa nor X. si-
nopudens but members of O. radicata var. furfuracea
complex .
Xerula setulosa ( Murrill ) R . H . Petersen et
T. J . Baroni , Mycotaxon 101: 114 . 2007 . ——— Gym-
nopus setulosusMurrill , North American Flora 9: 373 .
1916 . ——— Collybia setulosa (Murrill ) Murrill , Myco-
logia 8: 219 . 1916 . ——— Marasmius setulosus ( Mur-
rill ) Singer, Lilloa 22: 326 . 1951 ( non M. setulosus
Murrill , Bull . Torrey Bot . Club 67: 150 . 1940 ) .
——— Marasmius murrillianus Singer, Lilloa 25: 488 .
1952 . ——— Lentinus pilosus Rick, Lilloa 2: 310 .
1938 [ non Lentinuspilosus (Fr .) Fr ., Epicrisis Syst .
Mycol . 395 . 1838] . ——— Xerula pilosa ( Rick) Sing-
er, Lilloa 26: 86 . 1953 . ——— Oudemansiella pilosa
( Rick) Singer, Sydowia 15: 59 . 1962 . Figs. 31 - 36
Basidiomata miniature, brownish to dark brown,
covered with yellowish brown setae . Pileus margin often
with radially arranged striations .Basidiospores (Fig. 31)
[40?2?2] (10 .5) 11 .0 - 16 .0 (16 .5 )× ( 8 . 5) 9 .5 -
12 .0 ( 13 .0 ) μm [Q = ( 1 .04 ) 1 .09 - 1 .50 , Q =
1 .20±0 .11 ] , subgloboseor broadly ellipsoid to ellip-
soid . Basidia (Fig. 32 ) 4- or a mixture of 2- and 4-
spored . Pleurocystidia ( Figs. 32 - 33 ) slender fusi-
form, thick-walled ( 2 - 5μm) , occasionally subcapi-
tate, the apex always thin-walled and with yellowish
crystalline deposits . Stipe trama (Fig.35) monomitic,
composed of parallel , branching, hyaline, colorless,
thin-walled hyphaeconnectedwith slender, short celled-
hyphae ca . 2 - 5μm diam . Stipitipellis (Fig. 36 ) com-
posed of appressed, parallel hyphae 3 - 7 μm diam .
Caulocystidia ( Fig. 36 ) similar with pileocystidia .
Clamp connections rare .
Habitat—Under broad-leaved trees such as
Quercus and Coccoloba, etc . ( Petersen and Baroni ,
2007) .
Distribution—CENTRAL AND NORTHERN SO-
UTH AMERICA : Belize, Brazil ( Rio GrandedoSul ) ,
Jamaica, Mexico, US ( Puerto Rico) (Singer, 1964;
Petersen and Baroni , 2007) . New to Costa Rica .
046 云 南 植 物 研 究 30 卷
Figs. 31 - 36 . Xerula setulosa . 31 . Basidiospores; 32 . Hymeniumwith basidia and pleurocystidia; 33 . Pleurocystidia;
34 . Pileipellis and pileocystidia; 35 . Stipe strama; 36 . Stipitipellis and caulocystidia . All figures fromNavarro 1216
Materials examined—COSTA RICA : 27-VI-
2001 , E . Navarro 1216 ( INB) ; I . Lópe1472 ( INB) .
Notes: Petersen and Baroni ( 2007 ) suggested
that X. hispida may bea 2-sporedstateof X. setulosa .
Our molecular phylogenetic analyses ( data not shown)
showed that the two collections cited abovedo not clus-
tered with the holotype of X. hispida .
Discussion
Taxonomy and Systematics of Xerula s . str .
In study of the evolutionary relationships of My-
caureola diseaeMaire& Chemin, Binder et al. (2006)
provided a multigene molecular phylogenetic study of
the family Physalacriaceae and showed that“ O. ca-
narii”and O. mucida formed a well supported clade
1466 期 WANG Lan et al. : Synopsis and Systematic Reconsideration of Xerula s . str . (Agaricales)
with X. furfuracea ( Peck) Redhead et al . [ ≡ O. ra-
dicata var. furfuracea ( Peck ) Pegler ] and X. me-
galospora ( Clem .) Redhead, Ginns & Shoemaker ( fig-
ure 3 of Binder et al. , 2006 ) . Xerula pudens and
closely related taxa were not included in this analysis .
Interestingly, in the same publication, they presented
another analysis of a densely sampled ITS data set that
included X. pudens and X. hispida . These twospecies
formed a separated clade from X. furfuracea and“ O. -
canarii”although they all werelocated in thePhysalac-
riaceae clade ( figure 4 of Binder et al. , 2006 ) . Our
molecular phylogenetic analysis as inferred from ITS
and nLSU sequences also shows that Xerula s . str .
and Oudemansiella are independent lineages and
should be treated as different genera ( Zhang et al. ,
2003; unpublished data of Zhang) .
In this study, Xerula is proposed as adistinct ge-
nus in its strict sense, including X. pudens and its
close allies, viz . Xerula sect . Xerula in the sense of
D?rfelt (1984) and Contu ( 2000 ) . Species of Xerula
sect . Radicatae ( Clémen?on) D?rfelt should be ex-
cluded from Xerula and incorporated in thegenus Ou-
demansiella . Species with collybioid basidiomatawith a
dry pileus and apseudorhiza, and erect, thick-walled,
lanceolate, yellowish or brownish pilei- and caulocys-
tidia, subacerose basidioles and a fertile lamellar edge
with scattered cheilocystidia belong to Xerula, while
those with a gelatinous or viscid pileus and thin- to
slightly thick-walled and often collapsed, nearly color-
less, non-lanceolate pilei- and caulocystidia or without
dermatocystidia, without subacerose basidioles and a
sterile lamellar edge with crowded cheilocystidia are
Oudemansiella . Cheilocystidia are present in Xerula
but just scattered among thebasidiaof the fertile edge,
which were called marginal pleurocystidia by Corner
(1996 ) . The presence or absence of clamp connec-
tions, the form and the thickness of the wall of pleuro-
cystidia, and the size and the shape of basidiospores
are probably the best parameters to characterize the
species of Xerula effectively .
Species of X. sect . Albotomentosae ( Clémen?on)
D?rfelt and sect . Hyalosetae D?rfelt, such as X. ame-
ricana D?rfelt, X. caussei Maire and X. hongoi
D?rfelt, would seem to fit the genus Xerula . Until
more dataon these taxa are available, however, formal
inclusion is postponed . For example, X. hongoi , pos-
sesses a dry pileus, yellowish brown, thick-walled and
lanceolate pilei- and caulocystidia and, thus, might be
place in Xerula s . str . However, its marasmioid ba-
sidiomata with a short pseudorhiza share common fea-
tures with members of Rhizomarasmius R. H . Petersen
(Petersen, 2000 ) . In addition, Xerula mediterranea
(Pacioni et Lalli ) Quadr . et Lunghini , on which Xer-
ula subgen . Ixoflammula was introduced ( Contu,
2000) , might also belong to Rhizomarasmius according
to Petersen ( 2000) .
Biogeography of Xerula s . str .
It was proposed that the European X. pudens and
the American X. hispida occur in East Asia ( e. g .:
Lohwag, 1937; Imazeki and Hongo, 1965; D?rfelt,
1980a; Yang and Zang, 1993; Corner, 1996; Mueller
et al. , 2001 ) . Extensive and detailed studies have
now shown that such putative Eurasian disjunct distri-
butions of X. pudens, and eastern Asia and Central
America?northern South America disjunctions of
X. hispida are not well supported . Thus far, mushro-
oms of the genus Xerula s . str . can be found in many
parts of theworld . However, each species has a relative-
ly limited distribution . To date, the total number of re-
cognized Xerula species is six .Two of them, X. pudens
and X. melanotricha, are distributed in Europe, two,
X. sinopudens and X. strigosa, occur in East Asia and
Southeast Asia, and two species have been found in
Central and South America .
Species of Xerula from Africa and Oceania are
poorly known . Xerula melanotricha was reported from
Algeria, North Africa (D?rfelt, 1979 , 1980a) . Xerula
pudens ( under the name of Oudemansiella longipes)
was recorded from New Zealand (Segedin and Penny-
cook, 2001) , but its occurrence thereneeds to bever-
ified . Information on the diversity of species and their
distributions onboth continents will be useful for better
understanding thegeography of Xerula s . str .
Acknowledgements: The authors are very grateful to Dr . R .
Halling (New York, USA) , Prof . G . Kost (Marburg, Germa-
ny) , Prof . T. H . Li (Guangzhou, China) , Dr . Y . Mizuta (Ko-
246 云 南 植 物 研 究 30 卷
chi , Japan) , Mr . E . Nagasawa (Tottori , Japan) , Prof . R. H .
Petersen (Knoxville, USA) , Dr . K .H . Rexer (Marburg, Ger-
many) and Dr . S . Takehashi ( Hokkaido, Japan) for loaning
collections and assistance in providing some literature .
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书讯
新 书 介 绍
Gifts fromthe Gardens of China———The Introduction of Traditional
Chinese Garden Plants to Britain 1698 - 1862; Jane Kilpatrick, 288 p, 2007; Lon-
don: Frances Lincoln Ltd .
本书详细记载了清代英国从中国采集与引种植物的历史 (包括业余爱好
者和专业采集家 ) , 还有对当时中国的文化、社会考察与交流等情况。作者
是英国剑桥大学出身的历史学者 , 不但收集到非常珍贵的详细资料 , 而且还
是一个植物爱好者 , 并多次到中国进行野外实地考察。该书不仅记载了英国
从中国引种植物的历史 , 还从另一个侧面向读者展示了当代英国植物爱好者
对中国的园林与植物的酷爱程度 , 同时也包括对当年英法联军火烧圆明园的
痛恨。该书书末附有在华的英国植物采集人员名单与采集情况简介。
马金双
(美国纽约布鲁克林植物园 )
446 云 南 植 物 研 究 30 卷