蜜环菌生物种及鉴定方法研究进展
门金鑫, 邢晓科, 郭顺星

Biological species and identification methods of the genus Armillaria (Agaricales, Basidiomycota): a review
Jin-Xin MEN, Xiao-Ke XING, Shun-Xing GUO
表1 已报道的蜜环菌生物种
Table 1 The reported biological species of Armillaria
编号
No.
生物种
Biological species
形态特征
Morphological characteristics
地理分布
Geographical distribution
参考文献
References
菌环
Annulus
配合型
Mating types
菌盖
Pileus
担子
Basidia
担孢子
Basidiospore
根状菌索
Rhizomorph
1 蜜环菌
A. mellea (Vahl) P. Kumm. [=Agaricus melleus Vahl]
+ 异宗配合
Heterothallism
直径长达
90mm,从凸很快扩展到平凸,甚至变平
Up to 90mm, convex soon expanded to become plano-convex or even plane.
四孢型,没有锁状联合
Four-spored, without clamp connection.
2-8.9×5.6-6.7μm,宽椭圆形,有小水滴状斑点,表面光滑
2-8.9×5.6-6.7μm, broadly ellipsoid, guttulate, apparently smooth.
带状,二叉分枝
Belt-shaped, dichotomous.
欧洲,北美,非洲,中国,朝鲜,伊朗
Europe, North America, Africa, China, Korea, Iran
Ullrich & Anderson 1987; Watling et al. 1982
2 高卢蜜环菌
A. gallica Marxm. & Romagn. [=A. bulbosa (Barla) Kile & Watling=A. luten Gillet]
+ 异宗配合
Heterothallism
直径
1.8-5.5cm,从凸到平凸,最后变成平的中间略凹
1.8-5.5cm diam., convex then plano-convex and finally plane to slightly umbilicate.
四孢型,棍棒状,有锁状联合
Four-spored, clavate, with a clamp connection.
3.8-5×7-10.3μm,全部为白色,宽椭圆形到卵形,光滑,透明
3.8-5×7-10.3μm, spores ivory in mass, broadly ellipitical to ovate, smooth, hyaline.
圆柱状,顶端尖锐,单轴分枝,黑棕色到黑色
Cylindrical, and sharp tips with monopodial branches, dark brown to black.
欧洲,北美,非洲,日本,中国,朝鲜,伊朗
Europe, North America, Africa, Japan, China, Korea, Iran
Cha et al. 1992
待续
表1
3 奥氏蜜环菌
A. ostoyae (Romagn.) Herink [= A. solidipes Peck]
+ 异宗配合
Heterothallism
直径4-14cm,初时半球钟形或钝的圆锥形,之后平凸,最后变平
4-14cm diam., hemispherical- companulate or obtusely conic, then plano-convex and finally plane.
四孢型,棍棒状,多数有锁状联合
Four-spored, clavate, usually with a clamp connection.
5.2-7.5×7.5- 12.5μm,全部为白色,宽椭圆形到卵圆形,光滑,透明,有一个明显的细尖5.2-7.5×7.5- 12.5μm, spores ivory in mass, broadly elliptical to ovate, smooth, hyaline, with a distinct apiculus. 扁平,条状,黑棕色,初时白色带有二叉分枝
Flat and belt-shaped in shade of dark brown and white at first with dichotomous branches.
欧洲,北美,日本,中国,朝鲜
Europe, North America, Japan, China, Korea
Bérubé & Dessureault 1988; Cha et al. 1992
4 头柄蜜环菌
A. cepistipes Velen.
+ 异宗配合
Heterothallism
菌盖直径
20-100mm,边缘内卷,之后变直,内弯
20-100mm diam., margin involute, then straight, inflexed.
四孢型,狭的棍棒状,有锁状联合
Four-spored, rather narrowly clavate, with a clamp connection.
7.0-9.0(10)×4.5 -6.5(7.0)µm,多数宽椭圆形到近
球形
7.0-9.0(10)×4.5- 6.5(7.0)µm, mostly broadly ellipsoid to subglobose.
横切面为典型的椭圆形
Rhizomorphs with a typical oval cross-section.
欧洲,北美,日本,伊朗
Europe, North America, Japan, Iran
Antonín et al. 2009
5 北方蜜环菌
A. borealis Maxm. & Korhonen
+ 异宗配合
Heterothallism
直径3-10cm,初期半球形至扁半球形,后期扁平,四周鳞片稀少
3-10cm diam., hemispheric to flat hemispheric, then flat, few scales around.
四孢型,基部有锁状联合
Four-spored, with a clamp connection.
6-9×4.5-5μm,宽椭圆形,光滑,透明
6-9×4.5-5μm, broadly ellipsoid, smooth, hyaline.
栗棕色
Maroon brown.
欧洲,中国,伊朗
Europe, China, Iran
Korhonen 1978
待续
表1
6 假蜜环菌
A. tabescens (Scop.) Emel
同宗配合
Homothallism
直径2.8-8.5cm,幼时扁半球形,后渐平展,中部色深并有纤毛状小鳞片
2.8-8.5cm diam., flat hemispheric when young, later gradually flat. Central dark with fibrillous scales.
棍棒状,有锁状 联合
Clavate, with a clamp connection.
(6.0)6.5-8.9 (8.5)×(3.5)4.0-5.5 (6.0)µm,椭圆形至长椭圆形,光滑,透明
(6.0)6.5-8.9 (8.5)µm×(3.5)4.0-5.5 (6.0)µm,
ellipsoid to oblong, smooth, hyaline.
不致病
Nonpathogenic
欧洲,北美,日本,中国
Europe, North America, Japan, China
Schnabel et al. 2005; Marxmuller 1992
7 棘皮蜜环菌
A. ectypa (Fr.) Lamoure [=Agaricus ectypus Fr. =A. ectypa (Fr.) Herink= Armillariella ectypa (Fr.) Singer= Clitocybe ectypa (Fr.) Gillet= Omphalia ectypa (Fr.) Quél.]
同宗配合
Homothallism
直径
25-60(100)mm,凸到平凸,之后扁平,中央有深棕色小鳞片
25-60(100)mm diam., convex to plano-convex, later explanate with dark brown scales at center.
四孢型,棍棒状
Four-spored, clavate.
(6.5)7.0-8.5(9.0)×(5.0)5.5- 6.5(7.0)µm,宽椭圆形到椭圆形
(6.5)7.0-8.5(9.0)×(5.0)5.5-6.5 (7.0)µm, broadly ellipsoid to ellipsoid.
未见报道
Undescribed
欧洲Europe Stasińska 2015
待续
表1
8 双生蜜环菌
A. gemina Bérubé & Dessur.
+ 异宗配合
Heterothallism
直径2-10cm,半球-钟状或钝的抛物线形,之后变凸,最后变平
2-10cm diam., hemispherical- campanulate or obtusely-parabolic, then convex and finally plane.
四孢型,棍棒状,部分有锁状联合
Four-spored, clavate, some with a clamp connection.
8.2-10×5.2-7μm,全部白色,宽椭圆形到卵圆形,光滑,顶尖
8.2-10×5.2-7μm, spores ivory in mass, broadly elliptical to ovate, smooth, apiculate.
圆柱形,单轴分枝,顶端尖锐
Cylindrical with monopodial branches and sharp tips.
北美
North America
Bérubé & Dessureault 1989
9 光盖蜜环菌
A. calvescens Bérubé & Dessur.
+ 异宗配合
Heterothallism
直径2-10cm,球状,凸到
平凸
2-10cm diam., globose, convex then plano-convex.
四孢型,棍棒状,部分有锁状联合
Four-spored, clavate, some with a clamp connection.
8.5-10×5-7μm,全部白色,宽椭圆形到卵圆形,光滑,顶尖
8.5-10×5-7μm, spores ivory in mass, broadly elliptical to ovate, smooth, apiculate.
管状,有锐利的尖端,单轴
分枝
Tubular with sharp tips and monopodial branches.
北美
North America
Bérubé & Dessureault 1989
10 芥黄蜜环菌
A. sinapina Bérubé & Dessur.
+ 异宗配合
Heterothallism
直径2-6cm,圆锥-钟形到钟形,凸到平凸
2-6cm diam., conical- campanulate to campanulate, convex then plano-convex.
四孢型,棍棒状,部分有锁状联合
Four-spored, clavate, some with a clamp connection.
5.9-8×8.2-10μm,全部白色,宽椭圆形到卵形,光滑,顶尖
5.9-8×8.2-10μm, spores ivory in mass, broadly elliptical to ovate, smooth, apiculate.
管状带有锋利的尖,单轴分枝,浅棕色到深棕色
Tubular with sharp tips and monopodial, light brown to dark brown.
北美,日本,中国
North America, Japan, China
Bérubé & Dessureaul 1988
待续
表1
11 第九蜜环菌
A. nabsnona T.J. Volk & Burds.
+ 异宗配合
Heterothallism
直径4-7cm,初时凸状,后来
变平
4-7cm diam., at first convex, later plane.
四孢型,棍棒状,有锁状联合
Four-spored, clavate, with a clamp connection.
(6-)8-10×5.5-6.5µm,全部白色,卵圆形到近球形
(6-)8-10×5.5 -6.5µm, white in mass; ovoid to subglobose.
黑色,有分枝
Black, branched.
北美,日本
North America, Japan
Volk et al. 1996
12 A. altimontana Brazee, B. Ortiz, Banik & D.L. Lindner + 异宗配合
Heterothallism
直径可达3cm,平状,带有
鳞片
Up to 3cm diam., flat, with scales.
四孢型,圆的棍棒状,无锁状联合
Four-spored, cylindro-clavate, without clamp connection.
7.2-11.7×5.4 -6.3μm,宽椭圆形,光滑
7.2-11.7×5.4 -6.3μm, broadly elliptical, smooth.
单轴分枝,薄,与基部连接处圆柱状
Monopodial, and thin, cylindrical on attached substrates.
北美
North America
Brazee et al. 2012
13 A. heimii Pegler [=Clitocybe elegans R. Heim] 异宗配合
Heterothallism
4.5-5.2×3
-6.5mm,中央扁平,边缘弯曲
4.5-5.2×3-6.5mm diam., depressed at the centre, margin incurved.
四孢型,棍棒状,常为薄壁
Four-spored, clavate, generally thin-walled.
7.2-9×(4.4)5
-5.5μm,倒卵球形到有尖角的,光滑到稍不规则
7.2-9×(4.4)5-
5.5μm, obovoid to angular, smooth to very faintly irregular.
未见报道
Undescribed.
非洲
Africa
Kile & Watling 1988;
Pérez-Sierra et al. 2004;
Otieno & Termorshuizen 2003
14 A. fuscipes Petch 同宗配合
Homothallism
直径可达51mm(平均35mm),凸状,之后变平
Up to 51mm (average 35mm) diam., convex later broadly plane.
四孢型,棍棒状,薄壁,透明,担子底部无锁状联合
Four-spored, clavate, thin walled, hyaline, without clamp connection.
7-12(9)×4-7(6)μm,卵圆形到椭圆形,顶部细尖,透明
7-12(9)×4-7(6)μm, elliptic-ovoid to elliptic-oblong, apiculated, hyaline.
微红色到黑色,直径2-3mm
Reddish then black, 2-3mm diam.
非洲
Africa
Coetaee et al. 2000
待续
表1
15 A. mellea subsp. Nipponica J.Y. Cha & lgarashi [=A. mellea s. str.] + 同宗配合
Homothallism
直径1.6-7.5cm,幼时突的圆锥状到凸状,之后平凸,最后成平
1.6-7.5cm in diam., abruptly conic to convex when young, planoconvex, finally plane.
四孢型,棍棒状,基部无锁状联合
Four-spored, clavate, without clamp connection at the base.
10-13.5×6-8μm,全部白色,近球形,宽椭圆形
10-13.5×6-8μm, white in mass, subglobose, broadly elliptic.
带状,二叉分枝,有钝的尖端
Belt-shaped, dichotomous with obtuse tips.
非洲,日本
Africa, Japan
Cha & Igarashi 1994, 1995a
16 A. fellea (Hongo) Kile & Watling [=A. fellea Hongo] + 未见报道
Unknown
边缘有棕色的非常小的鳞片
Very small brown scales at margin.
四孢型,棍棒状,很少有锁状联合
Four-spored, clavate, rarely with a clamp connection.
7.2-8.3(-8.8)×(4.4)-5.5μm,椭圆形到宽椭圆形,表面光滑
7.2-8.3(-8.8)×(4.4)-5.5μm, ellipsoid to broadly ellipsoid, apparently smooth.
未见报道
Undescribed
巴布亚新几内亚
Papua New Guinea
Kile & Watling 1988
17 A. fumosa Kile & Watling 未见报道
Unknown
未见报道
Unknown
直径20-120mm,从凸扩展到平凸;边缘最初向内弯曲
20-120mm diam., convex expanding to plano-convex; margin initially incurved.
四孢型偶尔双孢型,棍棒状,无锁状联合
Four-spored occasionally two-spored, clavate, without clamp connection.
6.5-8.5(-9.5)×(4-) 4.5-6(-6.5)μm,幼时几乎全部白色,长椭圆形带有宽的突出的细尖
6.5-8.5(-9.5)×(4-) 4.5-6(-6.5)μm, almost white in mass when fresh, elongate-ellipsoid with abroad prominent apiculus.
扁平
Flat
澳大利亚
Australia
Kile & Watling 1983
待续
表1
18 A. limonea (G. Stev.) Boesew. + 未见报道
Unknown
带有棕黄色的小
鳞片
With claybank scales.
未见报道
Undescribed
6.5-9×3.5-5.0μm,有些细微的粗糙不平,成熟后稍厚
6.5-9×3.5-5.0μm, slightly rough, slightly thick when mature.
分支多
Branching is prolific.
新西兰
New Zealand
Singer 1969; Shaw et al. 1981
19 A. luteobubalina Watling & Kile + 异宗配合
Heterothallism
直径
40-100(-150)mm,初时平凸,之后展开,具有微凸起到凸起,中央有小鳞片
40-100(-150)mm diam., at first convex finally becoming expanded and subumbonate to umbonate, centre squamulose.
四孢型(有些双孢型),基部无锁状联合
Four-spored (some two-spored), without clamp connection at the base.
(5-)6.5-7.5(-8)× (4-)4.5-5.5(-6)mm,宽椭圆形,带着明显的宽的细尖,光滑
(5-)6.5-7.5(-8)× (4-)4.5-5.5(-6)mm, broadly ellipsoid with distinct broad apiculus, smooth.
圆柱形到扁平,分支较少
Cylindrical to flattened, sparsely-
branched.
澳大利亚
Australia
Kile 1981; Kile et al. 1983; Podger et al. 1978
20 A. novae-zelandiae (G. Stev.) Herink [= Armillariella sparrei Singer var. elaeodes Singer] + 未见报道
Unknown
直径
30-100(-150)mm,稍有凸起到凸起,之后变成平凸,最后多变为扁平
30-100(-150)mm diam., subumbonate to umbonate becoming plano-
convex and later often depressed.
四孢型,偶尔双孢型,棍棒状,基部无锁状联合
Four-spored occasionally two-spored, clavate, without clamp connection at the base.
7-8(-8.5)×4.5-5.0 (-5.5)μm,椭圆形到长椭圆形
7-8(-8.5)×4.5-5.0 (-5.5)μm, ellipsoid, to elongate-ellipsoid.
圆柱状,二叉分枝,光滑、窄的菌丝鞘
Cylindrical, with dichotomous branches at their tips and smooth, narrow, mycelial sheaths.
澳大利亚,新西兰,巴布亚新几内亚,南美洲
Australia, New Zealand, Papua New Guinea, South America
Kile & Watling 1983; Guillaumin et al. 1992; Shaw et al. 1981
待续
表1
21 A. pallidula Kile & Watling 未见报道
Unknown
未见报道
Unknown
直径
45-90mm,钟形,边缘内卷,之后凸起,扩展到平凸
45-90mm diam., campanulate with inrolled margin, then convex to subumbonate expanding to plano-convex.
四孢型,偶尔双孢型,细长的棍棒状,很少有锁状联合
Four-spored, occasionally two-spored, elongate-clavate, clamp connection very rare.
4.4-6.3×5.6- 10μm,细长到宽的椭圆形
4.4-6.3×5.6- 10μm, elongate to broadly ellipsoid.
未见报道
Undescribed
澳大利亚
Australia
Kile & Watling
1988
22 A. hinnulea Kile & Watling 未见报道
Unknown
未见报道
Unknown
直径20-80
(-120)mm,坚实,肉质,边缘有时有占直径1/3的条纹
20-80 (-120)mm diam., firm to fleshy, margin sometimes distinctly striate for up to one third of diameter.
四孢型,偶尔双孢型,棍棒状-圆柱状,无锁状联合
Four-spored occasionally two-spored, clavate-cylindric, without clamp connection.
6-8.5(-9)×(3.5-)4-6(-6.5)μm,宽椭圆形到卵圆形
6-8.5(-9)×(3.5-)4-6(-6.5)μm, broadly ellipsoid to ovoid.
褐色,圆柱形
Brown, cylindrical.
澳大利亚,新西兰
Australia, New Zealand
Kile & Watling
1983
待续
表1
23 A. paulensis Capelari + 未见报道
Unknown
直径0.9-9.2cm,初时凸,成熟后扁平状,中央有褐色的小鳞片
0.9-9.2cm diam., convex when young to aplanate-depressed as mature, brown squamules concentrated in the pileus centre.
双孢型到四孢型,圆柱状,无锁状联合
Two-four-spored, cylindric, without clamp connection.
8.5-10×5.5-6μm,长椭圆形,透明,稍微有些厚壁
8.5-10×5.5-6μm, longate-ellipsoid, hyaline, slightly thick-walled.
未见报道
Undescribed
非洲
Africa
Lima et al. 2008
24 A. montagnei (Singer) Herink [=Armillariella montagnei Singer=? Armillaria luteobubalina Watling & Kile] + 未见报道
Unknown
直径40-81mm,表面覆盖了丰富的小鳞片
40-81mm diam., covered with abundant, small squamules.
双孢型到四孢型,棍棒状
Two-four-spored, clavate.
(8-)9.5-11(-12)× (5-)5.5-7(-8.5)μm 部分扁平,分支无规则
Applanate in section, branching irregularly.
南美洲
South America
Singer 1956, 1970;
Pildain et al. 2010
25 A. umbrinobrunnea (Singer) Pildain & Rajchenb. [= Armillariella montagnei var. umbrinobrunnea Singer] + 未见报道
Unknown
直径15-50mm,散布着丰富、相对大而浓密的小鳞片
15-50mm diam., characterized by the presence of abundant, relatively large and dense squamules.
棍棒状
Clavate
(7-)9-11(-13)× (5-)5-6.5(-7)μm 圆柱形或扁平,光滑,不是严格的二叉分枝
Cylindrical or applanate, smooth, branched but not strictly dichotomous.
南美洲
South America
Pildain et al. 2010
待续
表1
26 A. sparrei (Singer) Herink [=Armillariella sparrei Singer] + 未见报道
Unknown
直径35-100mm,中间有一部分小鳞片,但并不多,成熟后消失
35-100mm diam., with few to many squamules at the center, but never abundant, that disappear with age.
棍棒状
Clavate
(7-)8.5-11(-12)× (4-)5-6.5(-7)μm 扁平,光滑,很多分支
Rhizomorphs applanate, smooth, heavily branched.
南美洲
South America
Pildain et al. 2010
27 单生蜜环菌
A. singula J.Y. Cha & Igarashi
+ 异宗配合
Heterothallism
直径2.5-3.8cm,球形,初时凸到半球形,之后平凸,有钝的凸起
2.5-3.8cm diam., globose, convex to hemispherical when young, piano-convex to plane later, obtusely umbonate.
四孢型,棍棒状,有锁状联合
Four-spored, clavate, with a clamp connection.
6.2-10.6×3.6 -6.2μm,宽椭圆形到卵圆形,带有细尖,表面光滑透明
6.2-10.6×3.6 -6.2μm, broadly elliptical to ovate, smooth, hyaline, with a apiculus.
顶端尖锐,圆柱形,单轴分枝
With cylindrical sharp tips and monopodial branches.
日本
Japan
Cha et al. 1994
28 北海道蜜环菌
A. jezöensis J.Y. Cha & Igarashi
+ 异宗配合
Heterothallism
直径4.7-6.8cm,幼时半球-凸状到凸,之后平凸再到平,有时稍有 凸起
4.7-6.8cm diam., hemispherical-convex to convex when young, then piano-convex to plane, sometimes slightly umbonate.
四孢型,棍棒状,有锁状联合
Four-spored, clavate, with a clamp connection.
6.3-10.3×4.8 -6.3μm,宽椭圆形到卵圆形,带有细尖,表面光滑透明
6.3-10.3×4.8
-6.3μm, broadly elliptical to ovate, smooth, hyaline, with a apiculus.
圆柱状,单轴 分枝
Cylindrical, with monopodial branches.
日本
Japan
Cha et al. 1994