担子菌Basidiomycete > 蘑菇目Agaricales > 蘑菇科Agaricaceae
> 青褶伞属Chlorophyllum
上一次编辑：2022/6/8
Last edit: 2022/6/8
作者：卷柏
By Jiiaming Cai, mycopedia
青褶伞属Chlorophyllum Massee (1898)
= Endoptychum Czern. (1845)
青褶伞属主要包含了一些体型中等至大型的伞菌种类。该属最常见的种类是大青褶伞Chlorophyllum molybdites，也叫铅绿褶伞。大青褶伞最突出的特征是其孢子印颜色——呈现铅绿色或橄榄绿色，而它的菌褶成熟后也会被孢子染上绿色调，“青褶”以及属名Chlorophyllum指的都是这个特点。这样的孢子印颜色整个菌物界都是独树一帜的存在。实际上，青褶伞属中只有大青褶伞孢子印呈现绿色，而其他的所有物种孢子印都是白色或淡黄色的（所以最常见的大青褶伞才是异类）。青褶伞属曾经只包含了大青褶伞一个物种，后来分子生物学研究揭示了以前一些归入大环柄菇属Macrolepiota、白环蘑属Leucoagaricus的物种的真实身份是青褶伞属的一员，即便它们的孢子印颜色和大青褶伞的孢子印颜色相去甚远。
青褶伞属经常和大环柄菇属混淆。大环柄菇属很多物种都是美味的食用菌，但青褶伞属很多物种有毒。宏观上看两者的区别是，大环柄菇属的菌柄上或多或少有鳞片，但青褶伞属的菌柄完全光滑。青褶伞属一些较为小型的物种容易和白环蘑属混淆，但这些小型的种类往往菌柄基部折断后会变红，白环蘑属物种一般不会。
青褶伞属下还有一个异类——陀螺青褶伞C.agaricoides。与那些长得有蘑菇样的青褶伞不同，陀螺青褶伞长得像一团白色的泡沫塑料，呈现梨形或者心脏形状，外面有一层薄薄的白色包被，包裹住里面扭曲的褐色孢体，底部还有一个短的菌柄。成熟时，整个菇体便从顶端开始裂开，散发出阵阵的褐色孢子粉。其形态如此诡异，以至于人们曾经建立了一个单独的属：粉菇包属Endoptychum，其中endo意为内部，指这位奇行种把孢子包裹在体内的特征。现在，菌物学家将这种特征命名为灰菇包型，被认为是介于伞菌型（正常蘑菇）和腹菌型（马勃一样的蘑菇）的中间体。许多伞菌都有朝着马勃演化的趋势，它们不再开伞，把菌褶特化成一团团的孢体，包裹在菇体内部，最终变为马勃样的形态。这个过程被称为腹菌化gasteromycetation，许多蘑菇都有这样的趋势，被认为是蘑菇面对干旱环境时的适应特征——将孢子包裹在体内可以更好的保护孢体，防止脱水，更好地在干燥的环境下生存下去。青褶伞属中也有一些腹菌化的完全体，例如C.arizonicum，整个蘑菇就是一马勃，只有那残余的一丁点菌柄暗示着它的来历。因此，各种马勃状的蘑菇（腹菌）之间并不是近亲，它们可能曾经是完全不一样的蘑菇。
核心特征：多数物种伞菌状。腐生，地生。菌褶离生。菌柄光滑无鳞片。菌盖白色，有褐色鳞片。菌环存在，通常宿存（长期留存）。孢子印白色、淡黄色或绿色。孢子无芽孔或有平截芽孔。除了伞菌状的物种，还有少数灰菇包状和腹菌状的物种。
Chlorophyllum – what does it have to do with
chlorophyll anyway? Remember - the green pigment in plant leaves? Well, chlorophyllum refers to the distinctive green spore print
of the type species, C.molybdites. C.molybdites
is a common giant mushroom often occur in gardens, lawns and farmlands, with a
big white cap and brown squamules on it, also a thick stem and obvious ring.
Besides C.molybdites, other species in the
genus are also common in urban areas, often forming significant fairy rings.
Several
species with white or pale yellow spore print in genus Chlorophyllum was
placed in Macrolepiota or Leucoagaricus (to be honest, C.molybdites are the only species with green
spore print, and used to be the only specie in Chlorophyllum). Recent
DNA studies has revealed their relationship with C.molybdites,
despite the different spore print color. Now, Chlorophyllum can be
distinguished from Macrolepiota by its smooth stem without any forms of
squamules or scales. As for Leucoagaricus, well, most species in Leucoagaricus
are pretty small compared to those in Chlorophyllum. Several small
species in Chlorophyllum, like C.hortense,
bruise reddish when injured, especially at the base of the stem, while most
species in Leucoagaricus do not change color.
Endoptychum was a genus created especially for
the special C.agaricoides, formerly named Endoptychum agaricoides
Czern. The species is secotioid, meaning it is
between the agaricoid (toadstool-form) and the gasteroid (puffball-form)
mushrooms. It looks like a pear or a heart, so bizarre that people created a
completely new genus for it, until DNA tests revealed the true identity of it.
Mycologists now believe that this is a form of adaptation in dry environments,
to protect the precious gleba (spores producing
structures - what a wired name) from dehydrating. The process is called gasteromycetation, and it could happen on any kinds of agaricoid
mushrooms. In fact, there are species in Chlorophyllum which are totally
gasterlized, e.g., C.arizonicum. So, not all secotioid /
gasteroid mushrooms were closely related, they can be very different mushrooms
all been through gasteromycetation.
Core Features: Mainly agaricoid. Saprobic,
terrestrial. Gills free. Stem smooth without squamules. Cap white, with
brownish squamules. Rings often persistent. Spore print white, pale yellow or
green. Spores without a germ pore or with a germ pore caused by depression in
the episporium on the truncate apex. Besides for
agaricoid species, there are a few secotioid and gasteroid species.
下属物种 Chlorophyllum Species in Mycopedia:
描述：
担子果中等至大型，多数伞菌状（agaricoid），少数灰菇包状（secotioid）或腹菌状（gasteroid）。菌柄（存在时）光滑无鳞片。孢子宽椭圆形、杏仁形、近卵形、球形或近球形，无色、浅绿色、淡黄色或淡褐色，无芽孔或平截芽孔，无孔盖，多少类糊精质。
伞菌型：菌盖幼时鼓棒状至半球形，成熟后平凸至平展。菌盖白色或近白色，表面覆盖淡黄色至褐色至紫褐色的膜质（hymenidermal）结构，随着菌盖开展被撕裂为菌环和鳞片，往往菌盖中央的鳞片完整而平滑，边缘的鳞片反卷而稀疏。菌褶离生，密至较密，不等长；幼时白色，逐渐转变为淡黄色、青绿色，或不变色。菌柄中生，圆柱形，光滑无鳞片，白色至褐色，基部略膨大或膨大为球形。菌环一般存在，膜质，单层或双层，上位，多数宿存。菌肉一般近白色，部分物种菌柄菌肉受伤后菌肉变红色、红褐色或黄褐色。孢子印白色、奶油色或绿色。担子棒状，多数4孢。锁状联合有或无。褶缘囊状体存在，无侧生囊状体。（例：C.molybdites）。
灰菇包型：子实体球形、近球形、梨形、陀螺形至圆锥形。包被白色或近白色。包被表面鳞片存在时褐色糠麸状，向下渐小渐稀疏。孢体迷路状，初期白色，成熟后褐色，至少部分变为粉末状。有中生短菌柄，向上延伸形成中柱。无菌环。例：C.agaricoides。
腹菌型：子实体被果型（angiocarpic），球形，近球形，梨形或圆锥形，偶尔有短柄。包被白色，成熟后常变褐色，光滑、龟裂或有褐色鳞片。中柱存在，通常延伸至菌体中部。孢体迷路状，紧致，乳白色至淡黄色，成熟后褐色。孢子球形、近球形至椭圆形，无芽孔，厚壁，淡褐色，常带有脐状附属物。例：C.arizonicum。
生境分布：腐生，地生，往往群生或散生。草地、林地、肥沃的田地上多见。广泛分布于世界各地，多见于亚热带和热带地区。
经济价值：多数物种有毒，部分物种可食用。大青褶伞是剧毒物种，能引起严重的肠胃炎反应，但一般不致死。
模式物种：Chlorophyllum
esculentum [=大青褶伞C.molybdites]
常见物种：大青褶伞C.molybdites、变红青褶伞C.hortens、拟乳头状青褶伞C.neomastoideum、陀螺青褶伞C.agaricoides
讨论：青褶伞属的主要特征是子实体中等至大型；菌盖（或包被）白色或近白色，多数物种被淡黄色至褐色鳞片；菌柄（如果存在）无细鳞片构成的花纹；菌环（如果存在）多宿存；孢子印淡绿色、灰绿色、橄榄绿色、白色至淡黄色；担孢子无芽孔或具有平截芽孔。该属的许多物种曾被归入大环柄菇属Macrolepiota、白环蘑属Leucoagaricus等属，但核酸特征表明它们应属于青褶伞属。青褶伞属的一些腹菌化物种，例如C.agaricoides曾被归入Endoptychum属中。
General Description: Basidiomata medium to large,
mostly agaricoid, a few species secotioid or gasteroid. Stem (when present)
smooth with no scales. Spores broadly ellipsoid, amygdaliform, subovoid, globose or subglobose; hyaline, pale green, pale
yellow or pale brown in color. Without a germ pore or with a germ pore caused
by depression in the episporium on the truncate apex,
more or less dextrinoid.
Agaricoid: Pileus hemispherical when young,
convex to nearly flat when mature; white to whitish, covered with yellow, brown
or purplish brown hymenidermal covering which tears into squamules and an
annulus as the pileus expands. Squamules become distance towards the margin.
Lamellae free, crowded, lamellulae (short-gills) present; whitish when young,
becoming pale yellow or greenish or not changing color. Stem central,
cylindrical, smooth with out squamules, usually white
to brown, base slightly swollen or swollen into ball shape. Annulus usually
present, hymenidermal, single or double layered, usually persistent. Context
whitish, many species staining reddish, reddish brown or yellowish brown on
bruising in stem (especially at the base). Spore print white, pale yellow or
green. Basidia clavate, mostly 4 spored. Clamps present or absent.
Cheilocystidia present, pleurocystidia absent. e.g., C.molybdites.
Secotioid:
Basidiomata
globose, conical to pyriform. Peridium whitish. Squamules (when present)
brownish, becoming small and distance towards the base. Gleba
labyrinthiform, whitish when young, becoming
brownish, powder-like (at least partly) when mature. Stem present, tapering
upward into a columella, tapering downward into a thick base. Columella extends
to the center, and usually not reaching the apex. Annulus absent. e.g., C.agaricoides
Gasteroid:
Basidiomata
angiocarpic, globose, subglobose, pyriform or
conical, occasionally with a short stem. Peridium whitish, often becoming
brownish when mature, smooth, cracking or with brown squamules. Columella
present, extends to the center, and usually not reaching the apex. Gleba labyrinthiform, crowded,
ivory to buff, and becoming brownish when mature. Spores globose, subglobose to
ellipsoid, with out germ pores, thick-walled, pale
brown, often with a hilar appendix. e.g., C.arizonicum.
Ecology & Distribution: Saprobic, terrestrial. Scattered.
On grassland, lawns, forestland, fertile farmland, etc.
Edibility: With a few exceptions (like C.rhacodes and C.brunneum),
most species are poisonous and inedible. C.molybdites are highly
poisonous and can causes severe gastrointestinal symptoms, but usually not
lethal.
Type species: Chlorophyllum esculentum [
= C.molybdites]
Common species in China: C.molybdites, C.hortens,
C.neomastoideum, C.agaricoides
Discussion: Main characters for genus Chlorophyllum is medium to
large sized basidiomata; white to whitish pileus (or peridium), often with
yellowish to brownish squamules; stem (if present) without squamules; annulus
(if present) usually persisting; spore print pale green, greyish green, olive
green, white to pale yellow; basidiospores without a germ pore or with a germ
pore caused by depression in the episporium on the
truncate apex.
图1 青褶伞属 显微特征（引自 Zai-Wei
Ge等，2006）
大青褶伞Chlorophyllum molybdites A.担孢子（平截芽孔） B.缘囊体，比例尺=10μm
Fig 1. Genus Chlorophyllum
Microscopic Feature (Retrieved from Zai-wei Ge et al.
2006)
Chlorophyllum molybdites A.
Basidiospores with with a germ pore caused by
depression in the episporium on the truncate apex B.
Cheilocystidia, Bars = 10μm
参考资料References
杨祝良主编. 中国真菌志第五十二卷环柄菇类（蘑菇科）[M].2019.科学出版社
杨祝良，吴刚，李艳春，王向华，蔡箐编著. 中国西南地区常见食用菌和毒菌[M].2021.科学出版社
Zai-Wei
Ge，Zhu
L. Yang. The genus Chlorophyllum (Basidiomycetes) in China, 2006
Else
Vellinga, Rogier de Kok,
Tom D Bruns.Phylogeny and Taxonomy of Macrolepiota
(Agaricaceae) [J].2003.Mycologia 95(3):442-56
Zai-Wei
Ge, Adriaana Jacobs, Else C. Vellinga, Phongeun Sysouphanthong, Retha
van der Walt, Carmine Lavorato, Yi-Feng An, and Zhu
L. Yang. A multi-gene phylogeny of Chlorophyllum (Agaricaceae, Basidiomycota):
new species, new combination and infrageneric classification[J]2018, MycoKeys
(32): 65–90.
Michael Loizides,
Pablo Alvarado, Elias Polemis, Matthias Gube. Multiple evolutionary origins of sequestrate species
in the agaricoid genus Chlorophyllum[J]2020. Mycologia
112(2):1–23
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引用 Cite as
蔡佳铭.青褶伞属Chlorophyllum. 菌物志. 2022,6 网站：www.mycopedia.top
Jiaming Cai. Genus Chlorophyllum
in China. Mycopedia. 2022,6 Website: www.mycopedia.top
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