担子菌Basidiomycete > 蘑菇目Agaricales > 蘑菇科Agaricaceae > 层蘑菇属Hymenagaricus
上一次编辑：2022/7/9
Last edit: 2022/7/9
作者：卷柏
By Jiiaming Cai, mycopedia
层蘑菇属Hymenagaricus Heinem. 1981
图1 最早描述于泰国的萨莎蒙层蘑菇Hymenagaricus saisamornae（暂拟，该种种加词是以泰国菌物学家Prof. Dr. Saisamorn
Lumyong命名），比例尺=10mm （引自Jaturong Kumla
et.al. 2021）
Fig
1. Hymenagaricus saisamornae originally
described in Thailand, bars = 10mm (Retrieved from Jaturong Kumla
et.al. 2021)
层蘑菇属是蘑菇科下的一个属，是一类体型小至中等的腐生真菌。“层”指的是其菌盖鳞片由膜皮状的细胞构成。该属和黄蘑菇属Xanthagaricus有着密切关系，黄蘑菇属曾经就作为层蘑菇属的一个亚属处理。该属和黄蘑菇属的区别是该属的孢子颜色一般较深，接近紫黑色，而黄蘑菇属则接近黄褐色；层蘑菇属盖表鳞片为膜皮型（hymeniderm，无球状细胞），或者上皮型加上膜皮状排列的菌丝（epithelial with hymeniform cells），而黄蘑菇属多数为上皮型（epithelial，可见球状细胞），少数物种表皮型（cutis）或毛皮型（trichoderm），且常常有黄色的胞内色素。不过随着分子树将越来越多的物种归入层蘑菇或黄蘑菇属，两属在形态学上的区别变得渐渐模糊，一些孢子颜色较淡，接近黄褐色的物种，其在分子树上的位置依然属于层蘑菇属，例如图1的萨沙蒙层蘑菇。而菌盖表皮的特征现在也开始变得混乱起来：一些盖表鳞片膜皮型的物种被归入了黄蘑菇属（没有球状细胞），它的盖表形态和一些层蘑菇属物种十分相似。
图2 左为萨莎蒙层蘑菇，右为黄蘑菇属的Xanthagaricus omanicus，该分类得到了分子树的支持。 左边盖表鳞片的被描述为“上皮型加上膜皮状排列的菌丝”，右边的被描述为“毛皮型”。虽然描述不同，但其实两者的结构很相似（反正我是没看出来什么区别）。前者的孢子颜色被描述为“黄褐色”，后者的孢子被描述为“淡黄褐色”，因此孢子颜色也无法准确区分它们。（左图引自Jaturong Kumla
et.al. 2021，右图引自Moza Mohammed AL-Kharousi
et.al.2022）
Fig
2. On the left, Hymenagaricus saisamornae. On
the right, Xanthagaricus omanicus. Both are
supported by molecular studies. The squamules of the former is described as
‘epithelial with hymeniform cells’, and the ones of the latter is described as
‘trichoderm’. Find the differences. (Left retrieved from Jaturong Kumla
et.al. 2021, right retrieved from Moza Mohammed AL-Kharousi et.al.2022）
我国很可能有该属物种的分布，但目前暂未发现。台湾层蘑菇H.taiwanensis因其孢子为黄褐色、盖表鳞片细胞近球形至长椭圆形至近棒状，以及分子证据的支持下，现在已经被转移到了黄蘑菇属，即台湾黄蘑菇X.taiwanensis
核心特征：伞菌状。腐生，地生。子实体小型至中等。菌盖有鳞片，鳞片膜皮型。菌褶成熟后紫黑色，偶尔颜色更淡。孢子一般紫黑色，少数近黄褐色。其他特征基本同黄蘑菇属Xanthagaricus。
Hymenagaricus is a small genus belongs to
Agaricaceae, contains a series of small to medium sized, saprobic agarics.
‘Hymen’ refers to its epithelial squamules composed of hymeniform cells. The
genus is close to Xanthagaricus phylogenetically, in fact Xanthagaricus
was treated as a subgenus under Hymenagaricus. Usually, Hymenagaricus
species have spores that are purplish black, while Xanthagaricus species
have yellowish ones; the former have hymeniderm
squamules pileipellis (without globose cells), or epithelial with hymeniform
cells, while the latter mostly have an epithelial pileipellis (with globose to
subglobose cells), few species have a cutis or a trichoderm pileipellis,
usually with yellowish intracellular pigment. However, as phylogenetic studies
putting more and more species into the two genera, the differences between Hymenagaricus
and Xanthagaricus are blurred. Some Hymenagaricus with pale
spores (yellowish-brown). Also, pileipellis structure seems to be variable.
It
is likely that Hymenagaricus is distributed in China, although not a single
species has been found. H.taiwanensis
is now transferred to Xanthagaricus, as X.taiwanensis.
Core Features: Agaricoid. Saprobic, terrestrial.
Basidiomata small to medium. Cap with squamules, mostly hymeniderm. Gills, when
mature, purplish black or paler. Spores purplish black or paler. Others same as
Xanthagaricus.
描述 Description：
担子果伞菌状，小型至中等。菌盖半球形至凸镜形，后逐渐平凸或平展。盖表有鳞片。菌褶离生，黑褐色、紫褐色至褐色。菌柄基本等长，圆柱状，中生，表面有鳞片。菌环存在，但可能消失。孢子印黑褐色、紫褐色至近褐色。担孢子常紫黑色，有时颜色淡，呈现褐色，椭圆形至近椭圆形。褶缘囊状体存在。侧生囊状体和锁状联合缺失。盖表鳞片膜皮型（hymeniderm）型。
生境分布：腐生，地生，往往群生或簇生。林地表多见。
经济价值：食用性未知。
模式物种：Hymenagaricus hymenopileus
General Description: Basidiomata agaricoid, small to
medium sized. Pileus hemispherical to convex, becoming plano-convex or nearly
flat. With squamules on the surface of the pileus. Lamellae free, blackish
brown, purplish brown to brown. Stem equal, cylindrical, central, with
squamules. Annulus present, but usually fugacious. Spore print blackish brown,
purplish brown to brown. Basidiospores usually purplish black, sometimes paler,
nearly brownish, ellipsoid to nearly so. Cheilocystidia present. Pleurocystidia
and clamps absent. Squamules a hymeniderm.
Ecology & Distribution: Saprobic, terrestrial. Scattered
or in clusters. Usually on forestland.
Type species: Hymenagaricus hymenopileus
图3 层蘑菇属的Hymenagaricus mlimaniensis，a-c子实体，d-e显微结构：i担孢子，ii担子，iii-iv盖表菌丝 （引自Zubura Mwanga et.al.2014）
Fig
3. Hymenagaricus mlimaniensis, a-c fruiting body, d-e
microscopic feature: I basidiospores, ii basidia, iii-iv pileipellis (Retrieved
form Zubura Mwanga et.al.2014)
分子树Phylogenetic Tree：
基于多篇参考文献，整理出黄蘑菇属、假环柄菇属、层蘑菇属和海氏蘑菇属的大致系统发育关系。青褶伞属为外群。
黄蘑菇属和假环柄菇属构成一个类群，海氏蘑菇属落入层蘑菇属内，但两者的宏观形态差异太大，因此没有合并。
粉尘环柄菇属Coniolepiota和Eriocybe属已被确认是姊妹群且位于该系统树内，但该姊妹群在系统发育树中的具体位置未定。蘑菇属Agaricus、小蘑菇属Micropsalliota、绿托褶菇属Clarkeinda在该系统树中的位置有争议。
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
Moza
Mohammed AL-Kharousi, Shah Hussain, Marwa A. Al-Muharabi, Zahra Al-Shabibi. The
Genus Xanthagaricus: An Updated Global Species Distribution and Phylogeny with
the Description of Two New Species from Oman[J]2022. Journal of Fungi — Open
Access Mycology Journal 8(2):173
Zai-Wei
GE, Chien-Ming CHEN, Zhu-Liang YANG. A new species of
the genus Hymenagaricus (Basidiomycota) from Taiwan and its phylogenetic
position inferred from ITS and nLSU sequences[J]2008,
Cryptogamie, Mycologie,
2008, 29(3): 259-265
Iqbal Hosen, Zong-ping
Song, Genevieve Gates, Tai-Hui Li. Xanthagaricus caeruleus ,
a new species with ink-blue lamellae from southeast China[J]2017, Mycoscience 59(2)
Jaturong
Kumla, Nakarin Suwannarach, Nopparat Wannathes. 2021. Hymenagaricus saisamornae
sp. nov. (Agaricales, Basidiomycota) from Northern
Thailand. Chiang Mai Journal of Science 48(3):827-836
Zuhura
Mwanga, Donatha Tibuhwa.
2014. Morphology and molecular taxonomy of Hymenagaricus mlimaniensis
species nov: A new Basidiomycota mushroom from Mlimani main campus, Tanzania. Journal of Yeast and Fungal
Research, Vol. 5(8), pp.96-102
版权 Copyright
除了标明“引自…”的图片引用于文献中外，其他图片的版权属拍摄者所有。
Pictures: photographer all rights
reserved, except for pictures captioned ‘Retrieved from …….’
引用 Cite as
蔡佳铭.层蘑菇属Hymenagaricus. 菌物志. 2022,7
Jiaming Cai. Genus Hymenagaricus
in China. Mycopedia. 2022,7
沪ICP备2022020698号