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灵芝科

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灵芝科
亮盖灵芝
科学分类 编辑
界: 真菌界 Fungi
门: 担子菌门 Basidiomycota
纲: 伞菌纲 Agaricomycetes
目: 多孔菌目 Polyporales
科: 灵芝科 Ganodermataceae
(Donk英语Marinus Anton Donk) Donk (1948)[1]
模式属
灵芝属
Ganoderma
P.Karst.英语Petter Adolf Karsten (1881)
  • 详见内文
异名[2]
  • Ganodermoideae Donk (1933)

灵芝科学名:Ganodermataceae)是担子菌门多孔菌目的一个,于1948年由荷兰真菌学家马里纳斯·安东·栋克英语Marinus Anton Donk发表描述。本科真菌一般生长在树木或腐木上,有腐生者,也有可造成白腐病(white rot)的植物病原菌,共有超过200种物种,广泛分布于世界各地[3],又以热带地区为大宗[4]东亚传统医学中的药材“灵芝”[注 1][5][6]即是本科模式属灵芝属的物种[7]

本科类群因种内外形变异很大,造成长久以来分类上的困难,其中椭圆形、两层细胞壁且内壁有突起的担孢子是本科的重要特征。传统上本科类群依子实体表面质地有无光泽、子实体表面菌丝的排列方式与担孢子的形态等外形特征分为灵芝属、假芝属鸡冠孢芝属网孢灵芝属松芝属等五个[3]

形态

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本科类群的特征为担孢子具有两层细胞壁,其中内壁有色、较厚并常有突起物,外壁则较薄且多为无色[1],此类担孢子被部分学者称为灵芝类孢子(ganodermatoid spores)[注 2];另外本科类群菌丝常具有三种型态,即薄壁透明的生殖菌丝、厚壁有色的骨骼菌丝与无色多分枝的缠绕菌丝,不过也有部分物种的菌丝只有两型或单型[9][10]

分类

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1933年,荷兰真菌学家马里纳斯·安东·栋克英语Marinus Anton Donk描述发表了多孔菌科下的一个新亚科灵芝亚科(Ganodermoideae),并于1948年将其独立为一新的科,即灵芝科(Ganodermataceae)[11]灵芝属为本科的模式属,亦为物种数最多的属,其担孢子有一端呈截形(truncate apex);假芝属Amauroderma)则是本科物种数第二多的属,其担孢子两端均不呈截形,且两属孢子内壁的突起形状稍有不同[3][12]松芝属Tomophagus)于1905年被发表,当时仅有松芝一物种,后来虽一度被认为是灵芝属的同物异名,但近年分子研究显示其为一独立演化支,已将松芝重新归入该属中[13][14];1972年,又有学者提出了鸡冠孢芝属Haddowia)与网孢灵芝属Humphreya)等新属[15]

过去即有学者认为假芝属可能为复系群,2017年,一篇分子种系发生学研究以核糖体DNA的部分序列与RNA聚合酶II大次单元(RPB1)、真核延伸因子1α(TEF-1α)等基因序列分析本科类群的亲缘关系,结果显示假芝属与灵芝属皆为复系群:灵芝属中咖啡网孢灵芝英语Humphreya coffeataG. coffeatum[注 3]多分枝灵芝英语Ganoderma ramosissimumG. ramosissimum)、粗皮灵芝瑞典语Trachyderma tsunodaeG. tsunodae[注 4]树脂灵芝G. subresinosum[注 5]页面存档备份,存于互联网档案馆)皆分别自成一演化支,其他类群则共为一单系群;假芝属中云南乌芝A. yunnanense)自成一演化支,惑乌芝瑞典语Amauroderma perplexumA. perplexum)、皱盖假芝英语Amauroderma rudeA. rude)与环皱乌芝瑞典语Amauroderma rugosumA. rugosum)组成一演化支,另外有两物种各自成一演化支,分别被归入ForaminisporaFurtadoa两个新属,其中后者只有一种型态的菌丝。假芝属的其他类群(狭义类群)则为一单系群。此外松芝属亦为单系群[3]

下级分类

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本科包括以下属:

  • 假芝属 Amauroderma (Patouillard) Torrend, 1920
  • Amaurodermellus Costa-Rezende, Drechsler-Santos & Góes-Neto
  • Cristataspora Robledo & Costa-Rezende
  • Furtadoa Costa-Rezende, Robledo & Drechsler-Santos[3]
  • Furtadoella
  • Furtadomyces Leonardo-Silva, Cotrim & Xavier-Santos
  • Ganodermites A.Fleischmann, M.Krings, H.Mayr & R.Agerer, 2007
  • Lazulinospora
  • Neoganoderma
  • Sanguinoderma Y.F.Sun, D.H.Costa & B.K.Cui, 2020
    • Sanguinoderma bataanense (Murrill) Y.F.Sun & B.K.Cui
    • 粗柄雪芝 Sanguinoderma elmerianum (Murrill) Y.F.Sun & B.K.Cui
    • Sanguinoderma flavovirens Y.F.Sun & B.K.Cui
    • Sanguinoderma guangdongense B.K.Cui & Y.F.Sun
    • Sanguinoderma infundibulare B.K.Cui & Y.F.Sun
    • Sanguinoderma laceratum Y.F.Sun & B.K.Cui
    • Sanguinoderma longistipitum B.K.Cui & Y.F.Sun
    • Sanguinoderma melanocarpum B.K.Cui & Y.F.Sun
    • Sanguinoderma microporum Y.F.Sun & B.K.Cui
    • Sanguinoderma microsporum B.K.Cui & Y.F.Sun
    • Sanguinoderma perplexum (Corner) Y.F.Sun & B.K.Cui
    • Sanguinoderma reniforme Y.F.Sun & B.K.Cui
    • Sanguinoderma rude (Berk.) Y.F.Sun, D.H.Costa & B.K.Cui
    • 乌血芝 Sanguinoderma rugosum (Blume & T.Nees) Y.F.Sun, D.H.Costa & B.K.Cui
    • Sanguinoderma sinuosum Y.F.Sun & B.K.Cui
    • Sanguinoderma tricolor B.K.Cui & Y.F.Sun
  • Sinoganoderma
  • 黑柄栓孔菌属 Whitfordia Murrill
    • 伸展黑柄栓孔菌 Whitfordia scopulosa (Berk.) Nú?ez & Ryvarden,为Amauroderma scopulosum (Berk.) Imazeki的异名
曾经在灵芝科的属:

化石纪录

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  • Ganodermites英语Ganodermites:2007年发表、出土于利比亚撒哈拉沙漠矿化子实体化石Ganodermites libycus,年代为距今2300万-1750万年前的中新世早期,形态与现生灵芝属物种相似,具有三种型态的菌丝,亦有两层细胞壁、一端呈截形的担孢子[4]
  • Archeterobasidium:1979年发表、出土于利比亚北部的矿化子实体化石Archeterobasidium syrtae,年代为中新世中期,其担孢子形态与现生灵芝科类群相似(但也有学者认为其担孢子与现生的多年异担子菌英语Heterobasidion annosumHeterobasidion annosum)相似),具有两种型态的菌丝[4][21][4]
  • 另外还有数个中新世、更新世的灵芝属物种化石曾被发表[4]

注脚

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  1. ^ 东亚传统医学中的药材灵芝过去被认为是亮盖灵芝G. lucidum),但分子鉴定显示其应为灵芝(G. lingzhi)、重伞灵芝英语Ganoderma multipileumG. multipileum)与四川灵芝英语Ganoderma sichuanenseG. sichuanense)等物种。
  2. ^ 但假芝属的A. coltricioides瑞典语Amauroderma coltricioides为一例外,其担孢子壁平滑,非灵芝类孢子[4][8]
  3. ^ 过去曾被部分学者归入网孢灵芝属Humphreya[3][16]
  4. ^ 过去曾被部分学者归入另一属Trachyderma,但后来该属被视为灵芝属的同物异名[3][17]
  5. ^ 过去曾被部分学者归入另一属Magoderna,但后来该属被视为灵芝属或假芝属的同物异名[3][15][18]

参考文献

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  1. ^ 1.0 1.1 Donk, Marinus A. Notes on Malesian fungi. I. Bulletin du Jardin botanique de Buitenzorg. 1948, 17 (III): 473–482. 
  2. ^ Ganodermataceae. MycoBank. [2019-10-22]. (原始内容存档于2019-04-10). 
  3. ^ 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 Costa-Rezende, D.H.; Robledo, G.L.; Góes-Neto, A.; Reck, M.A.; Crespo, E.; Drechsler-Santos, E.R. Morphological reassessment and molecular phylogenetic analyses of Amauroderma s.lat. raised new perspectives in the generic classification of the Ganodermataceae family. Persoonia英语Persoonia (journal). 2017, 39 (1): 254–269. ISSN 0031-5850. doi:10.3767/persoonia.2017.39.10. 
  4. ^ 4.0 4.1 4.2 4.3 4.4 4.5 Fleischmann, Andreas; Krings, Michael; Mayr, Helmut; Agerer, Reinhard. Structurally preserved polypores from the Neogene of North Africa: Ganodermites libycus gen. et sp. nov. (Polyporales, Ganodermataceae). Review of Palaeobotany and Palynology. 2007, 145 (1-2): 159–172. ISSN 0034-6667. doi:10.1016/j.revpalbo.2006.10.001. 
  5. ^ Hapuarachchi, KK; Wen, TC; Deng, CY; Kang, JC; Hyde, KD. Mycosphere Essays 1: Taxonomic Confusion in the Ganoderma lucidum Species Complex. Mycosphere (Mushroom Research Foundation). 2015, 6 (5): 542-559 [2023-06-01]. doi:10.5943/mycosphere/6/5/4. (原始内容存档于2023-06-01). 
  6. ^ Hennicke, Florian. Distinguishing commercially grown Ganoderma lucidum from Ganoderma lingzhi from Europe and East Asia on the basis of morphology, molecular phylogeny, and triterpenic acid profiles. Phytochemistry (The Phytochemical Society of Europe and The Phytochemical Society of North America). 2016, 127: 29-37 [2023-06-01]. doi:10.1016/j.phytochem.2016.03.012. (原始内容存档于2023-06-01) –通过Elsevier Science Direct. 
  7. ^ Wachtel-Galor, Sissi. Chapter 9 Ganoderma lucidum (Lingzhi or Reishi)A Medicinal Mushroom. Herbal Medicine: Biomolecular and Clinical Aspects. 2nd edition.. CRC Press Taylor and Francis. 2011 [2017-02-22]. ISBN 978-1-4398-0713-2. (原始内容存档于2015-09-10). 
  8. ^ Aime, M. Catherine; Henkel, Terry W.; Ryvarden, Leif. Studies in Neotropical Polypores 15: New and Interesting Species from Guyana. Mycologia. 2003, 95 (4): 614. ISSN 0027-5514. doi:10.2307/3761937. 
  9. ^ Seo GS, Kirk PM. Systematics of Ganoderma. J. Flood, P. D. Bridge, M. Holderness (编). Ganoderma Diseases of Perennial Crops. CABI Bioscience. 2000: 23–45. 
  10. ^ Hapuarachchi, KK. Ganodermataceae (Polyporales): Diversity in Greater Mekong Subregion countries (China, Laos, Myanmar, Thailand and Vietnam). Mycosphere. 2019, 10 (1): 221–309. ISSN 2077-7019. doi:10.5943/mycosphere/10/1/6. 
  11. ^ Furtado, Joao S. Relation of Microstructures to the Taxonomy of the Ganodermoideae (Polyporaceae) with Special Reference to the Structure of the Cover of the Pilear Surface. Mycologia. 1965, 57 (4): 588. ISSN 0027-5514. doi:10.2307/3756735. 
  12. ^ Hapuarachchi, KK. High diversity of Ganoderma and Amauroderma (Ganodermataceae, Polyporales) in Hainan Island, China. Mycosphere. 2018, 9 (5): 931–982. ISSN 2077-7019. doi:10.5943/mycosphere/9/5/1. 
  13. ^ Nowrousian, Minou; Loyd, A. L.; Barnes, C. W.; Held, B. W.; Schink, M. J.; Smith, M. E.; Smith, J. A.; Blanchette, R. A. Elucidating "lucidum": Distinguishing the diverse laccate Ganoderma species of the United States. PLOS ONE. 2018, 13 (7): e0199738. ISSN 1932-6203. doi:10.1371/journal.pone.0199738. 
  14. ^ Hong, Soon Gyu; Jung, Hack Sung. Phylogenetic analysis of Ganoderma based on nearly complete mitochondrial small-subunit ribosomal DNA sequences. Mycologia. 2004, 96 (4): 742–755. ISSN 0027-5514. doi:10.1080/15572536.2005.11832922. 
  15. ^ 15.0 15.1 R.L. Steyaert. Species of Ganoderma and related genera mainly of the Bogor and Leiden Herbaria 7 (1). Persoonia: 55-118. 1972. 
  16. ^ Humphreya coffeata. MycoBank. 
  17. ^ Trachyderma. MycoBank. [2019-10-22]. (原始内容存档于2018-07-27). 
  18. ^ Magoderna. MycoBank. [2019-10-22]. (原始内容存档于2018-07-27). 
  19. ^ Polyporopsis. MycoBank. [2019-10-22]. (原始内容存档于2018-07-27). 
  20. ^ Richter, Christian; Wittstein, Kathrin; Kirk, Paul M.; Stadler, Marc. An assessment of the taxonomy and chemotaxonomy of Ganoderma. Fungal Diversity. 2014, 71 (1): 1–15. ISSN 1560-2745. doi:10.1007/s13225-014-0313-6. 
  21. ^ Koeniguer, J.C., Locquin, M.V. Un polypore fossile à spores porées du Miocène de Libye: Archeterobasidium syrtae, gen. et sp. nov.. Ministère des Universités Comité des Travaux Historiques et Scientifiques (编). Comptes Rendus du 104e Congrès National des Sociétés Savantes, Bordeaux, 1979, fasc. I (Paléobotanique). 1979: 323-329 (法语).