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菌物学报, 2023, 42(1): 38-49 doi: 10.13346/j.mycosystema.220240

综述

黏菌分类学史研究

陈双林,,*

南京师范大学生命科学学院，江苏 南京 210023

The taxonomic history of Myxomycetes (Myxogastrea)

CHEN Shuanglin,,*

College of Life Sciences, Nanjing Normal University, Nanjing 210023, Jiangsu, China

收稿日期: 2022-06-29   接受日期: 2022-09-13  

基金资助: 国家自然科学基金(32070007)

Corresponding authors: * chenshuanglin@njnu.edu.cn;ORCID: CHEN Shuanglin (0000-0002-2043-6341)

Received: 2022-06-29   Accepted: 2022-09-13  

Fund supported: National Natural Science Foundation of China(32070007)

作者简介 About authors

陈双林，南京师范大学生命科学学院教授现任中国菌物学会第八届理事会常务理事、裸菌专业委员会主任等职曾获首届曾呈奎孢子植物分类学奖近40年来，一直从事菌物分类学、多样性和应用基础的研究 , E-mail：chenshuanglin@njnu.edu.cn

摘要

黏菌是菌物中一个特殊和有趣的类群。为了准确而深刻地认识黏菌的内涵、多样性和系统发育关系等分类学问题，本文从黏菌命名起点开始循览黏菌分类学的发展历程，结合黏菌分类学的最新研究进展，考证了黏菌纲Myxomycetes内涵的变化和分类系统的演变及黏菌分类学研究的重要事件，并论述了中国黏菌分类学研究史，从而系统阐述了国内外黏菌分类学研究历史的沿革和演化。

关键词： 原生质团黏菌; 分类系统; 命名起点; 类群内涵; 历史沿革

Abstract

Myxomycetes are a special and interesting group of eukaryotes mainly studied by mycologists. In order to profoundly and distinctly understand taxonomic problems of myxomycetes involving with the connotation, diversity and phylogeny, the developmental history of taxonomy was reviewed from the start point of nomenclature herein. The changes of myxomycete connotation, the evolution of the classification systems and the important events of taxonomic history were textually researched in combination with the latest progress of researches. The taxonomic development history of myxomycetes in China was also expounded and discussed. The evolution and history of taxonomic studies on myxomycetes at home and abroad was thereby systematically approached.

Keywords： plasmodial slime mould; taxonomic system; starting point for the nomenclature; taxon category; development of taxonomic history

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本文引用格式

陈双林. 黏菌分类学史研究[J]. 菌物学报, 2023, 42(1): 38-49 doi:10.13346/j.mycosystema.220240

CHEN Shuanglin. The taxonomic history of Myxomycetes (Myxogastrea)[J]. Mycosystema, 2023, 42(1): 38-49 doi:10.13346/j.mycosystema.220240

菌物是地球生物圈中一切真核菌类生物的总称，包括真菌界Fungi的真菌(fungi)、藻菌界Chromista的假菌(pseudofungi)或卵菌(oomycetes)以及原生动物界Protozoa的黏菌(slime moulds) (魏江春 2012)。然而，按照命名法规，作为普通名的“slime mould”如同作为普通名的“fungi”一样，并不是分类学上的学名，包括有网黏菌(net slime mould)、内寄生黏菌(endoparasitic slime mould)、集胞黏菌(acrasid cellular slime mould)、网柄黏菌(dictyostelid cellular slime mould)和原生质团黏菌(plasmodial slime mould)等异源异型、分属于不同生物界的类群(Cavalier-Smith 2003;Cavalier-Smith & Chao 2006;Adl et al. 2007;Kirk et al. 2008;Ruggiero et al. 2015)。国际上通称的、作为学名的黏菌(Myxomycetes或Myxogastrea)类群指的仅是原生质团黏菌(plasmodial slime mould)，又称真黏菌(true slime mould)、非细胞黏菌(acellular slime mould) (Alexopoulos et al. 1996)。目前，这个类群在系统发育上被认为是一个单系类群，并被归属于原生动物界Protozoa (Ruggiero et al. 2015;Leontyev et al. 2019)。从发现之初直至现在对这个类群仍有“黏菌(myxomycetes)”和“菌虫(mycetozoa)”之争，实际上反映了对其系统发育关系的困惑，但也使其分类学研究历史独具特色。研究黏菌分类学历史的发展，对于准确认识黏菌的内涵、范畴和多样性以及逐步揭示其中类群的系统发育关系具有学术意义。

1 黏菌名称的变化和分类学的建立

无论是原来的《国际植物命名法规》(International Code of Botanical Nomenclature, ICBN)，还是由其更名为晚近的《国际藻类、真菌和植物命名法规》(International Code of Nomenclature for Algae, Fungi, and Plants, ICN)，都按照1905年国际植物学大会通过的维也纳法规明确地将1753年作为黏菌命名的起点。这个年份是Linné Species Plantarum的出版日期，这部著作中记载有6种黏菌。在此之前更早记述黏菌的是Micheli在1729年出版的Nova Plantarum Genera，记载有4个黏菌属Clathroides、Clathroidastrum、Lycogala和Mucilago，前两个属含有现在的团网菌属Arcyria与发网菌属Stemonitis的黏菌物种，后两个属则含有现在的线膜菌属Reticularia、煤绒菌属Fuligo、发菌属Comatricha、绒泡菌属Physarum、钙皮菌属Didymium、粉瘤菌属Lycogala、团毛菌属Trichia和复囊钙皮菌属Mucilago的黏菌物种。

1833年，Link和Wallroth最早赋予这一类群以“Myxomycetes”这一名称(Yamamoto 1998;Lado & Eliasson 2017)，虽然都是作为亚目等级，但他们强调了这一类群在一定形状的孢子器内产生孢子这一与真菌相近的特征。1880年，Winter将Myxomycetes由目提升为纲(Lado & Eliasson 2017)。之后，Myxomycetes作为黏菌纲的学名被黏菌学者广泛采用，详见于Macbride (1899)、Martin & Alexopoulos (1969)、Farr (1976)、Nannenge-Bremekamp (1991)、Neubert et al. (1993, 1995, 2000)、Stephenson & Stempen (1994)、Ing (1999)和Poulain et al. (2011)等的专著。de Bary (1859)在通过大量实验揭示了黏菌的形态和生活史并详尽描述了子实体发育的2种方式之后，强调了这一类群的原生动物属性，命名为Mycetozoa，主要被Rostafiński (1873)、Lister (1894)、Hagelstein (1944)和Olive (1975)等所采用。

早期的真菌学家也注意到了一些具有酷似真菌子实体特征的黏菌类群，并开展了对黏菌的分类学研究。1801年Persoon在Synopsis Methodica Fungorum一书中，首先开始并使人们关注到黏菌的分类工作(Alexopoulos 1978;Yamamoto 1998)。他为黏菌设立1纲Angiocarpi，1目Dermatocarpi，2科Trichosperpi和Gymnospermi。Trichosperpi科含粉瘤菌属Lycogala、煤绒菌属Fuligo、刺皮菌属Spumaria、双皮菌属Diderma、绒泡菌属Physarum、团毛菌属Trichia、团网菌属Arcyria、发网菌属Stemonitis和筛菌属Cribraria 9属，Gymnospermi科则有无丝菌属Licea和筒菌属Tubifera 2属，这种安排被认为产生了黏菌分类系统的雏形。1829年Fries在Systema Mycologicum一书中鉴于一些黏菌(如粉瘤菌属Lycogala)的子实体与马勃相像，而将黏菌置于担子菌的腹菌目中设立一个亚目，即腹黏菌亚目Myxogastres (Alexopoulos 1978;Yamamoto 1998)，同时，他也首次给出了黏菌营养体——原生质团发育的一些知识，但却疏于显微形态的观察。Massee (1892)几乎是唯一一位在黏菌分类专著中为这一类群保留采用“Myxogastres”作为纲的名称的学者。在18世纪下半叶和19世纪初的几十年里，描述了大量黏菌新物种的分类学家主要有Batsch、Leers、Bulliard、Schrader、Curtis、Schweinitz、Berkeley、Cooke和Winter等，他们与Persoon和Fries命名的许多黏菌物种学名至今仍被使用。

通过对早期零散的分类研究的系统性整理和总结，de Bary的学生Rostafiński (1873)在Versuch Eines Systems der Mycetozoen一书中建立了第一个黏菌分类系统，其分类思想一直影响至今，以成熟子实体的形态特征为分类依据，主要涉及透射光下孢子的颜色、孢丝是否存在及其形态、石灰质有无等。黏菌经典分类也被认为是由Massee (1892)最早提出，他建立了包括子实体中是否存在孢丝和石灰质等特征的鉴定标准。直到Martin & Alexopolous (1969)的著作出版，黏菌分类系统仍不断被完善，但都能看到Rostafiński (1873)和Massee (1892)所建立的黏菌分类形态特征体系的印记。

2 异型类群的分类归属

生物分类单元的范畴是生物的系统学需要澄清的重要问题之一，但黏菌纲(Myxomycetes或Myxogastrea)的范畴给生物分类学家造成了长期困扰，争论经久不衰，随着系统发育关系研究的不断发展和深入，才逐渐得以厘清。

在作为普通名的“黏菌(slime mould)”所包括的类群中，网黏菌Labyrinthulomycota (net slime mould)的主要特征是具有一个由分枝联结的、无壁的、表面黏性的外质丝组成的菌体，在形态特征和系统发育上都与黏菌纲Myxomycetes差距较大，也从未被纳入黏菌纲Myxomycetes的范畴。Cavalier-Smith (1993)将网黏菌作为亚门Labyrinthista置于Chromista界Euchromista亚界Heterokonta门;Kirk et al. (2008)将网黏菌作为门Labyrinthulomycota置于Chromista界;Anderson & Cavalier-Smith (2012)将网黏菌作为纲Labyrinthulea置于Straminipila界Bigyra门Sagenista亚门;Ruggiero et al. (2015)将网黏菌作为纲Labyrinthulea置于Chromista界Harosa亚界Heterokonta超门Bigyra门。

根肿菌Plasmodiophoromycota (内寄生黏菌endoparasitic slime mould)的营养体虽为多核的原质团，但并不像黏菌的原质团那样可以移位活动;游动细胞的鞭毛为前生的两根不等长尾鞭;主要行吸收式营养;在植物体内专性寄生;孢子壁含甲壳质而非纤维素。最早由Schroeter (1889)作为1个目归于黏菌纲Myxomycetes，与之平行的另外2个目是集胞菌目Acrasida和腹黏菌目Myxogastres。由于Myxogastres就是现今的Myxomycetes，因此意味着Schroeter (1889)当时也并没有认为根肿菌是真正的黏菌。Macbride (1899)的黏菌纲Myxomycetes包括有寄生黏菌亚纲Phytomyxinae (即根肿菌)、外生孢子黏菌亚纲Exosporeae和腹黏菌亚纲Myxogastres。Alexopoulus & Mims (1979)安排根肿菌纲Plasmodiophoromycetes与壶菌纲Chytridiomycetes和丝壶菌纲Hyphochytridiomycetes一起组成单倍体鞭毛真菌亚门Haplomastigomycotina。Alexopoulos et al. (1996)将根肿菌升格为门Plasmodiophoromycota置于原生生物界Protists。Kirk et al. (2008)将根肿菌纲Plasmodiophorida置于Prorozoa界Cercozoa门;Ruggiero et al. (2015)将根肿菌作为纲Phytomyxea置于Chromista界Harosa亚界Cercozoa门Endomyxa亚门。

第一个加入黏菌纲也是最后一个被排除的异型类群是鹅绒菌，最主要的特点是孢子外生。多数的黏菌分类系统均包含鹅绒菌，一般将其作为黏菌纲Myxomycetes下的一个亚纲(Lister 1894;Macbride 1899;Martin & Alexopoulos 1969)。Olive & Stoianovitch (1960)发现原柄菌后，认为鹅绒菌的子实体由白色圆柱体组成，圆柱体表面是许多顶生单孢子的纤细小梗，孢子的产生方式相近于原柄菌而不同于孢子内生的腹黏菌，由此，Olive (1975)将鹅绒菌归于原柄菌。Gilbert (1935)认为鹅绒菌子实体外表着生的孢子实质是单孢子孢囊，就是说其孢子也是内生的;Scheetz (1972)发现鹅绒菌的孢子外面包有电射稠密层，认为可能相当于内生孢子的腹黏菌的囊被，这样，鹅绒菌就还是属于黏菌。Mims & Rogers (1973)发现黏菌的孢子具有双层壁，内壁为电子透明层，生有纹饰的外壁为电子稠密层，因此，鹅绒菌的孢子壁结构与黏菌的相同，其孢子是外生的，并不是单孢子孢囊;Furtado & Olive (1971)发现联会复合体出现在鹅绒菌的原孢子中，即在原孢子中发生减数分裂，与黏菌在幼孢子中发生减数分裂的情况似乎相同，但其成熟的孢子有4个单倍体核，而黏菌的成熟孢子均为单核体，所以并不相同。这都表明鹅绒菌与黏菌是异型的。按照近年的分子系统学研究，对于鹅绒菌更为合适的分类学处理是将其作为黏菌的姐妹群(Fiore-Donno et al. 2005，2010;Shadwick et al. 2009)。Spiegel (1990)和Corliss (1994)都认为鹅绒菌与原柄菌具有更密切的亲缘关系，鹅绒菌应该归属于原柄菌，这也是当今对鹅绒菌分类处理的主流观点。Kirk et al. (2008)将鹅绒菌作为科Ceratiomyxaceae置于Protozoa界Amoebozoa门Protostelia纲Protostelida目;而Ruggiero et al. (2015)将鹅绒菌作为亚纲Exosporeae置于Prorozoa界Sarcomastigota亚界Amoebozoa门Conosa亚门Myxogastrea纲。

集胞黏菌(acrasid slime molds)最早由Brefeld (1869)发现，从van Tieghem (1880)开始直到21世纪的今天，发现了越来越多的属于这一类群的物种(Lado & Eliasson 2017)。然而，它们具有比黏菌更小的子实体和更简单的生活循环，也并不是一个单系类群(Shadwick et al. 2009)。Schroeter (1889)将“黏菌”分成集胞菌、根肿菌和腹黏菌3组，这说明很早就已认识到集胞黏菌不是真黏菌。Olive (1975)根据营养体没有细胞壁裸露的原质团或假原质团，为菌虫这个复系类群建立了裸菌虫门Gymnomyxa，其中的菌虫亚门Mycetozoa下设真菌虫纲Eumycetozoa和集胞菌虫纲Acrasea，真菌虫纲又分为原柄亚纲Protostelia、网柄亚纲Dcityostelia和黏腹亚纲Myxogastria;Ainsworth & Bisby’s Dictionary of the Fungi第7版(1983)的黏菌门含有7个纲：原柄菌纲Protosteliomycetes、鹅绒菌纲Ceratiomycetes、网柄菌纲Dictyosteliomycetes、集胞菌纲Acrasiomycetes、黏菌纲Myxomycetes、根肿菌纲Plasmodiophoromycetes和网黏菌纲Labyrinthulomycetes;第8版(1995)将集胞菌、网柄菌和根肿菌升格为门，与黏菌门平行，置于原生动物界。这说明，组成细胞(状)黏菌(celluar slime moulds)的集胞菌(acrasid celluar slime moulds)和网柄菌(dictyostelid celluar slime moulds)已被认为是异源异型的2个类群，更不同于黏菌Myxomycetes。在Cavalier-Smith (1993)的原生动物界分类系统中，集胞菌作为一个纲Heterolobosea在Adictyozoa亚界Percolozoa门Tetramitia亚门Striatorhiza超纲，网柄菌作为亚门Dictyostelia在Dictyozoa亚界Opalomyxa超界Mycetozoa门。Kirk et al. (2008)将网柄菌作为纲Dictyostelea在Protozoa界Amoebozoa门，集胞菌作为纲Heterolobosea则是Protozoa界Percolozoa门的唯一纲。Ruggiero et al. (2015)将网柄菌作为纲Dictyostelea归在Protozoa界Sarcomastigota亚界Amoebozoa门Conosa亚门，集胞菌作为纲Heterolobosea属于Protozoa界Enzoa亚界Percolozoa门Tetramitia亚门。

原柄菌由Olive & Stoianovitch (1960)发现，是最后一个加入“slime mould”的异型类群。Olive (1970)为其设立原柄亚纲，隶属于菌虫纲Mycetozoa，其中包括鹅绒菌。Ainsworth & Bisby’s Dictionary of the Fungi第6版(1971)将原柄菌亚目归入黏菌门Myxomycota黏菌纲Myxomycetes腹黏菌亚纲Myxogastromycetidae。Alexopoulos & Mims (1979)将其划归真菌界Fungi裸菌门Gymnomycota原质体裸菌亚门Plasmodiogymnomycotina，为其建立了原柄菌纲Protosteliomycetes，与黏菌纲Myxomycetes平行。在Cavalier-Smith (1993)的原生动物界Protozoa中，黏菌、原柄菌和网柄菌在Dictyozoa亚界Opalomyxa超界Mycetozoa门，原柄菌纲Protostelea和黏菌纲Myxogastrea在Eumyxa亚门，网柄菌纲Dictyostelea在Dictyostelia亚门;Kirk et al. (2008)将原柄菌作为纲Protostelia置于Protozoa界，与黏菌纲Myxogastrea和网柄菌纲Dictyostelea共同组成Amoebozoa门;Ruggiero et al. (2015)将原柄菌作为纲Protostelea置于Prorozoa界Sarcomastigota亚界Amoebozoa门Conosa亚门Myxogastrea纲。

3 分类系统的演变

Rostafiński (1873)建立了第一个真黏菌分类系统，首先为黏菌建立了纲(class)，但在纲和目之间设立了一个“cohors”等级，然后再向下划分出目(ordo)，同时，他给出的科(family)等级名词是tribus，相当于“族”(表1)，并含有45个属级单元。

表1   历史上具有影响和代表性的黏菌分类系统

Table 1  Some important and representative classification systems in the history of myxomycete taxonomy

Rostafiński (1873)Massee (1892)Lister (1925)Olive (1975)
Class MycetozoaClass MyxogastresClass MycetozoaSubphylum Mycetozoa
Cohors ExosporeaeOrder PeritricheaeSubclass ExosporeaeClass Eumycetozoa
Tribus CeratiaceaeSuborder TubulinaeFamily CeratiomyxaceaeSubclass Protostelia
Suborder CribrariaeOrder Protosteliida
Cohors EndosporeaeOrder ColumelliferaeSubclass EndosporeaeFamily Cavosteliidae
Ordo EnteridieaeSuborder StemonitaeOrder LamprosporalesFamily Ceratiomyxidae
Tribus LycogalaceaeSuborder LamprodermaeSuborder AnemineaeFamily Protosteliidae
Ordo AnemeaeOrder LithodermeaeFamily HeterodermaceaeSubclass Myxogastria
Tribus Dictyosteliaceae Tribus LiceaceaeSuborder DidymeaeFamily LiceaceaeOrder Echinosteliida
Tribus LicaethaliaceaeSuborder PhysaraeFamily LycogalaceaeOrder Trichiida
Ordo HeterodermeaeOrder CalotricheaeFamily ReticulariaceaeOrder Stemonitida
Tribus CribrariaceaeSuborder TricheaeFamily TubulinaceaeOrder Physarida
Tribus DictydiaethaliaceaeSuborder ArcyriaeSuborder CalonemineaeOrder Liceida
Ordo ReticularieaeFamily ArcyriaceaeSubclass Dictyodtelia
Tribus ReticulariaceaeFamily MargaritaceaeClass Acrasea
Ordo CalonemeaFamily TrichiaceaeSubclass Acrasia
Tribus ArcyriaceaeOrder Amaurosporales
Tribus PerichaenaceaeSuborder Calcarineae
Tribus TrichiaceaeFamily Didymiaceae
Ordo CalcareaeFamily Physaraceae
Tribus CienkowskiaceaeSuborder Amaurochaetineae
Tribus DidymiaceaeFamily Amaurochaetaceae
Tribus PhysaraceaeFamily Collodermaceae
Tribus SpumariaceaeFamily Stemonitaceae
Ordo Amaurochaeteae
Tribus Amaurochaetaceae
Tribus Brefeldiaceae
Tribus Echinosteliaceae
Tribus Enerthemaceae
Tribus Stemonitidaceae
Martin et al. (1983)Cavalier-Smith (2013)Lado & Eliasson (2017)Leontyev et al. (2019)
Class MyxomycetesClass MyxomycetesClass MyxomycetesClass Myxomycetes
Subclass CeratiomyxomycetidaeSubclass ExosporeaSubclass CeratiomyxomycetidaeSubclass Lucisporomycetidae
Order CeratiomyxalesOrder CeratiomyxidaOrder CeratiomyxalesSuperorder Cribrariidia
Family CeratiomyxaceaeFamily CeratiomyxidaeFamily CeratiomyxaceaeOrder Cribrariales
Family Cribrariaceae
Subclass MyxogastromycetidaeSubclass MixogastriaSubclass MyxogastromycetidaeSuperorder Trichiidia
Order LicealesSuperorder LucisporidiaOrder EchinostelialesOrder Reticulariales
Family CribrariaceaeOrder LiceidaFamily ClatodermataceaeFamily Reticulariaceae
Family EnteridiaceaeFamily CribraridaeFamily EchinosteliaceaeOrder Liceales
Family LiceaceaeFamily DictydiaethalidaeOrder CribrarialesFamily Liceaceae
Order EchinostelialesFamily LiceidaeFamily CribrariaceaeOrder Trichiales
Family ClatodermataceaeFamily ListerillidaeFamily DictydiaethaliaceaeFamily Dianemataceae
Family EchinosteliaceaeFamily TubiferidaeFamily LiceaceaeFamily Trichiaceae
Order TrichialesOrder TrichiidaFamily Reticulariaceae
Family DianemaceaeFamily ArcyridaeOrder TrichialesSubclass Columellomycetidae
Family TrichiaceaeFamily DianematidaeFamily ArcyriaceaeSuperorder Echinosteliidia
Order PhysaralesFamily MinakatellidaeFamily DianemataceaeOrder Echinosteliales
Family DidymiaceaeFamily TrichidaeFamily MinakatellaceaeFamily Echinosteliaceae
Family ElaeomyxaceaeFamily TrichiaceaeSuperorder Stemonitidia
Family PhysaraceaeSuperorder CollumellidiaOrder PhysaralesOrder Clastodermatales
Order EchinosteliidaFamily DidymiaceaeFamily Clastodermataceae
Subclass StemonitomycetidaeFamily ClastodermatidaeFamily ElaeomyxaceaeOrder Meridermatales
Order StemonitidalesFamily EchinosteliidaeFamily PhysaraceaeFamily Meridermataceae
Family SchenellaceaeOrder FuscisporidaOrder Stemonitidales
Family StemonitidaceaeSuborder PhysarinaSubclass StemonitomycetidaeFamily Stemonitidaceae
Family DidymiidaeOrder StemonitidalesFamily Amaurochaetaceae
Family PhysaridaeFamily StemonitidaceaeOrder Physarales
Suborder LamproderminaFamily Lamprodermataceae
Family LamprodermidaeFamily Didymiaceae
Suborder StemonitinaFamily Physaraceae
Family Stemonitidae

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Lister (1894)的黏菌分类系统基本遵循了Rostafiński (1873)的分类标准，但进行了重组。1911年，由他的女儿修订出版的第2版除了内容增加外没有更改，而1925年的第3版则将分类等级全部进行了更新，还调整了一些分类标准(表1)。

Martin & Alexopoulos (1969)的专著The Myxomycetes被公认为是黏菌分类学的一个里程碑，全面总结了到1969年为止的所有黏菌的分类学知识。他们认为黏菌是一类真菌，相应地调整了它们的分类等级和名称。

Ross (1973)将发网菌目移出腹黏菌亚纲，升格并专设发网菌亚纲Stemonitiomycetidae，这源于其十几年前研究发现的确认。Ross (1959)在进行了系统性的个体发育比较研究后指出：黏菌子实体发育有2种不同类型，发网菌目与其他黏菌不同，为基质层上型，即原质团接触基物的底层变为基质层，上面的原生质集中隆起形成孢囊。在有柄的种类中，柄是以后从上升的原生质柱体中分化形成的，常为中空或纤维索状。由于没有原生质从柄内上升，柄内不含圆胞状或杂质颗粒物体，柄和孢囊的表面也没有基质层的薄鞘。

在Ross (1973)建立发网菌亚纲之后相当长的一段时间内，黏菌分类系统基本稳定为黏菌纲划分鹅绒菌亚纲Ceratiomycetidae、腹黏菌亚纲Myxogastromycetidae和发网菌亚纲Stemonitiomycetidae的分类方案，鹅绒菌亚纲只含有鹅绒菌目一个目，腹黏菌亚纲含有刺轴菌目、无丝菌目、团毛菌目和绒泡菌目，发网菌亚纲只含有发网菌目一个目。可见于Farr (1976)、Martin et al. (1983)、Neubert et al. (1993, 1995, 2000)、Yamamoto (1998)、李玉等(2008a, 2008b)和Poulain et al. (2011)等的黏菌分类专著。

Olive (1975)按照动物分类学进行了菌虫(黏菌)的分类，他的黏腹亚纲Myxogastria就是本文所指的黏菌，其中的目、科和属基本上都与Martin & Alexopoulos (1969)分类系统相同，但是纲、亚纲和目的名字后缀都采用了动物分类单元的词尾，而科的后缀仍采用了菌物分类单元的词尾，属和种的名字依然遵守国际植物命名法规(表1)。

进入本世纪，通过分子生物学对于黏菌系统发育关系的研究，将这一类群划归原生动物界已经基本形成共识。然而，黏菌的分类系统仍然主要保持3亚纲6目的方案，命名也依然遵守International Code of Nomenclature for Algae, Fungi, and Plants。或许，这样处理是为了避免大调整和改变而引起黏菌分类和命名的混乱，事实上，也正是如此使得黏菌分类系统和分类学研究在最近30年间得以稳定发展。

Cavalier-Smith (2013)采用动物命名法提出了一个基于分子进化和系统发育证据的黏菌分类方案(表1)。尽管对于将黏菌归属原生动物界的处理已经很少争议，但是Cavalier-Smith (2013)按照动物分类学所制定的黏菌分类方案并未能影响全世界的黏菌分类学家，也没有被接受和采用。不过，Cavalier-Smith (2013)的黏菌分类系统综合了新的和有趣的信息，包括了系统发育所涉及的更多特征，对于今后实现尽可能自然地建立黏菌分类系统可能具有重要意义。

根据系统发育的分子数据，有些其他特征似乎比一些传统分类标准能够更好地刻画黏菌纲内部的演化关系，例如孢丝与囊被的连接、孢囊柄形成的细节、孢子堆的颜色以及柄内孢子状细胞的存在与否等。因此，结合分子系统学研究证据，一个新的形态分类学特征标准体系被Leontyev et al. (2019)建立和提出，并被用于黏菌类群形态特征的修正描述，建立了一个全新的黏菌分类系统(表1)。在去除了鹅绒菌亚纲之后，黏菌纲仍然设置2个亚纲，但这2个亚纲已经不是腹黏菌亚纲和发网菌亚纲，从名称到范围都发生了根本性的变化，在形态学上分别对应浅色孢子黏菌类群和暗色孢子黏菌类群。Lucisporomycetidae (或可译为亮孢黏菌亚纲)含有4个目，筛菌目Cribrariales、孔膜菌目Reticulariales、无丝菌目Liceales和团毛菌目Trichiales，其中筛菌目Cribrariales位于这个亚纲系统演化的根部;Columellomycetidae (或可译为囊轴黏菌亚纲)含有5个目，刺轴菌目Echinosteliales、碎皮菌目Clastodermatales、斑皮菌目Meridermatales、发网菌目Stemonitidales和绒泡菌目Physarales，其中刺轴菌目Echinosteliales是这个亚纲分支的起点。这个系统有别于此前依据原生质团类型和子实体发育方式为基础的黏菌分类系统，是否足够合理还需要进一步的研究，例如，其团毛菌目包括了原来无丝菌目的一些成员、绒泡菌目包括了原来发网菌目的一些成员，忽略了对原生质团特征和子实体发育方式的已有认识，因此，这个系统是否能够被国际黏菌学家接受也还需要时间的考验。

4 中国黏菌分类学研究史

中国是最早认识和记述黏菌的国家，唐代陈藏器在《本草拾遗》(约公元739年)中就记载了一种接近于煤绒菌Fuligo sp.的黏菌“鬼屎”(李玉 2002)。然而，这一最早的黏菌记录描述过于简单，并且既没有命名基本法则中的模式概念，更不符合模式法，所以无法对它进行科学的鉴定(Martin 1960;Alexopoulos et al. 1996;李玉 2002)。

现代对于中国黏菌分类学的科学研究起步较晚，如同中国其他菌物类群分类学研究的境况一样，早期工作是由外国殖民者开始的。20世纪20年代末，中泽亮治研究了小畔四郎等于1924-1928年采集的中国台湾的黏菌标本，发表了一份关于中国台湾黏菌的名录，包括26属90种42变种，这是有关中国黏菌物种分布的最早报道(Nakazawa 1929)。之后，Skvortzow (1931)报道了采自黑龙江省哈尔滨和帽儿山的黏菌32种，其中有9个新种，周宗璜和李惠中(1978)复核考证后指出，这9个新种或应否定或存疑不能确定。不过，Skvortzow的标本是现今中国可见的最早的黏菌标本，还保存在中国科学院微生物研究所菌物标本馆中。日本的江本义数先后记述了中国辽宁省和河北省的一些种类，其中由栗田周彦和小林胜采自辽宁旅顺老铁山的Physarum puniceum是第一个发现于中国的黏菌新种(Emoto 1931)。

第一位报道中国黏菌的中国学者是邓叔群先生，他发表的中国黏菌的第一份文献记录有 8科16属29种，覆盖了中国南方的广大地区(Teng & Teng 1933)。此后，邓叔群先生还分别零散报道过浙江、福建、南京和甘肃等地的数种黏菌。第一位针对一个地区进行中国黏菌系统性分类学研究的中国学者是周宗璜先生，他发表的“河北小五台山黏菌志略”描述了8属22种。次年，他又报道了“华北之黏菌”9科17属 33种(Chow 1937)，其中有3个中国新记录属和15个中国新记录种。但是，在这之后的约40年间，中国黏菌仅由邓叔群(1963)在《中国的真菌》中记述了30属142种黏菌，而未再有其他分类学研究和文献。

中国地大物博，也完全应该具有较多的黏菌物种多样性。尽管国际上从事黏菌分类研究较真菌类群研究的科学工作者明显偏少，但是国外的黏菌学者非常希望获取中国的黏菌材料。改革开放以来，一些外籍人士多次努力前来中国开展黏菌的标本采集和分类学研究。Champion & Mitchell (1980)报道了从香港收集到的黏菌;Ing (1987)报道了从香港和广东采集的24种黏菌;Yamamoto et al. (2000，2002)先后报道了云南省的黏菌81种，其中，疣丝盖碗菌Perichaena verrucifera Y. Yamam. & Shuang L Chen和拟卵孢绒泡菌Physarum ovisporoides Y. Yamam. & Shuang L Chen是新种，齿孢团毛菌宽丝变种Trichia crenulata var. latitubularis Y. Yamam. & Shuang L Chen和暗孢钙皮菌钙柄变种Didymium melanospermum var. calcipes Y. Yamam. & Shuang L Chen是新变种，另有30种黏菌是中国新记录种;Ukkola et al. (2001)、Härkönen et al. (2004a, 2004b)先后报道了湖南省的黏菌127种，其中有20种黏菌为中国新记录种;Schnittler et al. (2013)通过对362份基物进行湿室培养和野外采集的206份标本的分类学研究，报道了新疆塔里木盆地北部和天山东部的黏菌80种，其中53种来自野外采集，32种来自湿室培养，Kelleromyxa和Protophysarum这2个单种属第一次在中国被发现。

进入20世纪80年代，有关中国黏菌的研究重新活跃起来。周宗璜(1981)撰文论述了“黏菌分类问题”，从而引导中国的菌物学家重视黏菌的分类学研究。在最近40多年间，中国的黏菌分类学研究者通过大量持续的工作，努力自己牢牢地掌握中国的黏菌多样性。刘锦惠从20世纪80年代起与她的学生连续报道了中国台湾的黏菌(Liu 1980;Liu & Chang 2007)，到目前为止，发现报道的黏菌新种9个、新变种2个。而对于中国黏菌分类学研究最重要的贡献无疑要归功于李玉院士，周宗璜和李玉(1983)发表了黏菌新种无节筛菌Cribraria enodis，这是第一个由中国科学家命名的黏菌新种。1989年，他主导完成了中国黏菌物种编目及其分布的文章在Mycotaxon上发表(Li & Li 1989)，这是中国学者的黏菌分类学研究成果第一次出现在国际期刊上，同时也使得国际黏菌分类学研究者开始了解

并重视中国同行的研究;1995年，他主导的对中国筛菌科Cribrariacea黏菌的比较形态学和系统分类学研究(Li & Li 1995)，充分论述了筛菌科Cribraria、Dictydium和Lindbladia 3个属既连续又间断的系统演化关系，证明了Cribraria和Dictydium应为2个不同的属，成为国际黏菌分

类学研究的经典之作;2008年，他主编的两卷本《中国真菌志·黏菌卷》出版，从此，在国际黏菌分类学研究领域中有了不可多得的中国专著;2014年，他担任在长春举办的第8届黏菌分类学及生态学国际会议的主席，使全世界的黏菌分类学家、生态学家首次汇聚亚洲，首次在中国进行学术研讨，奠基了中国在国际黏菌分类学研究的重要地位。在潜心治学的同时，他还培养了大批从事黏菌分类学研究的学生，带领和支持他们在全国各地开展黏菌物种多样性的野外研究工作，引导和帮助他们积极参与国际合作，从而推动中国黏菌分类学研究的可持续发展，因此在由Stephenson & Rojas主编的Myxomycetes: Biology, Systematics, Biogeography, and Ecology一书中专门指出，“自20世纪80年代以来，吉林农业大学的李玉和他的一些学生进行了黏菌的分类学、生活史、分子生物学和生物化学的研究，对于黏菌研究是重要的贡献”。迄今为止，世界各地已知分布的黏菌约为1 000种(Lado 2005-2022)，当然，这一数据排除了大量已发表的同物异名。通过李玉院士及其学生的研究，中国至今已知黏菌457种(http://www.sp2000.org.cn/)，接近全球已知种数的一半，较1989年中国黏菌已知的224种增加了1倍，同时，极大地增加、完善了黏菌物种在中国的分布格局。过去的35年以来，全世界报道的黏菌新种约为400种，其中，李玉院士及其学生发现于中国并报道的黏菌新种为55种，约占全世界已知种数的近6%，对于这35年间全世界新发现黏菌物种的贡献率约达14%。

黏菌具有十分丰富的多样性，分类学研究由于是国际黏菌研究的最主要方向之一而方兴未艾，同时黏菌也以诸多独有特征而对探索生物进化论中的科学问题具有重要价值。组学时代将使黏菌分类学取得创新的发展，基于基因型性状的分子系统学研究肯定会改变黏菌分类单元的划分，相信通过分类学工作者持续和坚韧地钻研，黏菌分类学一定会在传承和发展中不断前行。

致谢

谨以此文献给导师李玉院士八十华诞，祝敬爱的导师生日快乐，感谢导师数十年的关怀和教导。

参考文献

Adl SM, Leander BS, Simpson AGB, Archibald JM, Anderson RO, Bass D, Bowser SF, Brugerolle G, Farmer MA, Karpov SA, Kolisko M, Lane CE, Lodge DJ, Mann DG, Meisterfeld R, Mendoza L, Moestrup O, Mozley-Standridge SE, Smirnov A, Spiegel FW, 2007.

Diversity, nomenclature, and taxonomy of protists

Systematic Biology, 56(4): 684-689

PMID:17661235      [本文引用: 1]

Alexopoulos CJ, 1978. The evolution of the taxonomy of the myxomycetes. In: Subramanian CV (ed.) Proceedings of the international symposium on taxonomy of fungi, part I. University of Madras, Madras. 1-8

[本文引用: 2]

Alexopoulos CJ, Mims CW, 1979. Introductory mycology. 3rd. ed. John Wiley & Sons, Inc., New York. 47-98

[本文引用: 2]

Alexopoulos CJ, Mims CW, Blackwell M, 1996. Introductory mycology. 4th ed. John Wiley & Sons, Inc., New York. 743-808

[本文引用: 3]

Anderson OR, Cavalier-Smith T, 2012.

Ultrastructure of Diplophyrs parva, a new small freshwater species, and a revised analysis of Labyrinthulea (Heterokonta)

Acta Protozoologica, 51: 291-304

[本文引用: 1]

Cavalier-Smith T, 1993.

Kingdom Protozoa and its 18 phyla

Microbiological Reviews, 57(4): 953-994

DOI:10.1128/mr.57.4.953-994.1993      PMID:8302218      [本文引用: 3]

The demarcation of protist kingdoms is reviewed, a complete revised classification down to the level of subclass is provided for the kingdoms Protozoa, Archezoa, and Chromista, and the phylogenetic basis of the revised classification is outlined. Removal of Archezoa because of their ancestral absence of mitochondria, peroxisomes, and Golgi dictyosomes makes the kingdom Protozoa much more homogeneous: they all either have mitochondria and peroxisomes or have secondarily lost them. Predominantly phagotrophic, Protozoa are distinguished from the mainly photosynthetic kingdom Chromista (Chlorarachniophyta, Cryptista, Heterokonta, and Haptophyta) by the absence of epiciliary retronemes (rigid thrust-reversing tubular ciliary hairs) and by the lack of two additional membranes outside their chloroplast envelopes. The kingdom Protozoa has two subkingdoms: Adictyozoa, without Golgi dictyosomes, containing only the phylum Percolozoa (flagellates and amoeboflagellates); and Dictyozoa, made up of 17 phyla with Golgi dictyosomes. Dictyozoa are divided into two branches: (i) Parabasalia, a single phylum with hydrogenosomes and 70S ribosomes but no mitochondria, Golgi dictyosomes associated with striated roots, and a kinetid of four or five cilia; and (ii) Bikonta (16 unicellular or plasmodial phyla with mitochondria and bikinetids and in which Golgi dictyosomes are not associated with striated ciliary roots), which are divided into two infrakingdoms: Euglenozoa (flagellates with discoid mitochondrial cristae and trans-splicing of miniexons for all nuclear genes) and Neozoa (15 phyla of more advanced protozoa with tubular or flat [usually nondiscoid] mitochondrial cristae and cis-spliced spliceosomal introns). Neozoa are divided into seven parvkingdoms: (i) Ciliomyxa (three predominantly ciliated phyla with tubular mitochondrial cristae but no cortical alveoli, i.e., Opalozoa [flagellates with tubular cristae], Mycetozoa [slime molds], and Choanozoa [choanoflagellates, with flattened cristae]); (ii) Alveolata (three phyla with cortical alveoli and tubular mitochondrial cristae, i.e., Dinozoa [Dinoflagellata and Protalveolata], Apicomplexa, and Ciliophora); (iii) Neosarcodina (phyla Rhizopoda [lobose and filose amoebae] and Reticulosa [foraminifera; reticulopodial amoebae], usually with tubular cristae); (iv) Actinopoda (two phyla with axopodia: Heliozoa and Radiozoa [Radiolaria, Acantharia]); (v) Entamoebia (a single phylum of amoebae with no mitochondria, peroxisomes, hydrogenosomes, or cilia and with transient intranuclear centrosomes); (vi) Myxozoa (three endoparasitic phyla with multicellular spores, mitochondria, and no cilia: Myxosporidia, Haplosporidia, and Paramyxia); and (vii) Mesozoa (multicells with tubular mitochondrial cristae, included in Protozoa because, unlike animals, they lack collagenous connective tissue).

Cavalier-Smith T, 2003.

Protist phylogeny and the high-level classification of Protozoa

European Journal of Protistology, 39(4): 338-348

DOI:10.1078/0932-4739-00002      URL     [本文引用: 1]

Cavalier-Smith T, 2013.

Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa

European Journal of Protistology, 49(2): 115-178

DOI:10.1016/j.ejop.2012.06.001      PMID:23085100      [本文引用: 4]

I discuss how different feeding modes and related cellular structures map onto the eukaryote evolutionary tree. Centrally important for understanding eukaryotic cell diversity are Loukozoa: ancestrally biciliate phagotrophic protozoa possessing a posterior cilium and ventral feeding groove into which ciliary currents direct prey. I revise their classification by including all anaerobic Metamonada as a subphylum and adding Tsukubamonas. Loukozoa, often with ciliary vanes, are probably ancestral to all protozoan phyla except Euglenozoa and Percolozoa and indirectly to kingdoms Animalia, Fungi, Plantae, and Chromista. I make a new protozoan phylum Sulcozoa comprising subphyla Apusozoa (Apusomonadida, Breviatea) and Varisulca (Diphyllatea; Planomonadida, Discocelida, Mantamonadida; Rigifilida). Understanding sulcozoan evolution clarifies the origins from them of opisthokonts (animals, fungi, Choanozoa) and Amoebozoa, and their evolutionary novelties; Sulcozoa and their descendants (collectively called podiates) arguably arose from Loukozoa by evolving posterior ciliary gliding and pseudopodia in their ventral groove. I explain subsequent independent cytoskeletal modifications, accompanying further shifts in feeding mode, that generated Amoebozoa, Choanozoa, and fungi. I revise classifications of Choanozoa, Conosa (Amoebozoa), and basal fungal phylum Archemycota. I use Choanozoa, Sulcozoa, Loukozoa, and Archemycota to emphasize the need for simply classifying ancestral (paraphyletic) groups and illustrate advantages of this for understanding step-wise phylogenetic advances.Copyright © 2012. Published by Elsevier GmbH.

Cavalier-Smith T, Chao EEY, 2006. Phylogeny and megasystematics of phagotrophic heterokonts (Kingdom Chromista). Journal of Molecular Evolution, 62: 388-420

[本文引用: 1]

Champion CL, Mitchell DW, 1980.

Some myxomycetes collected in Hong Kong

Bulletin of the British Mycological Society, 14: 135-137

DOI:10.1016/S0007-1528(80)80009-5      URL     [本文引用: 1]

Chow CH, 1937.

Notes on myxomycetes from North China

Bulletin of the Fan Memorial Institute of Biology, Botanical Series, 7: 257-280

[本文引用: 1]

Corliss JO, 1994.

An interim utilitarian (‘user friendly’) hierarchial classification and characterization of the Protists

Acta Protozoologica, 33(1): 1-51

[本文引用: 1]

de Bary A, 1859.

Die Mycetozoen

Zeitschrift für Wissenschaftwehe Zoologie, 10: 88-175

[本文引用: 1]

Emoto Y, 1931.

Die myxomyceten der Suedmandschurei

Botanical Magazine, 45: 229-234

DOI:10.15281/jplantres1887.45.229      URL     [本文引用: 1]

Farr ML, 1976.

Flora Neotropica. Monograph No. 16. Myxomycetes

The New York Botanical Garden, New York. 1-305

[本文引用: 2]

Fiore-Donno AM, Berney CD, Pawlowski J, Baldauf SL, 2005.

Higher-order phylogeny of plasmodial slime molds (Myxogastria) based on elongation factor 1-A and small subunit rRNA gene sequences

Journal of Eukaryotic Microbiology, 52(3): 201-210

PMID:15926995      [本文引用: 1]

The Myxogastria are common soil microorganisms with a life cycle comprised of a plasmodial trophic stage and large fruiting bodies generally visible with the unaided eye. Until now, their classification has been based exclusively on a combination of morphological, ultrastructural, and developmental characters. Our study is the first attempt to examine phylogenetic relationships among these taxa using molecular data. Partial small-subunit ribosomal RNA and/or elongation factor 1-alpha gene sequences were obtained from eleven, mostly field-collected species representing the five orders of Myxogastria. Nineteen sequences were obtained and subjected to phylogenetic analysis together with 10 sequences available from GenBank. Separate and combined analyses of the two data sets support the division of Myxogastria into three distinct groups. The most basal clade consists of the Echinosteliales, an order considered to have affinities with Protostelia. The three species examined possess unpigmented or slightly pigmented spores. The second group consists of Liceales and Trichiales, taxa characterized by the presence of clear, but pigmented, spores. The third group consists of the two remaining orders, Physarales and Stemonitales, both possessing dark spores. This suggests that spore pigmentation is an evolutionarily conservative character in myxogastrians, and that the simple morphology of echinostelids is not a derived feature.

Fiore-Donno AM, Nikolaev SI, Nelson M, Pawlowski J, Cavalier-Smith T, Baldauf SL, 2010.

Deep phylogeny and evolution of slime moulds (mycetozoa)

Protist, 161(1): 55-70

DOI:10.1016/j.protis.2009.05.002      PMID:19656720      [本文引用: 1]

Mycetozoa, characterized by spore-bearing fruiting bodies, are the most diverse Amoebozoa. They traditionally comprise three taxa: Myxogastria, Dictyostelia and Protostelia. Myxogastria and Dictyostelia typically have multispored fruiting bodies, but controversy exists whether they are related or arose independently from different unicellular ancestors. Protostelid slime moulds, with single-spored fruiting bodies, are possible evolutionary intermediates between them and typical amoebae, but have received almost no molecular study. Protostelid morphology is so varied that they might not be monophyletic. We therefore provide 38 new 18S rRNA and/or EF-1alpha gene sequences from Mycetozoa and related species, including four protostelids and the enigmatic Ceratiomyxa fruticulosa. Phylogenetic analyses support the monophyly of Dictyostelia, Myxogastria, and Ceratiomyxa (here collectively called "macromycetozoa") and show that protostelids are Amoebozoa, mostly related to non-fruiting amoebae of the class Variosea, but may not be monophyletic; some phylogenetic relationships remain poorly resolved. Ceratiomyxa fruticulosa, originally regarded as a myxogastrid, but in recent decades included in Protostelia, is a deeply diverging sister to Myxogastria. The protostelids studied here plus varipodid amoebae and the flagellates Phalansterium and Multicilia together probably form the outgroup to macromycetozoa plus Archamoebae. Thus protostelids and Variosea are especially significant for understanding the evolutionary transition from solitary amoebae to macromycetozoa.Copyright 2009 Elsevier GmbH. All rights reserved.

Furtado JS, Olive LS, 1971.

Ultrastructural evidence of meiosis in Ceratiomyxa fruticulosa

Mycologia, 64: 413-416

[本文引用: 1]

Gilbert FA, 1935.

Critical events in the life history of Ceratiomyxa

American Journal of Botany, 22: 52-74

DOI:10.1002/j.1537-2197.1935.tb05008.x      URL     [本文引用: 1]

Hagelstein R, 1944.

The Mycetozoa of North America

Mineola, New York. 1-306

[本文引用: 1]

Härkönen M, Rikkinen J, Ukkola T, Enroth J, Virtanen V, Jääskeläinen K, Rinne E, Hiltunen L, Piippo S, He XL, 2004a.

Corticolous myxomycetes and other epiphytic cryptogams on seven native tree species in Hunan Province, China

Systematics and Geography of Plants, 74: 189-198

[本文引用: 1]

Härkönen M, Ukkola T, Zeng Z, 2004b.

Myxomycetes of the Hunan Province, China, Ⅱ

Systematics and Geography of Plants, 74: 199-208

[本文引用: 1]

Ing B, 1987.

Myxomycetes from Hong Kong and southern China

Mycotaxon, 30: 199-201

[本文引用: 1]

Ing B, 1999.

The Myxomycetes of Britain and Ireland - an identification handbook

The Richmond Publishing Co., Slough. 1-374

[本文引用: 1]

Kirk PM, Canon PF, David JC, Stalpers JA, 2008. Ainsworth & Bisby’s Dictionary of the fungi. 10th ed. CAB International, Wallingford. 747-771

[本文引用: 6]

Lado C, 2005- 2022.

An on line nomenclatural information system of eumycetozoa

Real Jardín Botánico, CSIC, Madrid, Spain. http://www.nomen.eumycetozoa.com (accessed 15 July 2022)

URL     [本文引用: 1]

Lado C, Eliasson U, 2017. Taxonomy and systematics:current knowledge and approaches on the taxonomic treatment of Myxomycetes. In: Stephenson SL, Rojas C (eds.) Myxomycetes: biology, systematics, biogeography and ecology. Academic Press, Elsevier, Amsterdam. 205-251

[本文引用: 4]

Leontyev DV, Schnittler M, Stephenson SL, Novozhilov YK, Shchepin ON, 2019.

Towards a phylogenetic classification of the Myxomycetes

Phytotaxa, 399(3): 209-238

DOI:10.11646/phytotaxa.399.3.5      [本文引用: 3]

The traditional classification of the Myxomycetes (Myxogastrea) into five orders (Echinosteliales, Liceales, Trichiales, Stemonitidales and Physarales), used in all monographs published since 1945, does not properly reflect evolutionary relationships within the group. Reviewing all published phylogenies for myxomycete subgroups together with a 18S rDNA phylogeny of the entire group serving as an illustration, we suggest a revised hierarchical classification, in which taxa of higher ranks are formally named according to the International Code of Nomenclature for algae, fungi and plants. In addition, informal zoological names are provided. The exosporous genus Ceratiomyxa, together with some protosteloid amoebae, constitute the class Ceratiomyxomycetes. The class Myxomycetes is divided into a bright- and a dark-spored clade, now formally named as subclasses Lucisporomycetidae and Columellomycetidae, respectively. For bright-spored myxomycetes, four orders are proposed: Cribrariales (considered as a basal group), Reticulariales, a narrowly circumscribed Liceales and Trichiales. The dark-spored myxomycetes include five orders: Echinosteliales (considered as a basal group), Clastodermatales, Meridermatales, a more narrowly circumscribed Stemonitidales and Physarales (including as well most of the traditional Stemonitidales with durable peridia). Molecular data provide evidence that conspicuous morphological characters such as solitary versus compound fructifications or presence versus absence of a stalk are overestimated. Details of the capillitium and peridium, and especially how these structures are connected to each other, seem to reflect evolutionary relationships much better than many characters which have been used in the past.

Li Y, 2002.

Textual research of “kwei shi”

Journal of Jilin Agricultural University, 24(2): 1-4 (in Chinese)

Li Y, Li HZ, 1989.

Myxomycetes from China. Ⅰ. A checklist of myxomycetes from China

Mycotaxon, 35(2): 429-436

[本文引用: 1]

Li Y, Li HZ, 1995.

Myxomycetes from China. Ⅲ. Description of a new species, Cribraria media, and discussion of the relationship between Cribraria and Dictydium

Mycotaxon, 53: 69-80

[本文引用: 1]

Li Y, Li HZ, Wang Q, Chen SL, 2008a. Flora fungorum sinicorum: Myxomycetes. Ⅰ. Ceratiomyxales, Echinosteliales, Liceales and Trichiales. Science Press, Beijing. 1-238 (in Chinese)

Li Y, Li HZ, Wang Q, Chen SL, 2008b. Flora fungorum sinicorum: Myxomycetes. Ⅱ. Physarales and Stemonitales. Science Press, Beijing. 1-204 (in Chinese)

Lister A, 1894.

A monograph of the Mycetozoa

The Trustees of the British Museum, London. 1-224

[本文引用: 3]

Lister A, 1925. A monograph of the Mycetozoa. 3rd ed. Revised by Lister G. The Trustees of the British Museum, London. 1-296

[本文引用: 1]

Liu CH, 1980. Myxomycetes of Taiwan. Ⅰ. Taiwania, 25: 141-151

[本文引用: 1]

Liu CH, Chang JH, 2007.

Myxomycetes of Taiwan. ⅩⅩ. A new species of Cribraria

Taiwania, 52(2): 164-167

[本文引用: 1]

Macbride TH, 1899.

North American slime moulds

Macmillan, New York. 1-231

[本文引用: 3]

Martin GW, 1960.

The systematic position of myxomycetes

Mycologia, 52: 119-129

DOI:10.1080/00275514.1960.12024883      URL     [本文引用: 1]

Martin GW, Alexopoulos CJ, 1969.

The Myxomycetes

University of Iowa Press, Iowa. 1-561

[本文引用: 5]

Martin GW, Alexopoulos CJ, Farr ML, 1983.

The genera of myxomycetes

University of Iowa Press, Iowa. 1-102

[本文引用: 2]

Massee G, 1892.

A monograph of the myxogastres

Methuen & Co. Ltd., London. 1-367

[本文引用: 4]

Mims CW, Rogers MA, 1973.

An ultrastructural study of spore germination in the myxomycete Stemonitis virginiensis

Protoplasma, 78: 243-354

DOI:10.1007/BF01275694      URL     [本文引用: 1]

Nakazawa H, 1929.

A list of Taiwan mycetozoa

Transaction of the Natural History Society of Taiwan, China, 19: 16-30

[本文引用: 1]

Nannenge-Bremekamp NE, 1991.

A guide to temperate Myxomycetes, An English translation by A. Feest and Y. Burggraaf of de Nederlandse Myxomyceten

Biopress Ltd., Bristol. 1-409

[本文引用: 1]

Neubert H, Nowotny W, Baumann K, 1993.

Die Myxomyceten Deutschlands und des angrenzenden Alpenraumes unter besonderer Berücksichtigung Österreichs, Band 1. Ceratiomyxales, Echinosteliales, Liceales, Trichiales

Baumann, Gomaringen. 1-343

[本文引用: 2]

Neubert H, Nowotny W, Baumann K, 1995.

Die Myxomyceten Deutschlands und des angrenzenden Alpenraumes unter besonderer Berücksichtigung Österreichs. Band 2. Physarales

Karlheinz Baumann Verlag, Gomaringen. 1-368

[本文引用: 2]

Neubert H, Nowotny W, Baumann K, Marx H, 2000.

Die Myxomyceten Deutschlands und des angrenzenden Alpenraumes unter besonderer Berücksichtigung Österreichs. Band 3. Stemonitales

Karlheinz Baumann Verlag, Gomaringen. 1-391

[本文引用: 2]

Olive LS, 1970.

The mycetozoa: a revised classification

The Botanic Review, 36: 59-89

DOI:10.1007/BF02859155      URL     [本文引用: 1]

Olive LS, 1975.

The mycetozoans

Academic Press, New York. 1-293

[本文引用: 5]

Olive LS, Stoinaovitch C, 1960.

Two new members of the Acrasiales

Bulletin of the Torrey Botanical Club, 87(1): 1-20

DOI:10.2307/2483057      URL     [本文引用: 2]

Poulain M, Meyer M, Bozonnet J, 2011. Myxomycétes, Vol. 1-2. Fédération Mycologique et Botanique Dauphiné-Savoie, Sevrier. 1-556

[本文引用: 2]

Ross IK, 1959. Fruiting in the myxomycetes. In: Anonymous (ed.) Proceedings of the Ninth International Botanical Congress, Montreal. Vol.2. 87-93

[本文引用: 1]

Ross IK, 1973.

The Stemonitomycetidae, a new subclass of Myxomycetes

Mycologia, 65: 477-485

DOI:10.1080/00275514.1973.12019457      URL     [本文引用: 2]

Rostafiński J, 1873.

Versuch eines systems der Mycetozoen

University Press, Strassburg. 1-115

[本文引用: 6]

Ruggiero MA, Gordon DP, Orrell TM, Bailly N, Bourgoin T, Brusca RC, Cavalier-Smith T, Guiry MD, Kirk PM, 2015.

A higher level classification of all living organisms

PLoS One, 10(6): e0130114

DOI:10.1371/journal.pone.0130114      URL     [本文引用: 7]

Scheetz RW, 1972.

The ultrastructure of Ceratiomyxa fruticulosa

Mycologia, 64: 38-54

PMID:4110694      [本文引用: 1]

Schnittler M, Novozhilov YK, Carvajal E, Spiegel FW, 2013.

Myxomycete diversity in the Tarim basin and eastern Tian-Shan, Xinjiang Prov., China

Fungal Diversity, 59: 91-108

DOI:10.1007/s13225-012-0186-5      URL     [本文引用: 1]

Schroeter J, 1889. Myxogasteres. In: Engler A, Prantl K (eds.) Die naturlichen Pflanzenfamilien I. The Bryologist, 1: 8-35

[本文引用: 3]

Shadwick LL, Spiegel FW, Shadwick JD, Brown MW, Silberman JD, 2009.

Eumycetozoa = Amoebozoa? SSU rDNA phylogeny of protosteloid slime molds and its significance for the amoebozoan supergroup

PLoS One, 4: e6754

DOI:10.1371/journal.pone.0006754      URL     [本文引用: 2]

Skvortzow BW, 1931.

Mycetozoa from north Manchuria, China

The Philippine Journal of Science, 46(1): 85-93

[本文引用: 1]

Spiegel FW, 1990. Phylum plasmodial slime molds, class Protostelia. In: Margulis L, Corliss JO, Melkonian M, Chapman DJ (eds.) Handbook of Protoctista. Jones and Bartlett, Boston. 484-497

[本文引用: 1]

Stephenson SL, Stempen H, 1994.

Myxomycetes:a handbook of slime molds

Timber Press Inc., Portland. 1-183

[本文引用: 1]

Teng SC, 1963.

Fungi of China

Science Press, Beijing. 5-46 (in Chinese)

Teng SC, Teng KL, 1933.

Notes on slime molds from China

Sinensia, 4: 61-81

[本文引用: 1]

Ukkola T, Harkonen M, Zeng Z, 2001.

Myxomycetes of Hunan Province, China. Ⅰ

Annales Botanici Fennici, 38: 305-328

[本文引用: 1]

Wei JC, 2012.

The pan-fungal biodiversity and prospect of R. ＆ D. of their resources

Journal of Fungal Research, 10(3): 125-129 (in Chinese)

Yamamoto Y, 1998.

The myxomycete biota of Japan

Toyo Shorin Publishing, Tokyo. 1-701

[本文引用: 4]

Yamamoto Y, Chen SL, Degawa Y, Hagiwara H, 2002.

Myxomycetes from Yunnan Province, China

Bulletin of the National Science Museum B, 28(3): 61-76

[本文引用: 1]

Yamamoto Y, Chen SL, Hagiwara H, 2000.

Myxomycetes from Yunnan Province, China, collected in 1998

Bulletin of the National Science Museum B, 26(1): 15-21

[本文引用: 1]

Zhou ZH, 1981.

The classification problem of slime moulds

Microbiology China, 8(3): 129-133 (in Chinese)

Zhou ZH, Li HZ, 1978.

Reexamination on the new myxomycetes described by Skvortzow

Acta Microbiologica Sinica, 18(3): 189-194 (in Chinese)

Zhou ZH, Li Y, 1983.

A new myxomycete: Cribraria enodis

Acta Mycologica Sinica, 2(1): 38-40 (in Chinese)

邓叔群, 1963. 中国的真菌. 北京: 科学出版社. 5-46

[本文引用: 1]

李玉, 2002.

“鬼屎”考

吉林农业大学学报, 24(2): 1-4

[本文引用: 2]

李玉, 李惠中, 王琦, 陈双林, 2008a.

中国真菌志:黏菌卷一,鹅绒菌目、刺轴菌目、无丝菌目、团毛菌目

北京: 科学出版社. 1-238

[本文引用: 1]

李玉, 李惠中, 王琦, 陈双林, 2008b.

中国真菌志:黏菌卷二,绒泡菌目、发网菌目

北京: 科学出版社. 1-204

[本文引用: 1]

魏江春, 2012.

菌物生物多样性及其资源研发前景

菌物研究, 10(3): 125-129

[本文引用: 1]

周宗璜, 1981.

黏菌分类问题

微生物学通报, 8(3): 129-133

[本文引用: 1]

周宗璜, 李惠中, 1978.

对Skvortzow报告的黏菌新种的复核

微生物学报, 18(3): 189-194

[本文引用: 1]

周宗璜, 李玉, 1983.

黏菌一新种:无节筛菌

真菌学报, 2(1): 38-40

[本文引用: 1]

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