中国石韦属植物系统分类学研究
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摘要
石韦属Pyrrosia Mirbel隶属于水龙骨科Polypodiaceae、石韦亚科Pyrrosioideae,全世界约有70种,是水龙骨科中较大的属。该属植物分布于旧大陆,东亚和东南亚是其主要分布区。该属植物一般附生在树上或岩石上,少数土生,是蕨类植物中比较进化的一类中小型蕨类。石韦属自成立以来,.以其特殊的脉序和星芒状的毛被而成为一个自然的类群,从而与水龙骨科其它属相区别,是研究水龙骨科系统分类的关键类群之一。
本文在文献考证及标本研究的基础上,从鳞片、脉序、毛被、孢粉纹饰、矿质元素等方面,对石韦属植物进行系统分类学研究,提出一个新的属下分类系统,确定中国产石韦属植物共27种、3变种、2变型、1存疑种。9个名称处理为新异名。
1鳞片解剖
对石韦属38种(《中国植物志》资料)代表植物的鳞片类型、形状、颜色、着生方式和边缘特征进行研究。本属植物的鳞片具有3种着生方式：盾状着生、拟盾状着生和基部着生；4种形状：披针形渐尖、披针形尾尖、卵钻形和卵圆形；3种颜色：浅棕色、深棕色及近黑色(颜色之间有一定的过渡,与植物的生长状态和生长阶段有关)；4种边缘：全缘、睫状齿、长纤毛和顶端毛发状。
2脉序类型
研究了石韦属10种代表植物和水龙骨科其它5属6种植物的脉序类型。结果表明,水龙骨科植物叶脉分为分离叶脉和结合叶脉,石韦属植物的脉序属于结合脉序,但与被研究的水龙骨科其他属的结合脉序不同。分为两种：弯脉蕨型与拟槲蕨型。前者与特产于美洲的水龙科弯脉蕨属脉序相同,可能二属有一定的亲缘关系。
3毛被特征
主要对石韦属38个种的叶背面毛被进行了显微和扫描电镜观察。石韦属植物毛被由特殊的星状毛组成。星状毛分支臂形状有3种类型：披针形、针状、卷曲绒毛状,分别为浅棕色、深棕色和白色；分支臂类型有2种：一型(分支臂仅1种形状)和二型(分支臂有2种形状,即同轴上长出披针形和长细丝状两种分支臂)；这些不同分支臂的星状毛相互组合,将石韦属叶下表面毛被分为以下5种类型：Ⅰ.一种披针形臂毛被；Ⅱ.一种针状臂毛被；Ⅲ.一种分支臂二型毛被；Ⅳ两种针状和披针形毛被；V.两种针状和卷曲绒毛状毛被。这些特征性性状对石韦属部分种类的分类鉴定具有一定的意义。
4孢粉特征
光镜和扫描电镜下观察石韦属19种植物的孢子特征,石韦属孢子黄色,形态均为肾形,两侧对称,单裂缝,说明该属植物是一个自然类群。表面纹饰类型有3种：Pyrrosia rupestris-type、Pyrrosia subfurfuracea-type及Pyrrosia christii-type。孢子表面纹饰特征相对稳定,在种间存在较大差异,可以为属下分类提供重要依据。
5矿质元素
采用原子吸收分光光度法(AAS)测定采自中国云南石韦属9种植物样品的矿物元素含量,分析表明,矿物元素含量在石韦属植物种间存在大幅度的变化,意味着该属每种植物都有一个固定的矿物元素的吸收和积累过程,从而可以控制植物药材及其制剂的质量、扩大天然药物的资源,并从化学分类学的角度进一步研究石韦属植物的系统分类,为研究蕨类植物的自然演化关系和化学分类学提供新的资料。
6属下分类
根据孢粉、叶形、叶脉和毛被等特征将石韦属分为2个亚属和6个组。2亚属：石韦亚属subgen. Pyrrosia和厚毛石韦亚属subgen. Porosae S. G. Lu et L. H. Yang, subgen. nov.;6组：钱币石韦组sect. Nummulariifoliae S. G. Lu et L. H. Yang, sect. nov.,石韦组sect. Pyrrosia,披针石韦组sect. Lanceolatae S. G. Lu et L. H. Yang, sect. nov.,毡毛石韦组sect. Drakeanae (Ching et Shing) S. G. Lu et L. H. Yang, stat. nov.,厚毛石韦组sect. Porosae,下延石韦组sect. Costatae (Ching et Shing) S. G. Lu et L. H. Yang, stat. nov.。综合这些特征列出属下检索表。
7种类修订
通过对国内外各大标本馆(PE, PYU, KUN, IBSC, HITBC, BM, K)馆藏标本和模式标本的研究考证,结合野外及栽培观察,综合分析有关分类学资料,对《中国植物志》上记载的38种中国产石韦属植物进行了新的分类学修订。经本次修订,确认中国产石韦属植物有27种、3变种、2变型,分属于6个组；9个名称处理为新异名：尾叶石韦P. caudifrons和佛海石韦P. fuohaiensis处理为纸质石韦P. heteractis的异名；冯氏石韦P. fengiana、相似石韦P. similis、波氏石韦P. bonii处理为布氏石韦P. boothii的异名；西南石韦P. gralla处理为华北石韦P. davidii的异名,并将其新拟为中国石韦；黄山石韦P. dimorpha处理为相近石韦P. assimilis的异名；钙生石韦P. adnascens f. calcicola处理为贴生石韦P. adnascens的异名；平绒石韦P. porosa var. mollissima处理为柔软石韦P. porosa的异名。3个名称下降为变种：贴生石韦P. adnascens和裸叶石韦P. nuda降为披针叶石韦P. lanceolata的变种；神农石韦P. shennongensis降为布氏石韦P. boothii的变种。2个降为变型：剑叶石韦P. ensata降为石韦P. lingua的变型；拟毡毛石韦P. pseudodrakeana Shing降为毡毛石韦P. drakeana的变型,另有1个种暂时存疑。
About70species of the genus Pyrrosia Mirbel (Polypodiaceae, Pyrrosioideae) are mainly distributed in the mainland of Asia, epiphytic or epilithic, sometimes terrestrial. The genus Pyrrosia Mirbel is one of the key groups in Polypodiaceae, and it is different from other genera of Polypodiaceae in special venation pattern and indumenta.
Based on the collection of literatures and checked in the specimens in herbaria, the scale, lamina indumenta, venation pattern, spore ornamentation and mineral element of the genus Pyrrosia were investigated in the present study. A new infrageneric system was supported, and all Chinese species were treated at27species,3variety and2forms in this genus. The major results of the dissertation were summarized as follows:
1Rhizome scale
The characters of scale of38species of the genus Pyrrosia were observed under the anatomical lens. It was showed there were3attachment, such as peltate, pseudopeltate and basifix;4shape types, such as lanceolate-acuminate, lanceolate-caudate, ovate-subulate and ovate;3colors, such as light brown, dark brown, brown-black and some intergradations (concerned growing age);4marginal types, including entire, dentate, ciliolate, and hair at apex.
2Venation pattern
The characters of venation pattern of10species of Pyrrosia and6species in other5genera were studied in Polypodiaceae. It was showed that free and anatomizing venation pattern were existed in Polypodiaceae. There were2venation patterns of Pyrrosia, Campyloneuroid type and Drynarioid type, included in anatomizing venation pattern. It was implied there be a evolutionary relationship between Pyrrosia and Campyloneurum.
3Lamina indument
The characters of lamina indument of38species of Pyrrosia were examined under anatomical lens and scanning electron microscopy (SEM). It was showed the hairs of the fronds were almost exclusively stellate with being composed of different rays. There were3shapes of rays, including lanceolate (light brown), aciculate (dark brown) and wooly (white);2types of rays, such as monomorphic and dimorphic. So,5types of indument were discovered in the Pyrrosia. These characters could be used to distinguish some species.
4Spore ornamentation
The spores of19taxa of the genus Pyrrosia were examined by means of light microscopy (LM) and SEM. It was showed the shapes of the spores were all kidney-shaped, bilaterally symmetric, monolcte, while the color of the spores was yellow. There were two main ornamentation types, such as verrucate and tuberculate. The spore morphological features were polymorphism, including Pyrrosia rupestris-type, Pyrrosia subfurfuracea-types and Pyrrosia christii-type. The evidence from spore ornamentations was stable, and could be used to differentiate the affined species in this genus.
5Mineral element
In this dissertation, the contents of9mineral elements in9species of Pyrrosia were determined by atomic absorption spectrophotometry (AAS). It was provided for baseline data on mineral element levels in fern genus Pyrrosia. The element contents change in wide range. It implied that the species should have an inherent characteristic absorption and accumulation for the determined mineral elements, and might control quality of herbal medicines, to provide chemotaxonomy information.
6Infrageneric system
Based on the characters of spore ornamentations, the shape of blade, venation pattern, scale, lamina indumenta, etc.. two subgenera and six sections of the genus were presented:subgen. Pyrrosia, subgen. Porosae S. G. Lu et L. H. Yang; sect. Nummulariifoliae S. G. Lu et L. H. Yang. sect. Pyrrosia. sect. Lanceolalae S. G. Lu et L. H. Yang, sect. Drakeanae (Ching et Shing) S. G. Lu et L. II. Yang. sect. Porosae, sect. Costalae (Ching et Shing) S. G. Lu et l..H. Yang. A key to the Pyrrosia species was given.
7Taxonomic revision
Based on the collection of literatures, checked in the specimens and type specimens in Herbaria all over the world (PE, PYU, KUN, IBSC, HITBC, BM, K), combining field, culture and morphological observation, all species in this genus Pyrrosia from China were made classification. As a result,27species,3variety,2forms were recognized from China in six sections. Nine species were treated as synonyms, such as P. caudifrons and P. fuohaiensis were treated as the synonym of P. heteractis; P. fengiana, P. bonii, and P. similis were treated as the synonym of P. bolhii; P. gralla was treated as the synonym of P. davidii; P. dimorpha was treated as the synonym of P. assimilis; P. adnascens f. calcicola was treated as the synonym of P. adnascens; P. porosa var. mollissima was treated as the synonym of P. porosa.3species was reduced to variety, i. e. P. adnascens and P. mida were reduced to the variety of P lanceolata:P. shennongensis was reduced to the variety of P. boothii.2species were reduced to forms, i. e. P. ensata was reduced to the form of P. lingua; P. pseudodrakeana was reduced to the form of P. drakeana, and1species were temperately treated as doubtful species in the present study.
本文在文献考证及标本研究的基础上,从鳞片、脉序、毛被、孢粉纹饰、矿质元素等方面,对石韦属植物进行系统分类学研究,提出一个新的属下分类系统,确定中国产石韦属植物共27种、3变种、2变型、1存疑种。9个名称处理为新异名。
1鳞片解剖
对石韦属38种(《中国植物志》资料)代表植物的鳞片类型、形状、颜色、着生方式和边缘特征进行研究。本属植物的鳞片具有3种着生方式：盾状着生、拟盾状着生和基部着生；4种形状：披针形渐尖、披针形尾尖、卵钻形和卵圆形；3种颜色：浅棕色、深棕色及近黑色(颜色之间有一定的过渡,与植物的生长状态和生长阶段有关)；4种边缘：全缘、睫状齿、长纤毛和顶端毛发状。
2脉序类型
研究了石韦属10种代表植物和水龙骨科其它5属6种植物的脉序类型。结果表明,水龙骨科植物叶脉分为分离叶脉和结合叶脉,石韦属植物的脉序属于结合脉序,但与被研究的水龙骨科其他属的结合脉序不同。分为两种：弯脉蕨型与拟槲蕨型。前者与特产于美洲的水龙科弯脉蕨属脉序相同,可能二属有一定的亲缘关系。
3毛被特征
主要对石韦属38个种的叶背面毛被进行了显微和扫描电镜观察。石韦属植物毛被由特殊的星状毛组成。星状毛分支臂形状有3种类型：披针形、针状、卷曲绒毛状,分别为浅棕色、深棕色和白色；分支臂类型有2种：一型(分支臂仅1种形状)和二型(分支臂有2种形状,即同轴上长出披针形和长细丝状两种分支臂)；这些不同分支臂的星状毛相互组合,将石韦属叶下表面毛被分为以下5种类型：Ⅰ.一种披针形臂毛被；Ⅱ.一种针状臂毛被；Ⅲ.一种分支臂二型毛被；Ⅳ两种针状和披针形毛被；V.两种针状和卷曲绒毛状毛被。这些特征性性状对石韦属部分种类的分类鉴定具有一定的意义。
4孢粉特征
光镜和扫描电镜下观察石韦属19种植物的孢子特征,石韦属孢子黄色,形态均为肾形,两侧对称,单裂缝,说明该属植物是一个自然类群。表面纹饰类型有3种：Pyrrosia rupestris-type、Pyrrosia subfurfuracea-type及Pyrrosia christii-type。孢子表面纹饰特征相对稳定,在种间存在较大差异,可以为属下分类提供重要依据。
5矿质元素
采用原子吸收分光光度法(AAS)测定采自中国云南石韦属9种植物样品的矿物元素含量,分析表明,矿物元素含量在石韦属植物种间存在大幅度的变化,意味着该属每种植物都有一个固定的矿物元素的吸收和积累过程,从而可以控制植物药材及其制剂的质量、扩大天然药物的资源,并从化学分类学的角度进一步研究石韦属植物的系统分类,为研究蕨类植物的自然演化关系和化学分类学提供新的资料。
6属下分类
根据孢粉、叶形、叶脉和毛被等特征将石韦属分为2个亚属和6个组。2亚属：石韦亚属subgen. Pyrrosia和厚毛石韦亚属subgen. Porosae S. G. Lu et L. H. Yang, subgen. nov.;6组：钱币石韦组sect. Nummulariifoliae S. G. Lu et L. H. Yang, sect. nov.,石韦组sect. Pyrrosia,披针石韦组sect. Lanceolatae S. G. Lu et L. H. Yang, sect. nov.,毡毛石韦组sect. Drakeanae (Ching et Shing) S. G. Lu et L. H. Yang, stat. nov.,厚毛石韦组sect. Porosae,下延石韦组sect. Costatae (Ching et Shing) S. G. Lu et L. H. Yang, stat. nov.。综合这些特征列出属下检索表。
7种类修订
通过对国内外各大标本馆(PE, PYU, KUN, IBSC, HITBC, BM, K)馆藏标本和模式标本的研究考证,结合野外及栽培观察,综合分析有关分类学资料,对《中国植物志》上记载的38种中国产石韦属植物进行了新的分类学修订。经本次修订,确认中国产石韦属植物有27种、3变种、2变型,分属于6个组；9个名称处理为新异名：尾叶石韦P. caudifrons和佛海石韦P. fuohaiensis处理为纸质石韦P. heteractis的异名；冯氏石韦P. fengiana、相似石韦P. similis、波氏石韦P. bonii处理为布氏石韦P. boothii的异名；西南石韦P. gralla处理为华北石韦P. davidii的异名,并将其新拟为中国石韦；黄山石韦P. dimorpha处理为相近石韦P. assimilis的异名；钙生石韦P. adnascens f. calcicola处理为贴生石韦P. adnascens的异名；平绒石韦P. porosa var. mollissima处理为柔软石韦P. porosa的异名。3个名称下降为变种：贴生石韦P. adnascens和裸叶石韦P. nuda降为披针叶石韦P. lanceolata的变种；神农石韦P. shennongensis降为布氏石韦P. boothii的变种。2个降为变型：剑叶石韦P. ensata降为石韦P. lingua的变型；拟毡毛石韦P. pseudodrakeana Shing降为毡毛石韦P. drakeana的变型,另有1个种暂时存疑。
About70species of the genus Pyrrosia Mirbel (Polypodiaceae, Pyrrosioideae) are mainly distributed in the mainland of Asia, epiphytic or epilithic, sometimes terrestrial. The genus Pyrrosia Mirbel is one of the key groups in Polypodiaceae, and it is different from other genera of Polypodiaceae in special venation pattern and indumenta.
Based on the collection of literatures and checked in the specimens in herbaria, the scale, lamina indumenta, venation pattern, spore ornamentation and mineral element of the genus Pyrrosia were investigated in the present study. A new infrageneric system was supported, and all Chinese species were treated at27species,3variety and2forms in this genus. The major results of the dissertation were summarized as follows:
1Rhizome scale
The characters of scale of38species of the genus Pyrrosia were observed under the anatomical lens. It was showed there were3attachment, such as peltate, pseudopeltate and basifix;4shape types, such as lanceolate-acuminate, lanceolate-caudate, ovate-subulate and ovate;3colors, such as light brown, dark brown, brown-black and some intergradations (concerned growing age);4marginal types, including entire, dentate, ciliolate, and hair at apex.
2Venation pattern
The characters of venation pattern of10species of Pyrrosia and6species in other5genera were studied in Polypodiaceae. It was showed that free and anatomizing venation pattern were existed in Polypodiaceae. There were2venation patterns of Pyrrosia, Campyloneuroid type and Drynarioid type, included in anatomizing venation pattern. It was implied there be a evolutionary relationship between Pyrrosia and Campyloneurum.
3Lamina indument
The characters of lamina indument of38species of Pyrrosia were examined under anatomical lens and scanning electron microscopy (SEM). It was showed the hairs of the fronds were almost exclusively stellate with being composed of different rays. There were3shapes of rays, including lanceolate (light brown), aciculate (dark brown) and wooly (white);2types of rays, such as monomorphic and dimorphic. So,5types of indument were discovered in the Pyrrosia. These characters could be used to distinguish some species.
4Spore ornamentation
The spores of19taxa of the genus Pyrrosia were examined by means of light microscopy (LM) and SEM. It was showed the shapes of the spores were all kidney-shaped, bilaterally symmetric, monolcte, while the color of the spores was yellow. There were two main ornamentation types, such as verrucate and tuberculate. The spore morphological features were polymorphism, including Pyrrosia rupestris-type, Pyrrosia subfurfuracea-types and Pyrrosia christii-type. The evidence from spore ornamentations was stable, and could be used to differentiate the affined species in this genus.
5Mineral element
In this dissertation, the contents of9mineral elements in9species of Pyrrosia were determined by atomic absorption spectrophotometry (AAS). It was provided for baseline data on mineral element levels in fern genus Pyrrosia. The element contents change in wide range. It implied that the species should have an inherent characteristic absorption and accumulation for the determined mineral elements, and might control quality of herbal medicines, to provide chemotaxonomy information.
6Infrageneric system
Based on the characters of spore ornamentations, the shape of blade, venation pattern, scale, lamina indumenta, etc.. two subgenera and six sections of the genus were presented:subgen. Pyrrosia, subgen. Porosae S. G. Lu et L. H. Yang; sect. Nummulariifoliae S. G. Lu et L. H. Yang. sect. Pyrrosia. sect. Lanceolalae S. G. Lu et L. H. Yang, sect. Drakeanae (Ching et Shing) S. G. Lu et L. II. Yang. sect. Porosae, sect. Costalae (Ching et Shing) S. G. Lu et l..H. Yang. A key to the Pyrrosia species was given.
7Taxonomic revision
Based on the collection of literatures, checked in the specimens and type specimens in Herbaria all over the world (PE, PYU, KUN, IBSC, HITBC, BM, K), combining field, culture and morphological observation, all species in this genus Pyrrosia from China were made classification. As a result,27species,3variety,2forms were recognized from China in six sections. Nine species were treated as synonyms, such as P. caudifrons and P. fuohaiensis were treated as the synonym of P. heteractis; P. fengiana, P. bonii, and P. similis were treated as the synonym of P. bolhii; P. gralla was treated as the synonym of P. davidii; P. dimorpha was treated as the synonym of P. assimilis; P. adnascens f. calcicola was treated as the synonym of P. adnascens; P. porosa var. mollissima was treated as the synonym of P. porosa.3species was reduced to variety, i. e. P. adnascens and P. mida were reduced to the variety of P lanceolata:P. shennongensis was reduced to the variety of P. boothii.2species were reduced to forms, i. e. P. ensata was reduced to the form of P. lingua; P. pseudodrakeana was reduced to the form of P. drakeana, and1species were temperately treated as doubtful species in the present study.
引文
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李春香,陆树刚,杨群.2004.蕨类植物起源与系统发生关系研究进展.植物学通报,19：478-485
李春香,陆树刚.2005.龙骨星蕨与膜叶星蕨的关系：来自叶绿体rbcL、rps4和trnL-trnF rps4-trnS序列的证据.生物多样性.13(2)：174-179
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李晓丹,肖娅萍,孙蓉蓉.2008.秦岭水龙骨科和剑蕨科植物孢子形态研究.内蒙古师范大学学报.37(3)：422-426.
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林尤兴.2000.石韦属.见：中国科学院中国植物志编辑委员会.中国植物志,6(2)[M].北京：科学出版社：116-151
刘红梅等.2008.石松类和蕨类植物研究进展：兼论国产类群的科级分类系统.植物分类学报， 46(6)：808-829.
刘家熙等.1998.北京水龙骨科孢子形态研究.植物学通报,15(1)：68-71.
鲁如坤等.1998.土城植物营养学原理和施肥[M].北京：化学工业出版社.
陆树刚.2007.蕨类植物学[M].北京：高等教育出版社.
陆树刚,李春香.2006.用叶绿体rbcL和rps4-trnS区序列确定亚洲特有单型属——篦齿蕨属的系统位置.植物分类学报,44(5)：494-502.
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王家玉.1992.植物营养元素交互作用研究.土壤学进展,20(2)：1-6.
王全喜,丁晶.2003.扫描电镜下真蕨目孢子表面纹饰的分类.云南植物研究,25(3)：313-320.
王任翔,陆树刚,邓晰朝等.2006.广西蕨类植物孢子形态的研究Ⅰ.水龙骨科,广西植物,26(5)：565-569
王任翔.2007.中国水龙骨科植物的系统学研究[D].昆明：云南大学.
王中仁,夏群.1984.一些中国产鳞毛蕨属和耳蕨属植物的细胞学观察.植物学报,26：24-27
王中仁,夏群,张志宪.1984.25种中国蕨类植物的细胞学观察.植物学报,26：595-604
文瑞良,伍兰.2000.相异石韦性状及显微鉴别.中药材,23(8)：446-447
吴兆洪,秦仁吕.1991.中国蕨类植物科属志[M].北京：科学出版社：521-523
谢语言·湖,江丰.2009HPLC法测定不同产地中药石韦的绿原酸含量.江西化工，2：62-64.
徐彦,陈佳正，吴春蕾等.2010.11种常用中药材在石蜡切片中的软化处理方法.西南民族大学学报(自然科学版),36(4)：603-605.
张碧波,王任翔,常艳芬,陆树刚.2006.中国西南地区水龙骨属植物(水龙骨科)孢子形态研究.武汉植物研究,24(2)：113-118.
张健.2009.丁香属(Syringa)一些植物形态解剖与园林应用探讨[D].福建：福建农业大学
张金谈.1979.从孢粉形态特征试论植物某些类群的分类与系统发育.植物分类学报,17(2)：1-8.
张金渝,王元忠,金航等.ICP-AES法测定滇重楼中的微量元素.光谱学与光谱分析,29(8)：2247-2249.
张晓林等.2001.土壤-烤烟矿质营养元素相互关系的主组分分析.土壤学报.38(2)：193-203.
张耀甲，于海峰,卢云霞等.1999.国产水龙骨科植物的气孔器类型及其系统学意义.兰州大学报，35(1)：130-139.
张玉龙,席以珍,张金谈等.1976.中国蕨类植物孢子形态[M].北京：科学出版社,363-365.
赵惠玲,王青,李晓玲.2002.木质化干枯植物材料的整体PAS反应及制片法.山西大学学报(自然科学版),26(3)：258-259.
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