川西南凤仙花属Impatiens L.植物区系及亲缘关系研究
摘要
在前人工作的基础上,对川西南地区凤仙花属植物区系的组成、特点、性质和来源及部分物种的分子生物学特征等方面进行了研究,获得了以下结论：
1.川西南地区凤仙花属植物种类组成及分布
川西南地区共有凤仙花属植物65种,占我国凤仙花属植物总数的25.4%,约占世界凤仙花属植物总数的7.1%。该属植物在川西南地区的分布中,以峨眉山和稻城-盐源区段种类最为丰富,分别有26种和19种,各占该区凤仙花属植物总数的40.0%和29.2%。凤仙花属植物在该区的垂直分布特征显著,从海拔800—4200米均有该属植物分布,其中海拔1600—2700米之间的区段种类分布最多,有43种。川西南地区凤仙花属植物特有现象十分明显,有中国特有种58种,占该区凤仙花属植物总数的90.6%,其中川西南特有种34种,占该区凤仙花属植物总数的58.7%。
2.川西南地区凤仙花属植物区系成分与性质
川西南地区凤仙花属植物区系分属于我国15种植物区系成分中的3种,其中热带亚洲分布5种,占该区凤仙花属植物总数的7.8%,北温带分布1种,占该区凤仙花属植物总数的1.6%,中国特有分布58种,占该区凤仙花属植物总数的90.6%。研究表明川西南地区的64种凤仙花属植物,属于热带性质的有26种,属于温带性质的有38种,分别占川西南地区该属植物总数的40.6%和59.4%。表明川西南地区凤仙花属植物区系虽然具有较大比重的温带成分,但总体上属于热带与温带过渡的性质。
3.川西南地区凤仙花属植物区系的地位
由于独特的地理位置和自然环境,川西南地区孕育了丰富的凤仙花属植物,这里不仅物种多样性丰富,而且既有该属较原始的类型,也有过渡类型和较进化的类型。如在该区低海拔地区分布有具多年生、总状花序、四枚萼片、花粉三沟型和辐射对称等原始特征的峨眉凤仙花I. omeiana Hook. f.等种类,而在中高海拔地段分布有具一年生、两枚萼片、花粉四沟型和左右对称等进化特征的类群。该区凤仙花属植物形态分化强烈,特有现象十分突出。川西南地区凤仙花属植物区系的丰富多样,不仅表明该区凤仙花属植物区系具有一定的古老性,而且表明该区是我国凤仙花属植物的重要分布和分化区域,在我国乃至世界凤仙花属区系研究中占有重要地位。
4.川西南地区凤仙花属植物区系的来源
研究表明,川西南地区凤仙花属植物区系主要来源于两个方面：一是该区凤仙花属区系继承了丰富的古热带植物区系成分,如该区的峨眉凤仙花I. omeiana Hook. f.和白花凤仙花I. wilsonii Hook. f.等古特有种是其典型代表。二是川西南地区凤仙花属植物区系与东喜马拉雅区系(特别是云南、西藏凤仙花属区系)交流、渗透的结果。川西南凤仙花属植物区系与云南高黎贡山、玉龙雪山及西藏区系的高度相似性,表明该区凤仙花属植物区系也是东喜马拉雅植物区系产生了密切的交流渗透,这种交流渗透加快了该区凤仙花属植物的分化和演化,孕育、分化出了如华丽凤仙花I. faberi Hook. f.、四裂片凤仙花I. quadriloba K. M. Liu & Y. L. Xiang等众多新特有成分。使该区凤仙花属植物区系呈现出种类繁多、形态多样、新老兼备、特有现象突出的现代分布格局。
5.ITS序列研究结果
选取华丽凤仙花I. faberi、峨眉凤仙花I. omeiana等22种凤仙花属植物作为代表类群,以凤仙花属的姊妹属水角属的水角Hydrocera triflora和凤仙花科的近缘科的2个种(蜜囊花科Marcgraviaceae的Noran tea guianensi和四籽树科Tetrameristaceae的Tetramerista sp.)为外类群,根据ITS序列构建了两个和凤仙花属植物形态特征基本相吻合的系统发育树。该系统发育树可分为3个分支,第一支是由具有四枚萼片、花粉三沟型和辐射对称、子房四室等原始性状的峨眉凤仙花I. omeiana组成,其位于该系统发育树的基部,组成这一分支的种类可视为川西南地区凤仙花属植物的原始类群。第二支是由花序具多数花(4朵以上),果实长线形或长棒状的种类群组成,但该分支内的小分支缺乏较高的支持率。第三支是由具少数花(通常1-3朵)、果实长线形或长棒状的种类组成,该分支在性状分化上比较强烈,其种类在一定程度上代表了该区凤仙花属植物较进化的类型。
6.川西南地区凤仙花属植物区系新发现
本研究发现凤仙花属新种1个,被命名为四裂片凤仙花I. quadriloba K. M. Liu & Y. L. Xiang[论文发表于Nordic Journal of Botany,2010(丹麦,SCI源刊)]；发现四川省凤仙花属植物区系新纪录种多个。
The author studied the flora of Impatiens L. in Southwest Sichuan, including structure and floristic featuresd, characteristic and sources, and molecular characteristics et albased on the previous work. The main results are summarized as follows:
1. Species structure and distribution of Impatiens in Southwest Sichuan
There are 65 Impatiens species in Southwest Sichuan,25.4% of the total species in China,7.1% of the total species in the World. Impatiens species unevenly distribution in Southwest Sichuan, species in subregion of Mount Emei and Daocheng-Yanyuan are most abundant. Accounting for 26 species and for 40.0% of the total species in MountEmei areas and 19species and 29.2% of the total species in Daocheng-Yanyuan areas. Vertical distribution of the Impatiens species is evident in Southwest Sichuan, and they distribution between 800m and 4200m alt. But mainly distribute between 1600m and 2700m (43species). Endemic species are rich in Southwest Sichuan, there are 58 species (accounting for 90.6% of the total species) endemic to the China,34species (accounting for 58.7% of the total species) endemic to Southwest Sichuan.
2. Floristic composition and properties of Impatiens in Southwest Sichuan
The Impatiens flora of Southwest Sichuan can be divided into 3 types of 15 distribution types from Chinese seed plants flora, there are 5 species of tropical Asian distribution (accounting for 7.8% of the total species),1 specie of north temperate distribution (accounting for 1.6% of the total species) and 58 species endemic to the China (accounting for 90.6% of the total species). This studies shows that 64 species of Impatiens plants in Southwest Sichuan, there are 26 species to be tropic character (accounting for 40.6% of the total species) and 38 species to be temperate character (accounting for 59.4% of the total species), this shows that the Impatiens flora of Southwest Sichuan have a large proportion of temperate character, But it possess tropical to temperate transition.
3. Status of Impatiens flora in Southwest Sichuan
Because of the unique geographical location and environment, The region of Southwest Sichuan give birth to a wealth of Impatiens plant, where not only rich in species diversity and more primitive type of the genus, but also transition type and evolution type. There are perennial, racemes, four sepals, three pollen groove and radial symmetry, other original features in low altitude region of Southwest Sichuan, such as I. omeiana Hook. f. et al. There are annual, two sepals, four pollen groove and symmetrical, and other evolution features in high altitude locations of Southwest Sichuan. The Morphological differentiation of Impatiens is strong, and endemism is evey prominent. The diversity of Impatiens flora in Southwest Sichuan, not only shows that the Impatiens flora is ancient, but also shows that the Impatiens flora of the region is important distribution and differentiation of China, the Impatiens flora play an important role in China and the World.
4. Source of Impatiens flora in Southwest Sichuan
The studying shows that the Impatiens flora of the region mainly from two aspects:first, the areas inherited a rich of the ancient tropical flora, such as I. omeiana Hook. f. and I.wilsonii Hook, f., and other ancient endemic species are typical representative; second, the results of the areas exchange and penetration with Eastern Himalayan flora (especially Yunnan, Tibet Impatiens flora). The Impatiens flora of Southwest Sichuan is highly similar with Gaoligong Mountain, Yulong Snow Mountain and the flora of Tibet, which indicated that the region Impatiens flora and Himalayan flora have a close and communicate, thereby speeding up the area Impatiens evolution and differentiation, the region Impatiens flora shows a modern distribution pattern of the species diversity, varied shapes, both old and new, endemical phenomenon.
5. ITS sequence results
Based the nrDNA ITS of 22 species, which represent most of the Impatiens types, two phylogenetic trees are constructed using Hydrocera triflora, Norantea guianensi and Tetramerista sp. At out groups. Three clades are recognized in high posterior probability value. CladeⅠlocates the root of phylogenetic tree, specie of this clade, such as I. omeiana Hook. f., have mang primordial character, such as four lateral sepals, three colpate pollen and radial symmetry and four chamber ovary et al. This group should be the primordial Impatiens group in Southwest Sichuan. CladeⅡis clustered by species with long lineshaped or clavate fruit and inflorescence, having number flowers (usually more than four flowers), however, the small branch of the branch lack of a high support rate. CladeⅢis clustered by species with long lineshaped or clavate fruit and inflorescence, having Less flowers (usually one to three flowers), The branch is relatively strong in character differentiation, to some extent, this species represent more evolutionary type.
6. New dicovery of Impatiens flora in Southwest Sichuan
One new specie (I. quadriloba K. M. Liu & Y. L. Xiang published in Nordic Journal of Botany) and seven new record species of Impatiens are found.
1.川西南地区凤仙花属植物种类组成及分布
川西南地区共有凤仙花属植物65种,占我国凤仙花属植物总数的25.4%,约占世界凤仙花属植物总数的7.1%。该属植物在川西南地区的分布中,以峨眉山和稻城-盐源区段种类最为丰富,分别有26种和19种,各占该区凤仙花属植物总数的40.0%和29.2%。凤仙花属植物在该区的垂直分布特征显著,从海拔800—4200米均有该属植物分布,其中海拔1600—2700米之间的区段种类分布最多,有43种。川西南地区凤仙花属植物特有现象十分明显,有中国特有种58种,占该区凤仙花属植物总数的90.6%,其中川西南特有种34种,占该区凤仙花属植物总数的58.7%。
2.川西南地区凤仙花属植物区系成分与性质
川西南地区凤仙花属植物区系分属于我国15种植物区系成分中的3种,其中热带亚洲分布5种,占该区凤仙花属植物总数的7.8%,北温带分布1种,占该区凤仙花属植物总数的1.6%,中国特有分布58种,占该区凤仙花属植物总数的90.6%。研究表明川西南地区的64种凤仙花属植物,属于热带性质的有26种,属于温带性质的有38种,分别占川西南地区该属植物总数的40.6%和59.4%。表明川西南地区凤仙花属植物区系虽然具有较大比重的温带成分,但总体上属于热带与温带过渡的性质。
3.川西南地区凤仙花属植物区系的地位
由于独特的地理位置和自然环境,川西南地区孕育了丰富的凤仙花属植物,这里不仅物种多样性丰富,而且既有该属较原始的类型,也有过渡类型和较进化的类型。如在该区低海拔地区分布有具多年生、总状花序、四枚萼片、花粉三沟型和辐射对称等原始特征的峨眉凤仙花I. omeiana Hook. f.等种类,而在中高海拔地段分布有具一年生、两枚萼片、花粉四沟型和左右对称等进化特征的类群。该区凤仙花属植物形态分化强烈,特有现象十分突出。川西南地区凤仙花属植物区系的丰富多样,不仅表明该区凤仙花属植物区系具有一定的古老性,而且表明该区是我国凤仙花属植物的重要分布和分化区域,在我国乃至世界凤仙花属区系研究中占有重要地位。
4.川西南地区凤仙花属植物区系的来源
研究表明,川西南地区凤仙花属植物区系主要来源于两个方面：一是该区凤仙花属区系继承了丰富的古热带植物区系成分,如该区的峨眉凤仙花I. omeiana Hook. f.和白花凤仙花I. wilsonii Hook. f.等古特有种是其典型代表。二是川西南地区凤仙花属植物区系与东喜马拉雅区系(特别是云南、西藏凤仙花属区系)交流、渗透的结果。川西南凤仙花属植物区系与云南高黎贡山、玉龙雪山及西藏区系的高度相似性,表明该区凤仙花属植物区系也是东喜马拉雅植物区系产生了密切的交流渗透,这种交流渗透加快了该区凤仙花属植物的分化和演化,孕育、分化出了如华丽凤仙花I. faberi Hook. f.、四裂片凤仙花I. quadriloba K. M. Liu & Y. L. Xiang等众多新特有成分。使该区凤仙花属植物区系呈现出种类繁多、形态多样、新老兼备、特有现象突出的现代分布格局。
5.ITS序列研究结果
选取华丽凤仙花I. faberi、峨眉凤仙花I. omeiana等22种凤仙花属植物作为代表类群,以凤仙花属的姊妹属水角属的水角Hydrocera triflora和凤仙花科的近缘科的2个种(蜜囊花科Marcgraviaceae的Noran tea guianensi和四籽树科Tetrameristaceae的Tetramerista sp.)为外类群,根据ITS序列构建了两个和凤仙花属植物形态特征基本相吻合的系统发育树。该系统发育树可分为3个分支,第一支是由具有四枚萼片、花粉三沟型和辐射对称、子房四室等原始性状的峨眉凤仙花I. omeiana组成,其位于该系统发育树的基部,组成这一分支的种类可视为川西南地区凤仙花属植物的原始类群。第二支是由花序具多数花(4朵以上),果实长线形或长棒状的种类群组成,但该分支内的小分支缺乏较高的支持率。第三支是由具少数花(通常1-3朵)、果实长线形或长棒状的种类组成,该分支在性状分化上比较强烈,其种类在一定程度上代表了该区凤仙花属植物较进化的类型。
6.川西南地区凤仙花属植物区系新发现
本研究发现凤仙花属新种1个,被命名为四裂片凤仙花I. quadriloba K. M. Liu & Y. L. Xiang[论文发表于Nordic Journal of Botany,2010(丹麦,SCI源刊)]；发现四川省凤仙花属植物区系新纪录种多个。
The author studied the flora of Impatiens L. in Southwest Sichuan, including structure and floristic featuresd, characteristic and sources, and molecular characteristics et albased on the previous work. The main results are summarized as follows:
1. Species structure and distribution of Impatiens in Southwest Sichuan
There are 65 Impatiens species in Southwest Sichuan,25.4% of the total species in China,7.1% of the total species in the World. Impatiens species unevenly distribution in Southwest Sichuan, species in subregion of Mount Emei and Daocheng-Yanyuan are most abundant. Accounting for 26 species and for 40.0% of the total species in MountEmei areas and 19species and 29.2% of the total species in Daocheng-Yanyuan areas. Vertical distribution of the Impatiens species is evident in Southwest Sichuan, and they distribution between 800m and 4200m alt. But mainly distribute between 1600m and 2700m (43species). Endemic species are rich in Southwest Sichuan, there are 58 species (accounting for 90.6% of the total species) endemic to the China,34species (accounting for 58.7% of the total species) endemic to Southwest Sichuan.
2. Floristic composition and properties of Impatiens in Southwest Sichuan
The Impatiens flora of Southwest Sichuan can be divided into 3 types of 15 distribution types from Chinese seed plants flora, there are 5 species of tropical Asian distribution (accounting for 7.8% of the total species),1 specie of north temperate distribution (accounting for 1.6% of the total species) and 58 species endemic to the China (accounting for 90.6% of the total species). This studies shows that 64 species of Impatiens plants in Southwest Sichuan, there are 26 species to be tropic character (accounting for 40.6% of the total species) and 38 species to be temperate character (accounting for 59.4% of the total species), this shows that the Impatiens flora of Southwest Sichuan have a large proportion of temperate character, But it possess tropical to temperate transition.
3. Status of Impatiens flora in Southwest Sichuan
Because of the unique geographical location and environment, The region of Southwest Sichuan give birth to a wealth of Impatiens plant, where not only rich in species diversity and more primitive type of the genus, but also transition type and evolution type. There are perennial, racemes, four sepals, three pollen groove and radial symmetry, other original features in low altitude region of Southwest Sichuan, such as I. omeiana Hook. f. et al. There are annual, two sepals, four pollen groove and symmetrical, and other evolution features in high altitude locations of Southwest Sichuan. The Morphological differentiation of Impatiens is strong, and endemism is evey prominent. The diversity of Impatiens flora in Southwest Sichuan, not only shows that the Impatiens flora is ancient, but also shows that the Impatiens flora of the region is important distribution and differentiation of China, the Impatiens flora play an important role in China and the World.
4. Source of Impatiens flora in Southwest Sichuan
The studying shows that the Impatiens flora of the region mainly from two aspects:first, the areas inherited a rich of the ancient tropical flora, such as I. omeiana Hook. f. and I.wilsonii Hook, f., and other ancient endemic species are typical representative; second, the results of the areas exchange and penetration with Eastern Himalayan flora (especially Yunnan, Tibet Impatiens flora). The Impatiens flora of Southwest Sichuan is highly similar with Gaoligong Mountain, Yulong Snow Mountain and the flora of Tibet, which indicated that the region Impatiens flora and Himalayan flora have a close and communicate, thereby speeding up the area Impatiens evolution and differentiation, the region Impatiens flora shows a modern distribution pattern of the species diversity, varied shapes, both old and new, endemical phenomenon.
5. ITS sequence results
Based the nrDNA ITS of 22 species, which represent most of the Impatiens types, two phylogenetic trees are constructed using Hydrocera triflora, Norantea guianensi and Tetramerista sp. At out groups. Three clades are recognized in high posterior probability value. CladeⅠlocates the root of phylogenetic tree, specie of this clade, such as I. omeiana Hook. f., have mang primordial character, such as four lateral sepals, three colpate pollen and radial symmetry and four chamber ovary et al. This group should be the primordial Impatiens group in Southwest Sichuan. CladeⅡis clustered by species with long lineshaped or clavate fruit and inflorescence, having number flowers (usually more than four flowers), however, the small branch of the branch lack of a high support rate. CladeⅢis clustered by species with long lineshaped or clavate fruit and inflorescence, having Less flowers (usually one to three flowers), The branch is relatively strong in character differentiation, to some extent, this species represent more evolutionary type.
6. New dicovery of Impatiens flora in Southwest Sichuan
One new specie (I. quadriloba K. M. Liu & Y. L. Xiang published in Nordic Journal of Botany) and seven new record species of Impatiens are found.
引文
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