青藏高原东缘种子大小变异的进化生态学研究
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摘要
对青藏高原东缘106科412属1130种被子植物种子大小的变异模式、种子大小变异与系统发育、植物分布区类型、植物生活史特征、植被类型之间的相关性进行了分析研究,并从不同角度分析了海拔对种子大小变异的影响,以期揭示青藏高原东缘种子大小变异的进化及生态适应意义。结果如下：
1、青藏高原东缘区系被子植物的种子大小呈现正态分布,种子大小从10-3mg到104mg相差约为6～7个数量级。种子大小主要集中在0.1～10.0mg范围之内,占到总种数的70.53%,属于小种子占优势的区系。整体上种子大小属于中度大小的种子的物种较多,而大种子和小种子的物种较少的分布格局类型。
2、青藏高原东缘区系中,分类学水平纲、目、科均显著影响种子大小,其中,纲能解释11.6%的种子大小变异,目能解释31.4%,科能解释58.2%,与影响和控制种子大小的其它因素相比,科所占的比例最高,因而系统发育是决定种子大小的一个关键因子。
3、无论纲、目、科级分类学水平,种子大小的变异都没有表现出一致的演化方向或演进模式,基本上是靠近基部类群的种子居于中间大小,随着演化水平提高,种子大小向大或者向小的方向变化发展。因而,现存被子植物的种子大小具有明显的适应辐射的演化特点。
4、植物分布区类型不同,种子大小变异存在较大的差异。一般来说,古老类群具有相对较大的种子大小,而年轻类群的种子较小。
5、种子大小变异与植物生活史各种性状如种子扩散方式、植物功能类群、植物生活型、植株高度、寿命、果实类型、开花季节、花期持续时间长短都存在一定的相关性,即使除去系统发育的影响。其中对种子大小变异影响最大的生活史性状是果实类型,其次是扩散模式和植株高度。
6、植被类型也可以显著地影响青藏高原东缘植物区系种子大小的变异。植被类型中种子大小从大到小依次是森林、灌丛、农田、草甸及草地、水生植被、高山流石滩植被、垫状植被、沼泽。
7、青藏高原东缘区系种子大小与海拔呈极显著的线性负相关性。但从整体上看,在海拔3100m处的种子大小居中,随海拔降低或升高,种子分别趋向于变大或变小的方向变化,即种子大小具有适应辐射演化的特点。
8、海拔梯度变化对不同扩散模式、不同分布区类型、不同生活史特征、不同植被类型及各科内种子大小变异的影响没有统一的模式,但存在一个共同的特点就是,无论从哪个方面分析,种子大小都有适应辐射的特点,即以3100m为起点,随海拔降低或升高,种子分别趋向于变大或变小的方向变化,具有明显的趋异分化的特点。
Seed size is an important feature of the attribute constellation of a plant species. Under the pressures of long-term selection and evolutionary, most plants have different seed size to adapt to the habitat and ensure the viability of offspring. As an important biological character in botany and ecology, seed size has relationship with other characters of plant life history, including dispersal capacity of seed, life form, plant individuals in a community, the trait of seed germination, seeding livability, the dispersal mode of seed, etc. In this study, the fresh seed size of 1130 angiosperm species which belong to 412 genera in 106 families were collected from the eastern Qinghai-Tibet plateau. We examined the pattern of the seed size variation in the flora, the relationship of seed size with phylogeny, areal-types of plants, dispersal mode, growth form, plant height, perenniality, fruit type, onset of flowering, duration of flowering, habitat and altitude.The results were shown:
1. Within the flora of eastern Qinghai-Tibet plateau, the frequency of seed size classes on a log scale produced a normal distribution.The seed size is ranged from 10-3 to 104mg, which has 6-7orders of magnitude.In addition, the mostly seed size is mainly concentrated from 0.1 to 10.0mg, which hold 70.53%.Especially large or small seeds are very scarce in the flora. So we consider that small seed is preponderant on the eastern Qinghai-Tibetan Plateau.
2.Seed size was strong related to phylogeny. Class, order and family membership accounted for most of the variation in log seed size(11.6%,31.4%and 58.2%, respectively).Therefore, the seed size variation was determined mainly by the phylogeny.
3.From whaterever taxonomic levels, seed size variation showed no consistent direction of evolution or evolutionary model.Generally, the seeds near the base of APGⅡsystem are likely to be in the middle size. Trends in the evolution of seed size has gone in many directions,towards larger or smaller seeds.Thus, the seed size of angiosperm has the obvious characteristics of adaptive radiation.
4.With different areal-types of plants, the seed size variation is significant difference. The primitive families and genera have the larger seeds, and the more evolved families and genera have the smaller seeds relatively in general.
5.Seed size was correlated with many other plant attributes such as dispersal mode, growth form, plant height, perenniality, fruit type, onset of flowering and duration of flowering, respectively,even in after phylogenic effect. The strongest of the plant attributes in seed size are fruit type, and dispersal mode, followed by plant height and growth form.
6. Seed size variation is also significantly associated with vegetation types.The mean log scale seed size from large to small in different vegetation types is forests, shrubs, cultivated vegetation, steppes and medows, aquatic vegetation, open vegetation on flowing rock hillside, cushion vegetation and swamp vegetation.
7. There is significantly negative relationship between the variation in seed size and altitude in the flora, and seed size decreases with altitude on the whole.But one interesting phenomenon is that the seeds with middle size are distrbuted at 3100m, and seed size becomes to larger or smaller respectively along with the altitude increasing or decreasing respectively. That is to say, the seed size has the evolutionary pattern of adaptive radiation.
8.The effect of altitude on seed size variation has different patterns among dispersal modes, areal-types of plants, growth forms, fruit types, different plant height, different perennialities, vegetation types, families, and even among species in a family. But there is a commen feature:from every aspect to examine effect of altitude on seed size variation, it seems that the seed size has a vertical distribution implications of adaptive radiation. That is, taking the altitude of 3100m as a starting point, the seed size becomes to larger or smaller respectively along with the altitude increasing or decreasing respectively.
1、青藏高原东缘区系被子植物的种子大小呈现正态分布,种子大小从10-3mg到104mg相差约为6～7个数量级。种子大小主要集中在0.1～10.0mg范围之内,占到总种数的70.53%,属于小种子占优势的区系。整体上种子大小属于中度大小的种子的物种较多,而大种子和小种子的物种较少的分布格局类型。
2、青藏高原东缘区系中,分类学水平纲、目、科均显著影响种子大小,其中,纲能解释11.6%的种子大小变异,目能解释31.4%,科能解释58.2%,与影响和控制种子大小的其它因素相比,科所占的比例最高,因而系统发育是决定种子大小的一个关键因子。
3、无论纲、目、科级分类学水平,种子大小的变异都没有表现出一致的演化方向或演进模式,基本上是靠近基部类群的种子居于中间大小,随着演化水平提高,种子大小向大或者向小的方向变化发展。因而,现存被子植物的种子大小具有明显的适应辐射的演化特点。
4、植物分布区类型不同,种子大小变异存在较大的差异。一般来说,古老类群具有相对较大的种子大小,而年轻类群的种子较小。
5、种子大小变异与植物生活史各种性状如种子扩散方式、植物功能类群、植物生活型、植株高度、寿命、果实类型、开花季节、花期持续时间长短都存在一定的相关性,即使除去系统发育的影响。其中对种子大小变异影响最大的生活史性状是果实类型,其次是扩散模式和植株高度。
6、植被类型也可以显著地影响青藏高原东缘植物区系种子大小的变异。植被类型中种子大小从大到小依次是森林、灌丛、农田、草甸及草地、水生植被、高山流石滩植被、垫状植被、沼泽。
7、青藏高原东缘区系种子大小与海拔呈极显著的线性负相关性。但从整体上看,在海拔3100m处的种子大小居中,随海拔降低或升高,种子分别趋向于变大或变小的方向变化,即种子大小具有适应辐射演化的特点。
8、海拔梯度变化对不同扩散模式、不同分布区类型、不同生活史特征、不同植被类型及各科内种子大小变异的影响没有统一的模式,但存在一个共同的特点就是,无论从哪个方面分析,种子大小都有适应辐射的特点,即以3100m为起点,随海拔降低或升高,种子分别趋向于变大或变小的方向变化,具有明显的趋异分化的特点。
Seed size is an important feature of the attribute constellation of a plant species. Under the pressures of long-term selection and evolutionary, most plants have different seed size to adapt to the habitat and ensure the viability of offspring. As an important biological character in botany and ecology, seed size has relationship with other characters of plant life history, including dispersal capacity of seed, life form, plant individuals in a community, the trait of seed germination, seeding livability, the dispersal mode of seed, etc. In this study, the fresh seed size of 1130 angiosperm species which belong to 412 genera in 106 families were collected from the eastern Qinghai-Tibet plateau. We examined the pattern of the seed size variation in the flora, the relationship of seed size with phylogeny, areal-types of plants, dispersal mode, growth form, plant height, perenniality, fruit type, onset of flowering, duration of flowering, habitat and altitude.The results were shown:
1. Within the flora of eastern Qinghai-Tibet plateau, the frequency of seed size classes on a log scale produced a normal distribution.The seed size is ranged from 10-3 to 104mg, which has 6-7orders of magnitude.In addition, the mostly seed size is mainly concentrated from 0.1 to 10.0mg, which hold 70.53%.Especially large or small seeds are very scarce in the flora. So we consider that small seed is preponderant on the eastern Qinghai-Tibetan Plateau.
2.Seed size was strong related to phylogeny. Class, order and family membership accounted for most of the variation in log seed size(11.6%,31.4%and 58.2%, respectively).Therefore, the seed size variation was determined mainly by the phylogeny.
3.From whaterever taxonomic levels, seed size variation showed no consistent direction of evolution or evolutionary model.Generally, the seeds near the base of APGⅡsystem are likely to be in the middle size. Trends in the evolution of seed size has gone in many directions,towards larger or smaller seeds.Thus, the seed size of angiosperm has the obvious characteristics of adaptive radiation.
4.With different areal-types of plants, the seed size variation is significant difference. The primitive families and genera have the larger seeds, and the more evolved families and genera have the smaller seeds relatively in general.
5.Seed size was correlated with many other plant attributes such as dispersal mode, growth form, plant height, perenniality, fruit type, onset of flowering and duration of flowering, respectively,even in after phylogenic effect. The strongest of the plant attributes in seed size are fruit type, and dispersal mode, followed by plant height and growth form.
6. Seed size variation is also significantly associated with vegetation types.The mean log scale seed size from large to small in different vegetation types is forests, shrubs, cultivated vegetation, steppes and medows, aquatic vegetation, open vegetation on flowing rock hillside, cushion vegetation and swamp vegetation.
7. There is significantly negative relationship between the variation in seed size and altitude in the flora, and seed size decreases with altitude on the whole.But one interesting phenomenon is that the seeds with middle size are distrbuted at 3100m, and seed size becomes to larger or smaller respectively along with the altitude increasing or decreasing respectively. That is to say, the seed size has the evolutionary pattern of adaptive radiation.
8.The effect of altitude on seed size variation has different patterns among dispersal modes, areal-types of plants, growth forms, fruit types, different plant height, different perennialities, vegetation types, families, and even among species in a family. But there is a commen feature:from every aspect to examine effect of altitude on seed size variation, it seems that the seed size has a vertical distribution implications of adaptive radiation. That is, taking the altitude of 3100m as a starting point, the seed size becomes to larger or smaller respectively along with the altitude increasing or decreasing respectively.
引文
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