黄土高原土壤种子库与植被恢复研究
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摘要
以黄土高原典型草原区、森林草原区和荒漠草原区为研究区域，采用资料收集、野外调查和室内分析相结合的方法，研究黄土高原不同地带性植被土壤种子库的特征，分析土壤种子库在黄土高原植被构建中的作用机理以及在退化植被恢复过程中的作用与潜能，以期为黄土高原退化生态系统的恢复与重建提供科学依据和理论支持。主要结论如下：
1.阐明了黄土高原地区土壤种子库的物种组成、多样性、大小和空间分布格局等特征。在不同地带性植被土壤种子库中，均为草本植物占优势，以禾本科和菊科植物为主；黄土高原拥有丰富的种子库资源，种子库密度在典型草原为1388~9853seeds·m-2，森林草原为567~1210seeds·m-2，荒漠草原为31~482seeds·m-2；土壤种子库物种丰富度和多样性在三个地带性植被中排序为：典型草原>森林草原>荒漠草原；黄土高原地区有活力的种子主要分布在10cm土层以上。荒漠草原持久种子库中0~5cm和5~10cm的种子储量比例接近，其它地带性植被土壤种子库的储量均随土壤深度的增加呈下降趋势，具有明显的垂直分布格局（枯枝落叶层除外）。
2.揭示了黄土高原地区土壤种子库与地上植被的关系，以及土壤种子库在黄土高原植被结构建成和退化植被恢复中的作用与潜力。黄土高原地区土壤种子库与地上植被的相似性较低，Jaccard相似性指数在典型草原为0.29~0.38，森林草原为0.15~0.21，荒漠草原为0.05~0.27。在典型草原封禁演替过程中，土壤种子库在演替前期对植被的贡献较演替后期大，但从整体看土壤种子库在封禁演替过程中发挥的作用较小，而且土壤种子库也不能准确地预测未来植被的发展方向和演替趋势；森林草原区乔木实生苗成苗率低，依靠实生苗进行森林种群更新演替困难。土壤种子库是森林维持物种多样性的一个重要来源，但不是森林更新的主要方式，优势树种种群的天然更新主要依靠萌蘖苗。荒漠草原依靠持久种子库恢复灌木层的植被潜能很小。可见，黄土高原地区种子库对地上植被的贡献较低，仅仅依靠土壤种子库实现退化植被的自然修复的可能性非常小，地上植被的自然更新很有可能依赖于种子扩散和无性繁殖。
3.研究了干扰措施（封禁和放牧）对典型草原土壤种子库的影响。封禁和放牧措施未能显著改变土壤种子库优势物种组成的变化，但放牧使土壤种子库中一些伴生种消失，而封禁保护了这些物种在土壤种子库中存留；短期封禁显著提高土壤种子库的物种多样性，长期封禁则降低了物种多样性；封禁和放牧措施对土壤种子库密度影响显著，长期封禁使土壤种子库密度显著增加。封禁和放牧对土壤种子库密度的影响主要集中在枯枝落叶层和0~5cm土层，封禁主要提高了种子库中5cm以上的种子数量，对5cm以下的种子数量无显著改变；封禁和放牧措施通过改变草地枯落物生物量和厚度影响枯枝落叶层的种子数量，这正是封禁和放牧措施影响种子库数量的内部驱动因素之一；放牧地土壤种子库密度的变异系数最大，短期封禁使土壤种子库密度的变异系数略微降低，而长期封禁则使土壤种子库密度的变异系数明显降低；封禁和放牧措施对地上植被与土壤种子库之间的相似性影响不显著。
4.研究了典型草原地上植被与土壤种子库在封禁演替过程中的特征变化。在典型草原区，本氏针茅群落随着封禁年限的延长，其地上植被表现出明显的演替趋势，但是这种演替趋势未能在土壤种子库中得以体现，说明土壤种子库对外界干扰的反应滞后于地上植被。随着封禁演替年龄的增加，土壤种子库密度呈递增趋势；土壤种子库物种组成的相似性递减，说明土壤种子库物种更替速率逐渐增加；地上植被与土壤种子库之间的相似性先升高后降低，说明土壤种子库对地上植被的贡献在演替前期大于演替后期。
5.研究了森林草原优势树种种子质量与命运。动物捕食、虫蛀是土壤种子库优势树种种子损失的主要原因，占种子储量的30%；霉变是引起土壤种子库中优势树种种子消耗的另一个重要因素，占到15%；仍有55%的种子为优势树种种群更新提供种源，但只有0.1%的机会幼苗能成功建植。
6.分析了地理坐标和海拔对荒漠草原持久土壤种子库特征的影响。在荒漠草原区，持久土壤种子库密度与纬度之间呈显著负相关，而与经度的相关性不显著；经纬度对持久种子库物种丰富度、多样性，以及地上植被-土壤种子库的相似性影响不显著；海拔对持久种子库密度影响不显著，但对持久种子库的物种丰富度、多样性，以及地上植被-土壤种子库的相似性影响显著，随着海拔的升高，物种丰富度、多样性，以及地上植被-土壤种子库的相似性降低。
7.分析了黄土高原地区主要物种种子在土壤中的持久性及种子质量、形状与持久性之间的关系。黄土高原地区145种植物中，具有持久种子库的物种占48%，具有瞬时种子库的物种占44%。种子质量、形状均与持久性呈显著负相关，说明种子质量与形状是决定植物种子在土壤中的命运及持久性的重要特征，是预测种子在土壤中的持久性的重要指标。易于埋藏是这种关系存在的最主要的内在机制。
The research aimed to explore the potential role of soil seed bank in the process of plantcommunity construction, the vegetation restoration and reconstruction in the degradedecosystems. Moreover, this study provided a theoretical basis and scientific foundation forrestoration and reconstruction of degraded ecosystems. Study was carried out in three zonalvegetations on the Loess Plateau including typical grasslands, forest grasslands and desertgrasslands. The methods of a combined operation of historical data collection with fieldinvestigation and laboratory analysis were used to systematically study the characteristics ofsoil seed bank, and its relationship with vegetation in three zonal vegetations. The mainresults were concluded as follows:
1. The basic characteristics of soil seed bank on the Loess Plateau were clarified, suchas species composition, species diversity, density and spatial distribution pattern. Soil seedbank was dominated by Poaceae, Compositae species, and herbaceous plants. Soil seed bankdensity ranged as1388~9853seeds·m-2in typical grasslands,567~1210seeds·m-2in forestgrasslands and31~482seeds·m-2in desert grasslands, indicated that restoration of degradedecosystems on the Loess Plateau was not seed limited. Species richness and diversity of thesoil seed bank had the same position in three grassland types, and ranked as follows: typicalgrasslands> forest grasslands> desert grasslands. Most viable seeds were scattered at the≤10cm depth of the soil. Soil seed bank had obvious vertical distribution pattern that seeddensity significantly reduced with the depth increasing (except seed density in the litter layer),except that seed density at the0~5cm depth and at the5~10cm depth were similar in desertgrasslands.
2. The relationship between soil seed bank and vegetation, and the potential contributionof soil seed bank to the process of plant community construction, and the vegetationrestoration and reconstruction in the degraded ecosystems were revealed. The similarity indexbetween soil seed bank and vegetation was very low on the Loess Plateau, Jaccard similarityindex ranged as0.29~0.38in typical grasslands,0.15~0.21in forest grasslands,0.05~0.27indesert grasslands. In typical grasslands, soil seed bank played a minimal role in the process ofvegetation succession under the grazing exclusion, and could not accurately predict thedirection of vegetation succession and development. In forest grasslands, although soil seed bank played an important role in maintaining the species diversity, it was not an importantmeans for forest natural regeneration. The seedlings had extremely low survival rate and weregreatly sparse, the natural population regeneration depended mainly on sprouting instead ofsoil seed bank. In desert grasslands, it was likely that the potential of shrub layers restorationdepending on soil seed bank was very low. Overall, the establishment of the vegetation inthese communities is more likely to rely on seed dispersal and asexual recruitment, and thepotential role of degraded ecosystems restoration depending on soil seed bank was very lowon the Loess Plateau.
3. Impacts of disturbance (grazing and grazing exclusion) on soil seed bankcharacteristics in typical grasslands were studied. The results showed that grazing and grazingexclusion measures did not significantly change the dominant species composition of the soilseed bank, however, grazing measures made the companion species disappear from the soilseed bank and grazing exclusion protected these species to remain in the soil. Moderategrazing significantly increased the species numbers of soil seed bank, but significantreduction in long-term grazing exclusion grasslands. Species diversity of soil seed bank wassignificantly improved by short-term grazing exclusion, but was significantly reduced bylong-term grazing exclusion. Grazing and grazing exclusion significantly affected the densityof soil seed bank by changing the density at0~5cm depth, short-term grazing exclusion didnot significantly increase the density of soil seed bank, but significant increase was found inlong-term grazing exclusion grasslands. Litter biomass and thickness significantly affected theseed density, not the species number, which was the internal driver for the effects of grazingand grazing exclusion on soil seed bank. Coefficient of variation (CV) of seed bank densityfor grazing grasslands was largest, and slightly was reduced in short-term grazing exclusiongrasslands, and significantly reduced in long-term grazing exclusion grasslands. Grazing andgrazing exclusion did not significantly affect the similarity between soil seed bank andvegetation.
4. The effects of succession on vegetation and soil seed bank were analyzed. During thecourse of succession under grazing exclusion measures, the vegetation showed a clearsuccession trend, however, this trend is not reflected in soil seed bank. The response of soilseed bank to disturbance is latter than vegetation. Soil seed density increased，and thesimilarity of species composition decreased with succession age increasing. The soil seedbank played an important role in the vegetation restoration in the early succession stage, andseed bank′role became weaker and weaker with succession.
5. Seed quality and fate after dispersal of dominant trees were analyzed. Seed lossoccurred mainly because of vermiculating, predation and removal by animals (30%), and mildewing (15%), among which predation and removal by animals was the most importantfactor that caused seed bank depletion. Germinating or sound seeds accounted for55%,which provided enough seed source for population natural regeneration, however, theseedlings had only0.1%survival rate.
6. Impacts of the geographic location (Longitude, Latitude, altitude) on thecharacteristics of the persistent seed bank in desert grasslands were revealed. In desertgrasslands, persistent seed bank density was significantly and negatively correlated withlatitude, but no significant correlation with longitude. Longitude and Latitude did notsignificantly affect the species richness and diversity of the persistent soil seed bank, and thesimilarity between the persistent soil seed bank and vegetation. Altitude did not significantlyaffect the density of the persistent seed bank, however, it significantly affected the speciesrichness and diversity of the persistent soil seed bank, and the similarity between thepersistent soil seed bank and vegetation. Species richness and diversity of the persistent seedbank, and its similarity with vegetation decreased with the altitude increasing.
7. Seed persistence in the soil for145species belonging to Loess Plateau were tested,and whether and how seed mass and shape are related to seed persistence in soil were studied.Among145species with a range of growth forms, from a wide range of families, in the LoessPlateau, the species with persistent seed bank accounted for48%, and the species withtransient seed bank accounted for44%. Seed mass and shape were found to be significantlyand negatively correlated to persistence in the soil. The value of seed mass and shape as goodpredictors of persistence in the soil could be applied to the Loess Plateau. Although manyfactors affect the relationship between seed mass, shape and persistence, ease of burialappears to be the main potential interpretation.
1.阐明了黄土高原地区土壤种子库的物种组成、多样性、大小和空间分布格局等特征。在不同地带性植被土壤种子库中，均为草本植物占优势，以禾本科和菊科植物为主；黄土高原拥有丰富的种子库资源，种子库密度在典型草原为1388~9853seeds·m-2，森林草原为567~1210seeds·m-2，荒漠草原为31~482seeds·m-2；土壤种子库物种丰富度和多样性在三个地带性植被中排序为：典型草原>森林草原>荒漠草原；黄土高原地区有活力的种子主要分布在10cm土层以上。荒漠草原持久种子库中0~5cm和5~10cm的种子储量比例接近，其它地带性植被土壤种子库的储量均随土壤深度的增加呈下降趋势，具有明显的垂直分布格局（枯枝落叶层除外）。
2.揭示了黄土高原地区土壤种子库与地上植被的关系，以及土壤种子库在黄土高原植被结构建成和退化植被恢复中的作用与潜力。黄土高原地区土壤种子库与地上植被的相似性较低，Jaccard相似性指数在典型草原为0.29~0.38，森林草原为0.15~0.21，荒漠草原为0.05~0.27。在典型草原封禁演替过程中，土壤种子库在演替前期对植被的贡献较演替后期大，但从整体看土壤种子库在封禁演替过程中发挥的作用较小，而且土壤种子库也不能准确地预测未来植被的发展方向和演替趋势；森林草原区乔木实生苗成苗率低，依靠实生苗进行森林种群更新演替困难。土壤种子库是森林维持物种多样性的一个重要来源，但不是森林更新的主要方式，优势树种种群的天然更新主要依靠萌蘖苗。荒漠草原依靠持久种子库恢复灌木层的植被潜能很小。可见，黄土高原地区种子库对地上植被的贡献较低，仅仅依靠土壤种子库实现退化植被的自然修复的可能性非常小，地上植被的自然更新很有可能依赖于种子扩散和无性繁殖。
3.研究了干扰措施（封禁和放牧）对典型草原土壤种子库的影响。封禁和放牧措施未能显著改变土壤种子库优势物种组成的变化，但放牧使土壤种子库中一些伴生种消失，而封禁保护了这些物种在土壤种子库中存留；短期封禁显著提高土壤种子库的物种多样性，长期封禁则降低了物种多样性；封禁和放牧措施对土壤种子库密度影响显著，长期封禁使土壤种子库密度显著增加。封禁和放牧对土壤种子库密度的影响主要集中在枯枝落叶层和0~5cm土层，封禁主要提高了种子库中5cm以上的种子数量，对5cm以下的种子数量无显著改变；封禁和放牧措施通过改变草地枯落物生物量和厚度影响枯枝落叶层的种子数量，这正是封禁和放牧措施影响种子库数量的内部驱动因素之一；放牧地土壤种子库密度的变异系数最大，短期封禁使土壤种子库密度的变异系数略微降低，而长期封禁则使土壤种子库密度的变异系数明显降低；封禁和放牧措施对地上植被与土壤种子库之间的相似性影响不显著。
4.研究了典型草原地上植被与土壤种子库在封禁演替过程中的特征变化。在典型草原区，本氏针茅群落随着封禁年限的延长，其地上植被表现出明显的演替趋势，但是这种演替趋势未能在土壤种子库中得以体现，说明土壤种子库对外界干扰的反应滞后于地上植被。随着封禁演替年龄的增加，土壤种子库密度呈递增趋势；土壤种子库物种组成的相似性递减，说明土壤种子库物种更替速率逐渐增加；地上植被与土壤种子库之间的相似性先升高后降低，说明土壤种子库对地上植被的贡献在演替前期大于演替后期。
5.研究了森林草原优势树种种子质量与命运。动物捕食、虫蛀是土壤种子库优势树种种子损失的主要原因，占种子储量的30%；霉变是引起土壤种子库中优势树种种子消耗的另一个重要因素，占到15%；仍有55%的种子为优势树种种群更新提供种源，但只有0.1%的机会幼苗能成功建植。
6.分析了地理坐标和海拔对荒漠草原持久土壤种子库特征的影响。在荒漠草原区，持久土壤种子库密度与纬度之间呈显著负相关，而与经度的相关性不显著；经纬度对持久种子库物种丰富度、多样性，以及地上植被-土壤种子库的相似性影响不显著；海拔对持久种子库密度影响不显著，但对持久种子库的物种丰富度、多样性，以及地上植被-土壤种子库的相似性影响显著，随着海拔的升高，物种丰富度、多样性，以及地上植被-土壤种子库的相似性降低。
7.分析了黄土高原地区主要物种种子在土壤中的持久性及种子质量、形状与持久性之间的关系。黄土高原地区145种植物中，具有持久种子库的物种占48%，具有瞬时种子库的物种占44%。种子质量、形状均与持久性呈显著负相关，说明种子质量与形状是决定植物种子在土壤中的命运及持久性的重要特征，是预测种子在土壤中的持久性的重要指标。易于埋藏是这种关系存在的最主要的内在机制。
The research aimed to explore the potential role of soil seed bank in the process of plantcommunity construction, the vegetation restoration and reconstruction in the degradedecosystems. Moreover, this study provided a theoretical basis and scientific foundation forrestoration and reconstruction of degraded ecosystems. Study was carried out in three zonalvegetations on the Loess Plateau including typical grasslands, forest grasslands and desertgrasslands. The methods of a combined operation of historical data collection with fieldinvestigation and laboratory analysis were used to systematically study the characteristics ofsoil seed bank, and its relationship with vegetation in three zonal vegetations. The mainresults were concluded as follows:
1. The basic characteristics of soil seed bank on the Loess Plateau were clarified, suchas species composition, species diversity, density and spatial distribution pattern. Soil seedbank was dominated by Poaceae, Compositae species, and herbaceous plants. Soil seed bankdensity ranged as1388~9853seeds·m-2in typical grasslands,567~1210seeds·m-2in forestgrasslands and31~482seeds·m-2in desert grasslands, indicated that restoration of degradedecosystems on the Loess Plateau was not seed limited. Species richness and diversity of thesoil seed bank had the same position in three grassland types, and ranked as follows: typicalgrasslands> forest grasslands> desert grasslands. Most viable seeds were scattered at the≤10cm depth of the soil. Soil seed bank had obvious vertical distribution pattern that seeddensity significantly reduced with the depth increasing (except seed density in the litter layer),except that seed density at the0~5cm depth and at the5~10cm depth were similar in desertgrasslands.
2. The relationship between soil seed bank and vegetation, and the potential contributionof soil seed bank to the process of plant community construction, and the vegetationrestoration and reconstruction in the degraded ecosystems were revealed. The similarity indexbetween soil seed bank and vegetation was very low on the Loess Plateau, Jaccard similarityindex ranged as0.29~0.38in typical grasslands,0.15~0.21in forest grasslands,0.05~0.27indesert grasslands. In typical grasslands, soil seed bank played a minimal role in the process ofvegetation succession under the grazing exclusion, and could not accurately predict thedirection of vegetation succession and development. In forest grasslands, although soil seed bank played an important role in maintaining the species diversity, it was not an importantmeans for forest natural regeneration. The seedlings had extremely low survival rate and weregreatly sparse, the natural population regeneration depended mainly on sprouting instead ofsoil seed bank. In desert grasslands, it was likely that the potential of shrub layers restorationdepending on soil seed bank was very low. Overall, the establishment of the vegetation inthese communities is more likely to rely on seed dispersal and asexual recruitment, and thepotential role of degraded ecosystems restoration depending on soil seed bank was very lowon the Loess Plateau.
3. Impacts of disturbance (grazing and grazing exclusion) on soil seed bankcharacteristics in typical grasslands were studied. The results showed that grazing and grazingexclusion measures did not significantly change the dominant species composition of the soilseed bank, however, grazing measures made the companion species disappear from the soilseed bank and grazing exclusion protected these species to remain in the soil. Moderategrazing significantly increased the species numbers of soil seed bank, but significantreduction in long-term grazing exclusion grasslands. Species diversity of soil seed bank wassignificantly improved by short-term grazing exclusion, but was significantly reduced bylong-term grazing exclusion. Grazing and grazing exclusion significantly affected the densityof soil seed bank by changing the density at0~5cm depth, short-term grazing exclusion didnot significantly increase the density of soil seed bank, but significant increase was found inlong-term grazing exclusion grasslands. Litter biomass and thickness significantly affected theseed density, not the species number, which was the internal driver for the effects of grazingand grazing exclusion on soil seed bank. Coefficient of variation (CV) of seed bank densityfor grazing grasslands was largest, and slightly was reduced in short-term grazing exclusiongrasslands, and significantly reduced in long-term grazing exclusion grasslands. Grazing andgrazing exclusion did not significantly affect the similarity between soil seed bank andvegetation.
4. The effects of succession on vegetation and soil seed bank were analyzed. During thecourse of succession under grazing exclusion measures, the vegetation showed a clearsuccession trend, however, this trend is not reflected in soil seed bank. The response of soilseed bank to disturbance is latter than vegetation. Soil seed density increased，and thesimilarity of species composition decreased with succession age increasing. The soil seedbank played an important role in the vegetation restoration in the early succession stage, andseed bank′role became weaker and weaker with succession.
5. Seed quality and fate after dispersal of dominant trees were analyzed. Seed lossoccurred mainly because of vermiculating, predation and removal by animals (30%), and mildewing (15%), among which predation and removal by animals was the most importantfactor that caused seed bank depletion. Germinating or sound seeds accounted for55%,which provided enough seed source for population natural regeneration, however, theseedlings had only0.1%survival rate.
6. Impacts of the geographic location (Longitude, Latitude, altitude) on thecharacteristics of the persistent seed bank in desert grasslands were revealed. In desertgrasslands, persistent seed bank density was significantly and negatively correlated withlatitude, but no significant correlation with longitude. Longitude and Latitude did notsignificantly affect the species richness and diversity of the persistent soil seed bank, and thesimilarity between the persistent soil seed bank and vegetation. Altitude did not significantlyaffect the density of the persistent seed bank, however, it significantly affected the speciesrichness and diversity of the persistent soil seed bank, and the similarity between thepersistent soil seed bank and vegetation. Species richness and diversity of the persistent seedbank, and its similarity with vegetation decreased with the altitude increasing.
7. Seed persistence in the soil for145species belonging to Loess Plateau were tested,and whether and how seed mass and shape are related to seed persistence in soil were studied.Among145species with a range of growth forms, from a wide range of families, in the LoessPlateau, the species with persistent seed bank accounted for48%, and the species withtransient seed bank accounted for44%. Seed mass and shape were found to be significantlyand negatively correlated to persistence in the soil. The value of seed mass and shape as goodpredictors of persistence in the soil could be applied to the Loess Plateau. Although manyfactors affect the relationship between seed mass, shape and persistence, ease of burialappears to be the main potential interpretation.
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