三峡库区消落带土壤理化性质及种子库研究
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摘要
三峡库区消落带水位周期性涨落,产生了干湿交替的生态环境,其土壤理化性质及种子库物种势必会发生较大变化。本文针对三峡库区消落带水淹初期的特殊情况,连续两年对土壤理化性质和种子库的动态研究,探讨其对水位变化的响应趋势,为未来三峡库区消落带的植被恢复和重建提供理论依据。
在地理条件相似的条件下,选取消落带内现存的次生灌丛和弃耕地固定样地4块,采用海拔梯度代替水淹时间,以海拔145 m、155 m、165 m、175 m和185 m为分界线,将消落带划分为145-155 m,155-165 m、165-175 m三段高程,将存有原生植被的175-185 m作为对照样带,对地上植被、土壤表层水分物理性质、土壤化学元素、土壤种子库季节性变化、年际变化等进行系统研究,探讨水淹初期消落带土壤理化性质与种子库如何变化,以及种子库优势物种与环境因子的关系,结果表明:
1.不同月份表层土壤含水率,除了最大持水量、毛管持水量、最小持水量、非毛管孔隙度、土壤通气度9月份最高外,土壤质量含水量、土壤体积含水量、毛管空隙度、总孔隙度5月份最高,7月份最低;土壤容重在5月份和7月份相差不大,而9月份则最小。随着海拔梯度的升高,各月份土壤表层土壤含水率未表现出一致的变化趋势,但以水淹时间最长的145-155m区段土壤物理性质最差。
2.连续水淹两年后,土壤化学元素含量降低的分别是土壤pH值1.18%、全氮20.16%、全磷30.80%、全钾6.87%、速效磷103.75%、速效钾16.90%、全铜12.12%、全锰16.17%、全锌50.16%,而含量升高的分别是有机质7.22%、水解性氮3.03%、全铁6.88%、全钙31.89%、全镁84.80%、全钠98.44%;随着海拔梯度的升高,水淹和人为因素干扰导致各种元素变化方式差异较大;随着土壤层次的加深,除速效磷、全铜、全铁、全锰、全锌、全钙、全镁、全钠未表现出规律性变化趋势外,其他元素在2008年表现为逐渐降低,2009年则表现为逐渐增加。
3.水淹初期两种植被类型的土壤种子库储量存在极显著差异,次生灌丛种子密度为6991±954.33粒/m2、弃耕地种子密度为22582±6217.36粒/m2。已淹区段种子密度最低,未淹区段最高。随着土壤层次的加深,种子库密度逐渐下降。土壤种子库萌发出现的物种分属45科97属118种,以一年生和多年生草本植物为主,菊科、禾本科、玄参科和十字花科为优势科;个体数量占土壤种子库总储量<0.01%的有34种,占所有物种的28.82%。两种植被类型内土壤种子库出现的物种数较接近,物种多样性指数、均匀度都较高,但优势物种组成差异很大,生态优势度较低。未淹区段的生物多样性最高,而已淹区段的生态优势度最高,已淹区段和未淹区段相似性指数最大。
4.已淹区段地表植被分属47科71属100种,未淹区段地表植被分属84科163属223种,优势科主要包括菊科、豆科、禾本科、蔷薇科、莎草科。与对照样带比较,已淹区段地表植被物种多样性指数和丰富度指数都降低,未淹区段明显增加;已淹区段、未淹区段土壤种子库具有物种79种和108种,分别比对照样带增加了9种和38种;与对照样带土壤种子库总密度相比,已淹区段降低了32.32%,未淹区段增加了103.12%。土壤种子库植物以草本占绝大多数比例,且1年生草本所占比例大于多年生草本所占比例。土壤种子库的生物多样性指数、丰富度指数和均匀度指数与地表植被变化相一致。已淹区段、未淹区段、对照样带地表植被和土壤种子库共同出现的物种数分别为9,40和15种,土壤种子库和地表植被间的相似性系数分别为0.328,0.241,0.186。
5.随着月份的延伸,次生灌丛土壤种子库密度逐渐增加,弃耕地逐渐降低。随着海拔梯度的升高,5、7、9月份土壤种子库密度都表现为逐渐增加。5月份土壤种子库共出现了79种物种,隶属于36科66属;7月份土壤种子库共出现了80种物种,隶属于35科69属;9月土壤种子库共出现了73种物种,隶属于32科64属,各月共同出现的主要优势科为菊科、禾本科、玄参科、十字花科。土壤种子库物种生物多样性指数逐渐降低,但相差范围不是很大。各月份消落带土壤种子库的物种生活型非常接近,主要以一年生植物、多年生植物种子为主,而相对缺少乔木种子。
6.随着水淹年限的延伸,次生灌丛和弃耕地土壤种子库密度呈现降低的趋势,且弃耕地降低的比例较次生灌丛高。2009年土壤种子库共出现了85种,隶属于35科70属,较前一年共减少了10科27属33种。次生灌丛地上植被减少的植物种类为105种37科78属,弃耕地减少56种17科44属,优势乔灌隶属于大戟科、豆科、蔷薇科等,优势草本属于禾本科、菊科、莎草科、玄参科等。地上植被与土壤种子库物种类型的相似性显著提高。
7.应用典范对应分析(CCA)研究土壤表层水分物理性质、化学性质与各月份土壤种子库优势物种分布的关系,海拔梯度、土壤容重、通气度对种子库内物种储存分布具有较一定的影响;pH值、全磷、速效磷、全钠、全钙、全铁、全锰等化学元素在解释不同月份种子库优势物种的分布影响较强。
The periodical fluctuation of water-level-fluctuating zone in Three Gorges Reservoir caused the variation of the wet and dry state of the soil ecological environment, which would inevitably lead to the difference of soil physical and chemical properties and seed bank species. In this paper, we studied soil physical and chemical properties and seed bank species changes in two consecutive years to explore the response tendency of seed bank species to the water level changes. It can provide theoretical basis for vegetation rehabilitation and reconstruction of water-level-fluctuating zone in Three Gorges Reservoir Area.
According to the similar geographical conditions, we chose 4 sample lands in secondary bush and abandoned farmland which are the existing vegetation types in water-level-fluctuating zone as our research objects. We divided the sample lands into 145-155m, 155-165m, 165-175m three sections, and kept 175-185m as the control area to keep native vegetation according to the criteria of replacing elevation gratitude by the flooded time, and making 145m, 155m, 165m, 175m, 185m as the boundary. We studied the standing vegetation, surface soil moisture physical properties, soil chemical properties, seasonal changes of soil seed bank and inter-annual changes in systematic way to investigate the changes of the soil physical and chemical properties, seed bank species in the initial stage of river-flooding, and the relationship between dominant specie of soil seed bank and environmental factors from dry to flooded states. The results showed that:
1. In addition to maximum water holding capacity, capillary water holding capacity, minimum water holding capacity, non-capillary porosity, degree of soil aeration , surface soil moisture content in different months, were highest in September. Soil quality water content, soil volumetric water content, capillary porosity and total porosity were highest in May and lowest in July. Soil bulk density in May and in July has no obvious difference, but was lowest in September. With the elevation gradient increases, the soil moisture content did not display the same variation trend, but soil moisture physical properties in the 145-155m section was the lowest due to its longest flooded time.
2. Comparison of soil chemical properties in flooded lands for two years, the nutrient element including soil pH, total nitrogen, total phosphorus, total potassium, available phosphorous, available potassium, total Cu, total Mn, total Zn content decreased. The value was 1.18%, 20.16%, 30.80%, 6.87%, 103.75%, 16.90%, 12.12%, 16.17%, 50.16%, separately. In contrast, organic matter, hydrolysis of nitrogen, total Fe, total Ca, total Mg, total Na increased, the value were separately 7.22%, 3.03%, 6.88%, 31.89%, 84.80%, 98.44%; With the increase of elevation gradient, various elements was very different caused by water flooded and human disturbance; With the increase of soil layers, except total phosphorus, total Cu, total Fe, total Mn, total Zn, total Ca, total Mg, total Na did not display the regularity of changes in trend, other elements decreased gradually in 2008, while increased in 2009.
3. We divided water-level-fluctuating zone into flooded section, non-flooded section and control section, and studied the characteristics of soil seed banks in two main vegetation types by germination method in 2008. The results showed that average seed density in secondary bush was 6991±954.33 seed per m2 while in abandoned farmland 22582±6217.36 seed per m2 and there was significant difference between them. Seed density in non-flooded area was more than the control area which is close to water-level-fluctuating zone, and seed density in flooded area was the minimum. Soil seed density decreased with soil depth. A total of 118 species which belonged to 45 family and 97 genera were detected in the soil of the two vegetation types. Most of them were annual and perennial herbage species belonging to the main family Asteraceae, Poaceae, Scrophulariaceae, and Cruciferae. Moreover, the number of the species less than 0.01%, was 34. This accounted for 28.8% in all species. Species number in the soil seed of the two vegetation types was similar. The species diversity index and evenness index were relative high. However, the significant difference between dominant species of the two vegetation types and ecological dominance was relatively low. In addition, species diversity in the non-flooded section and ecological dominance in the flooded section were the highest, and the similarity index between the flooded and non-flooded section was the highest.
4. The standing vegetation in flooded section and non-flooded section was composed of 100 species in 71 genera of 47 families, and 223 species 163 genera of 84 families. The main family consists of Asteraceae, Leguminosae, Poaceae, Rosaceae, Cyperaceae. In the standing vegetation in flooded section, species diversity indices and richness indices compared to control section decreased significant. And it increased significantly in non-flooded section. Evenness indices were all decreased in these areas. The species number of the soil seed bank in flooded section and non-flooded section were 79 and 108, with 9 and 38 species more than control section; Compared to the seed bank density of control section, seed bank density decreased by 32.32% in flooded section and increased by 103.12%, and herbaceous species seeds accounted for majority proportion in these two sites. The proportion of annual herbaceous species seeds was higher than of that perennial herbaceous. The trends of indices biodiversity, richness, evenness of the soil seed bank were consistent with the standing vegetation. However, the extent of the change was not obvious. In flooded section, non-flooded section and control section, the co-occurrence species in standing vegetation and the soil seed bank were 9, 40, and 15. The similarity coefficients of species composition between the soil seed bank and the standing vegetation is 0.328, 0.241 and 0.186.
5. The soil seed bank density of two vegetation types in hydro-fluctuation belt displayed different changes trend along with extension of the month, The secondary bush increased gradually, the value were separately 7534.2 seed per m2, 13710 seed per m2, 16746 seed per m2. The abandoned farmland decreased, the value were separately 19423 seed per m2, 14977 seed per m2, 11869 seed per m2. With the increase of elevation gradient, the density of soil seed bank in May, July, September have shown gradual upward trend. In May, July, September, the soil seed bank of water-level-fluctuating zone of Three Gorges Reservoir was composed of 79 species in 66 genera of 36 families, and 80 species 69 genera of 35 families, 73 species 64 genera of 32 families. The main family of co-occurrence of each month were Compositae, Gramineae, Scrophulariaceae, Brassicaceae. Soil seed bank of species biodiversity index showed decreased trend as an extension of the month, however, the range of difference was not very great. Hydro-fluctuation belt of the soil seed bank of species life-form in different month is very similar. They are mainly annuals, perennials seed based, while being relatively lack of the seed of arbor.
6. As the extension for years of flooded, soil seed bank density of secondary bush and abandoned farmland of hydro-fluctuation belt showed a falling trend. The decreased proportion of the soil seed bank reserves of abandoned farmland was higher than that of secondary bush. Compare to last year, a total of 85 species which belonged to 35 family and 70 genera in 2009 reduced by 33 species 27 genera of 10 families. Compared with the change of two standing vegetation, reduction of vegetation type of secondary bush on the ground belonged to 105 species 78 genera of 37 families. While the abandoned farmland of 56 species belongs to 44 genera of 17 families. The main dominant family of arbor and shrub in the standing vegetation belongs to Euphorbiaceae, Leguminosae, Rosaceaein, and the main dominant herbage belongs to Gramineae, Asteraceae, Cyperaceae, Scrophulariaceae and so on. With the increase of flooded years, the similarity coefficients of species composition between the soil seed bank and the standing vegetation increased markedly.
7. We analyzed the relationship between physical properties, chemical properties of surface soil and the distribution of dominant species of soil seed bank in different months by CCA method. The distribution of seed bank species had closely relationship with elevation gradient, soil bulk density, degree of soil aeration. In addition, soil pH value, total phosphorus, available phosphorus, total sodium, total calcium, total iron and total manganese could interpret perfectly the distribution of dominant species of soil seed bank in different months.
在地理条件相似的条件下,选取消落带内现存的次生灌丛和弃耕地固定样地4块,采用海拔梯度代替水淹时间,以海拔145 m、155 m、165 m、175 m和185 m为分界线,将消落带划分为145-155 m,155-165 m、165-175 m三段高程,将存有原生植被的175-185 m作为对照样带,对地上植被、土壤表层水分物理性质、土壤化学元素、土壤种子库季节性变化、年际变化等进行系统研究,探讨水淹初期消落带土壤理化性质与种子库如何变化,以及种子库优势物种与环境因子的关系,结果表明:
1.不同月份表层土壤含水率,除了最大持水量、毛管持水量、最小持水量、非毛管孔隙度、土壤通气度9月份最高外,土壤质量含水量、土壤体积含水量、毛管空隙度、总孔隙度5月份最高,7月份最低;土壤容重在5月份和7月份相差不大,而9月份则最小。随着海拔梯度的升高,各月份土壤表层土壤含水率未表现出一致的变化趋势,但以水淹时间最长的145-155m区段土壤物理性质最差。
2.连续水淹两年后,土壤化学元素含量降低的分别是土壤pH值1.18%、全氮20.16%、全磷30.80%、全钾6.87%、速效磷103.75%、速效钾16.90%、全铜12.12%、全锰16.17%、全锌50.16%,而含量升高的分别是有机质7.22%、水解性氮3.03%、全铁6.88%、全钙31.89%、全镁84.80%、全钠98.44%;随着海拔梯度的升高,水淹和人为因素干扰导致各种元素变化方式差异较大;随着土壤层次的加深,除速效磷、全铜、全铁、全锰、全锌、全钙、全镁、全钠未表现出规律性变化趋势外,其他元素在2008年表现为逐渐降低,2009年则表现为逐渐增加。
3.水淹初期两种植被类型的土壤种子库储量存在极显著差异,次生灌丛种子密度为6991±954.33粒/m2、弃耕地种子密度为22582±6217.36粒/m2。已淹区段种子密度最低,未淹区段最高。随着土壤层次的加深,种子库密度逐渐下降。土壤种子库萌发出现的物种分属45科97属118种,以一年生和多年生草本植物为主,菊科、禾本科、玄参科和十字花科为优势科;个体数量占土壤种子库总储量<0.01%的有34种,占所有物种的28.82%。两种植被类型内土壤种子库出现的物种数较接近,物种多样性指数、均匀度都较高,但优势物种组成差异很大,生态优势度较低。未淹区段的生物多样性最高,而已淹区段的生态优势度最高,已淹区段和未淹区段相似性指数最大。
4.已淹区段地表植被分属47科71属100种,未淹区段地表植被分属84科163属223种,优势科主要包括菊科、豆科、禾本科、蔷薇科、莎草科。与对照样带比较,已淹区段地表植被物种多样性指数和丰富度指数都降低,未淹区段明显增加;已淹区段、未淹区段土壤种子库具有物种79种和108种,分别比对照样带增加了9种和38种;与对照样带土壤种子库总密度相比,已淹区段降低了32.32%,未淹区段增加了103.12%。土壤种子库植物以草本占绝大多数比例,且1年生草本所占比例大于多年生草本所占比例。土壤种子库的生物多样性指数、丰富度指数和均匀度指数与地表植被变化相一致。已淹区段、未淹区段、对照样带地表植被和土壤种子库共同出现的物种数分别为9,40和15种,土壤种子库和地表植被间的相似性系数分别为0.328,0.241,0.186。
5.随着月份的延伸,次生灌丛土壤种子库密度逐渐增加,弃耕地逐渐降低。随着海拔梯度的升高,5、7、9月份土壤种子库密度都表现为逐渐增加。5月份土壤种子库共出现了79种物种,隶属于36科66属;7月份土壤种子库共出现了80种物种,隶属于35科69属;9月土壤种子库共出现了73种物种,隶属于32科64属,各月共同出现的主要优势科为菊科、禾本科、玄参科、十字花科。土壤种子库物种生物多样性指数逐渐降低,但相差范围不是很大。各月份消落带土壤种子库的物种生活型非常接近,主要以一年生植物、多年生植物种子为主,而相对缺少乔木种子。
6.随着水淹年限的延伸,次生灌丛和弃耕地土壤种子库密度呈现降低的趋势,且弃耕地降低的比例较次生灌丛高。2009年土壤种子库共出现了85种,隶属于35科70属,较前一年共减少了10科27属33种。次生灌丛地上植被减少的植物种类为105种37科78属,弃耕地减少56种17科44属,优势乔灌隶属于大戟科、豆科、蔷薇科等,优势草本属于禾本科、菊科、莎草科、玄参科等。地上植被与土壤种子库物种类型的相似性显著提高。
7.应用典范对应分析(CCA)研究土壤表层水分物理性质、化学性质与各月份土壤种子库优势物种分布的关系,海拔梯度、土壤容重、通气度对种子库内物种储存分布具有较一定的影响;pH值、全磷、速效磷、全钠、全钙、全铁、全锰等化学元素在解释不同月份种子库优势物种的分布影响较强。
The periodical fluctuation of water-level-fluctuating zone in Three Gorges Reservoir caused the variation of the wet and dry state of the soil ecological environment, which would inevitably lead to the difference of soil physical and chemical properties and seed bank species. In this paper, we studied soil physical and chemical properties and seed bank species changes in two consecutive years to explore the response tendency of seed bank species to the water level changes. It can provide theoretical basis for vegetation rehabilitation and reconstruction of water-level-fluctuating zone in Three Gorges Reservoir Area.
According to the similar geographical conditions, we chose 4 sample lands in secondary bush and abandoned farmland which are the existing vegetation types in water-level-fluctuating zone as our research objects. We divided the sample lands into 145-155m, 155-165m, 165-175m three sections, and kept 175-185m as the control area to keep native vegetation according to the criteria of replacing elevation gratitude by the flooded time, and making 145m, 155m, 165m, 175m, 185m as the boundary. We studied the standing vegetation, surface soil moisture physical properties, soil chemical properties, seasonal changes of soil seed bank and inter-annual changes in systematic way to investigate the changes of the soil physical and chemical properties, seed bank species in the initial stage of river-flooding, and the relationship between dominant specie of soil seed bank and environmental factors from dry to flooded states. The results showed that:
1. In addition to maximum water holding capacity, capillary water holding capacity, minimum water holding capacity, non-capillary porosity, degree of soil aeration , surface soil moisture content in different months, were highest in September. Soil quality water content, soil volumetric water content, capillary porosity and total porosity were highest in May and lowest in July. Soil bulk density in May and in July has no obvious difference, but was lowest in September. With the elevation gradient increases, the soil moisture content did not display the same variation trend, but soil moisture physical properties in the 145-155m section was the lowest due to its longest flooded time.
2. Comparison of soil chemical properties in flooded lands for two years, the nutrient element including soil pH, total nitrogen, total phosphorus, total potassium, available phosphorous, available potassium, total Cu, total Mn, total Zn content decreased. The value was 1.18%, 20.16%, 30.80%, 6.87%, 103.75%, 16.90%, 12.12%, 16.17%, 50.16%, separately. In contrast, organic matter, hydrolysis of nitrogen, total Fe, total Ca, total Mg, total Na increased, the value were separately 7.22%, 3.03%, 6.88%, 31.89%, 84.80%, 98.44%; With the increase of elevation gradient, various elements was very different caused by water flooded and human disturbance; With the increase of soil layers, except total phosphorus, total Cu, total Fe, total Mn, total Zn, total Ca, total Mg, total Na did not display the regularity of changes in trend, other elements decreased gradually in 2008, while increased in 2009.
3. We divided water-level-fluctuating zone into flooded section, non-flooded section and control section, and studied the characteristics of soil seed banks in two main vegetation types by germination method in 2008. The results showed that average seed density in secondary bush was 6991±954.33 seed per m2 while in abandoned farmland 22582±6217.36 seed per m2 and there was significant difference between them. Seed density in non-flooded area was more than the control area which is close to water-level-fluctuating zone, and seed density in flooded area was the minimum. Soil seed density decreased with soil depth. A total of 118 species which belonged to 45 family and 97 genera were detected in the soil of the two vegetation types. Most of them were annual and perennial herbage species belonging to the main family Asteraceae, Poaceae, Scrophulariaceae, and Cruciferae. Moreover, the number of the species less than 0.01%, was 34. This accounted for 28.8% in all species. Species number in the soil seed of the two vegetation types was similar. The species diversity index and evenness index were relative high. However, the significant difference between dominant species of the two vegetation types and ecological dominance was relatively low. In addition, species diversity in the non-flooded section and ecological dominance in the flooded section were the highest, and the similarity index between the flooded and non-flooded section was the highest.
4. The standing vegetation in flooded section and non-flooded section was composed of 100 species in 71 genera of 47 families, and 223 species 163 genera of 84 families. The main family consists of Asteraceae, Leguminosae, Poaceae, Rosaceae, Cyperaceae. In the standing vegetation in flooded section, species diversity indices and richness indices compared to control section decreased significant. And it increased significantly in non-flooded section. Evenness indices were all decreased in these areas. The species number of the soil seed bank in flooded section and non-flooded section were 79 and 108, with 9 and 38 species more than control section; Compared to the seed bank density of control section, seed bank density decreased by 32.32% in flooded section and increased by 103.12%, and herbaceous species seeds accounted for majority proportion in these two sites. The proportion of annual herbaceous species seeds was higher than of that perennial herbaceous. The trends of indices biodiversity, richness, evenness of the soil seed bank were consistent with the standing vegetation. However, the extent of the change was not obvious. In flooded section, non-flooded section and control section, the co-occurrence species in standing vegetation and the soil seed bank were 9, 40, and 15. The similarity coefficients of species composition between the soil seed bank and the standing vegetation is 0.328, 0.241 and 0.186.
5. The soil seed bank density of two vegetation types in hydro-fluctuation belt displayed different changes trend along with extension of the month, The secondary bush increased gradually, the value were separately 7534.2 seed per m2, 13710 seed per m2, 16746 seed per m2. The abandoned farmland decreased, the value were separately 19423 seed per m2, 14977 seed per m2, 11869 seed per m2. With the increase of elevation gradient, the density of soil seed bank in May, July, September have shown gradual upward trend. In May, July, September, the soil seed bank of water-level-fluctuating zone of Three Gorges Reservoir was composed of 79 species in 66 genera of 36 families, and 80 species 69 genera of 35 families, 73 species 64 genera of 32 families. The main family of co-occurrence of each month were Compositae, Gramineae, Scrophulariaceae, Brassicaceae. Soil seed bank of species biodiversity index showed decreased trend as an extension of the month, however, the range of difference was not very great. Hydro-fluctuation belt of the soil seed bank of species life-form in different month is very similar. They are mainly annuals, perennials seed based, while being relatively lack of the seed of arbor.
6. As the extension for years of flooded, soil seed bank density of secondary bush and abandoned farmland of hydro-fluctuation belt showed a falling trend. The decreased proportion of the soil seed bank reserves of abandoned farmland was higher than that of secondary bush. Compare to last year, a total of 85 species which belonged to 35 family and 70 genera in 2009 reduced by 33 species 27 genera of 10 families. Compared with the change of two standing vegetation, reduction of vegetation type of secondary bush on the ground belonged to 105 species 78 genera of 37 families. While the abandoned farmland of 56 species belongs to 44 genera of 17 families. The main dominant family of arbor and shrub in the standing vegetation belongs to Euphorbiaceae, Leguminosae, Rosaceaein, and the main dominant herbage belongs to Gramineae, Asteraceae, Cyperaceae, Scrophulariaceae and so on. With the increase of flooded years, the similarity coefficients of species composition between the soil seed bank and the standing vegetation increased markedly.
7. We analyzed the relationship between physical properties, chemical properties of surface soil and the distribution of dominant species of soil seed bank in different months by CCA method. The distribution of seed bank species had closely relationship with elevation gradient, soil bulk density, degree of soil aeration. In addition, soil pH value, total phosphorus, available phosphorus, total sodium, total calcium, total iron and total manganese could interpret perfectly the distribution of dominant species of soil seed bank in different months.
引文
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