不同改良措施下退化羊草（Leymus chinensis）草原群落恢复演替规律研究
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摘要
放牧过度导致草原退化是近几十年来内蒙古草原的突出生态学问题。采取不同的经营管理方式和改良途径促进退化草原的恢复既是国家的需求,也是草原生态学研究的重要课题。
本研究在锡林郭勒盟,中国科学院内蒙古草原生态系统定位研究站样地完成。研究区位于北纬43°20′N～44°00′N,116°06′E～117°05′E。改良前植被为因过度放牧导致的羊草草原退化变体,即含小叶锦鸡儿(Caraganamicrophylla)灌丛的冷蒿(Artemisia frigida)+丛生小禾草+羊草群落。于1983年6月,用围栏封育、耙地、浅耕翻三个措施进行处理,围栏内样地没有进行任何方式的利用。采用植物样方法、土壤常规分析方法和土壤微生物分析方法对经过25年恢复演替草原样地内地植物群落特征、土壤理化性状和土壤微生物特征进行了分析,结果如下:
1.浅耕翻:群落地上生物量和密度组成中,禾本科占主导地位。最高时禾本科生物量占到总生物量的95%,总密度的90%以上,在整个恢复演替过程中一直保持在60%以上。以羊草为主的根茎禾草随着演替进程的延续一直处于优势地位。高丛生禾草在恢复演替的后期增加。群落总地上生物量与降水相关关系不显著,主要受改良措施造成的群落结构变化的影响。浅耕翻处理恢复过程中多样性指数、均匀性指数总变化趋势为开口向上的抛物线状。以生物量为指标的多样性指数(DI)与恢复年限(x)之间的关系可以用如下方程表示:DI=0.0004x~2+0.0113x+0.228(r=0.7777).浅耕翻处理恢复过程中群落主要植物的种群密度、生物量与多样性指数存在着显著的相关关系。浅耕翻改良处理,25年的恢复演替过程可划分为四个阶段:第一阶段,即处理初期(1983～1985),群落类型为羊草+一、二年生杂类草群落;第二阶段(1986～1991年)的群落类型为羊草群落;第三阶段(1992～1996)的群落类型为羊草+冰草+多年生杂类草的过渡型群落类型;第四阶段,即演替后期(1997～2007),群落类型为羊草+丛生禾草、多年生杂类草群落,此时羊草的重要值已基本上类似于原生群落的。
2.耙地处理:群落总地上生物量和密度由于耙地改良处理持续恢复,禾本科植物生物量作为生物量构成的主体,处于建群地位。根茎禾草作为耙地的受益者在恢复演替的前期呈现持续增加的趋势,中期呈现高位维持,而后有所下降。小半灌木和丛生禾草受耙地处理的抑制,多年生植物在演替后期增加,使群落更接近与天然草原群落。群落总地上生物量与降水的相关关系不显著,主要与改良措施造成的群落结构变化有关。耙地处理后,均匀性指数、多样性指数的变化随着恢复演替的进程前期呈现波动下降的趋势,恢复中后期有所升高。以生物量为指标的多样性指数(DI)与恢复年限(x)之间的关系可以用如下方程表示:DI=0.001x~2-0.042x+0.896(r=0.7169)耙地处理后羊草的密度和生物量的变化与多样性指数具有极显著的负相关关系。耙地改良处理25年的恢复演替过程可划分为四个阶段:第一阶段,即处理初期(1983～1986),群落类型为羊草+一、二年生杂类草+冷蒿、冰草群落;第二阶段(1987～1996年),群落类型为羊草+小丛生禾草+冰草、杂类草群落;第三阶段(1997～2000),群落类型为羊草+冰草+丛生禾草、多年生杂类草的过渡型群落类型;第四阶段(2001～2007),群落类型为羊草+大针茅+冰草、丛生禾草群落,此时羊草的重要值已基本上接近原生群落,而随着针茅等丛生禾草的增加,群落更趋近于天然群落。
3.围栏封育自然恢复:围栏封育自然恢复群落的地上生物量和密度就功能群组成来看,根茎禾草在恢复演替的前9年呈现快速增加、缓慢增加并维持的趋势,丛生禾草后期增加,在退化群落中占主导地位的以冷蒿为主的小半灌木持续减少并维持较低的比例。与前两种改良措施相比,围栏封育自然恢复处理下群落生物量的恢复较滞后。退化草原围栏封育自然恢复过程中植物群落的多样性指数和均匀性指数总体上基本稳定,变幅不大。退化草原围栏封育自然恢复过程中羊草、冰草、冷蒿的生物量和密度的变化与多样性指数具有显著的负相关关系,星毛委陵菜的生物量和密度的变化与多样性指数具有显著的正相关关系。围封自然恢复,25年的恢复演替过程可划分为四个阶段:第一阶段,为封育后1—3年(1983～1985),群落类型为冷蒿+丛生小禾草+羊草群落;第二阶段(1986～1988)为由冷蒿群落向羊草群落过渡的演替阶段;第三阶段(1989～1991)群落类型为羊草+冰草、冷蒿+丛生禾草群落;第四阶段(1992～2007)群落已演变成羊草+针茅+杂类草群落,已基本接近原生羊草草原群落。随着恢复年限的延续,针茅种群的群落重要性将继续增加。
4.环境差异变化:经过24年的恢复改良,土壤理化性质明显改善,但仍有差别。耕翻、耙地、自然恢复处理后的土壤容重分别比围栏外降低22.30%、21.62%、11.49%,且浅耕翻和耙地处理后的土壤容重显著低于自然恢复处理后的土壤容重;土壤含水量分别比围栏外土壤含水量提高93.71%、91.36%、86.39%,改良措施之间差异不显著;土壤L1层细菌的数量分别比围栏外增加了91.56%、89.66%、73.56%,改良措施之间并无显著差异;真菌的数量分别比围栏外提高了53.06%、50.34%、22.45%,自然恢复显著低于另外两种改良方法;放线菌的数量分别比围栏外提高了41.72%、38.89%、22.63%,浅耕翻、耙地处理后的显著高于自然恢复的。因此,浅耕翻和耙地更有利于土壤微生物数量的增加。
5.几种措施的长期效果评价
就与天然羊草草原的相似度来讲,对于恢复A.frigida草原这种广泛分布的退化草原,除去了食草动物的自然恢复要优于两种机械改良的方法。对于那些不可能通过自然恢复的严重退化或沙漠化的草原而言,人类的干预也许是可行的。
Objectives:Grazing-induced grassland degradation in Inner Mongolia is the primary impediment to the sustainable socioeconomic development of the region. Understanding the natural succession processes of the degraded grasslands is necessary for managing these prestigious natural resources.Practical techniques are urgently needed to add or accelerate the restoration of the degraded grasslands. Effects of two mechanical intervention techniques,shallow plowing and harrowing,in facilitating restoration of a degraded Leymus chinensis grassland were assessed,in comparison with grassland natural recovery,based on a long-term motoring research.
Methods:The research was conducted in the Inner Mongolia Grassland Ecosystem Research Station of the Chinese Academy of Sciences,which is located in the Xilingol League of Inner Mongolia(43°20′N-44°00′N,116°06′E-117°05′E).The degraded L chinensis grassland was fenced in 1983(foe excluding grazing animals), when grassland community was dominated by Artemisia frigida,Cleistogenes squarrosa and Agropyron michnoi,etc.Shallow plowing and harrowing were applied on the grassland in designated areas in the same year.The plant species composition and biomass production of these treated grasslands and that in natural recovery were monitored following the treatments at plant growing season.The integrative analysis of the long-term data set resulted in following main findings:
Results:
1.Shallow plowing." The grasses were dominant in the whole restoration period, having higher herbage production and population density.Rhizomateous L.chinensis had extremely high biomass,especially in the early stage.Grassland production was not significantly related with rainfall in the season,but related with restoration years. This indicated that the changes in plant community structures over the period were related with restoration following the treatment.The species diversity(DI) and evenness indices of plant community(in terms of species biomass percentage) decreased first then increased during the observation period.DI had following relation with restoration years(x):DI=0.0004x~2 +0.0113x+0.228;r=0.7777.
Four restoration stages can be distinguished over the restoration period in grassland community succession:(1) L.chinensis + annual or biennial forbs community(1983-85);(2) L.chinensis predominant community(1986-91);(3) L. chinensis + Agropyron michnoi + perennial forbs community(1992-96);and(4) L. chinensis + bunchgrasses + perennial forbs community(1997-07),which were very similar to the climax community in the region.
2.Harrowing:Grasses restored following the harrowing,gradually becoming dominant in plant communities.Rhizomateous L.chinensis was facilitated by the treatment,which increased quickly at the early stage,but decreased later due to the increase of other species.Semi-shrub and bunchgrasses were inhibited by the treatment at the early stage.Species diversity(DI) and evenness indices also decreased in the early stage,but increased later.DI had following relation with restoration years(x):DI= 0.001x~2-0.042x + 0.896;r = 0.7169 The four stages following the treatment are(1) L.chinensis + annual and biennual forbs + Artemisia frigida community(1983-86);(2) L.chinensis + short bunchgrasses + forbs community(1987-96);(3) L.chinensis + Agropyron michnoi + bunchgrasses + forbs community(1997-2000);and(4) L.chinensis + Stipa grandis + short bunchgrasses community,in which L.chinensis had the same important value as it was in climax community,and community structure was very similar to natural climax community.
3.Natural recovery:The change in plant community was gradual.The biomass and population density of grasses increased while that of Artemisia frigida (semi-shrub) deceased in the early recovery stage,but the composition of species biomass maintained a relatively stable proportion in the later stage.The recovery of grassland biomass was slower than that of the shallow-plowed and harrowed grasslands.The species diversity and evenness indices had no significant change trend during the period.The four recovery stages were(1) Artemisia frigida + short bunchgrasses + L.chinensis community(1983-85);(2) Transitional stage from A. frigid to L chinensis dominant communities(1986-88);(3) L.chinensis + Agropyron michnoi + short bunchgrasses community(1989-91);and(4) L.chinensis + Stipa grandis + short bunchgrasses communities(1991-2007).
4.Environmental changes:Soil physical and chemical properties improved during the restoration for all the treatments,but the differences among the treatments last for the whole period.Soil bulk density of the plowed,harrowed grassland and in that of natural recovery in the exclosure(animal grazing excluded) were 22.3%,21.6%and 11.5%lower than that of public open grazing grassland at the end of observation period.Soil moisture content increased by 93.71%,91.36%and 86.39%respectively when compared with the soil moisture in grazed grassland.The soil bacteria number in soil layer L1 also increased by 91.56%,89.66%and 73.56%,and Actinomyces increased by 41.72%,38.89%and 22.63%,in comparison with that in the soil of grazed grassland.These results indicated that shallow plowing and harrowing benefited soil micro-organisms.
5.Natural recovery was superior to the two mechanical interventions for restoring the widely distributed A.frigida grassland,derived from L chinensis or other native grasslands.Human intervention might be justifiable for the severely-degraded or desertified grassland beyond the threshold of natural regeneration.
本研究在锡林郭勒盟,中国科学院内蒙古草原生态系统定位研究站样地完成。研究区位于北纬43°20′N～44°00′N,116°06′E～117°05′E。改良前植被为因过度放牧导致的羊草草原退化变体,即含小叶锦鸡儿(Caraganamicrophylla)灌丛的冷蒿(Artemisia frigida)+丛生小禾草+羊草群落。于1983年6月,用围栏封育、耙地、浅耕翻三个措施进行处理,围栏内样地没有进行任何方式的利用。采用植物样方法、土壤常规分析方法和土壤微生物分析方法对经过25年恢复演替草原样地内地植物群落特征、土壤理化性状和土壤微生物特征进行了分析,结果如下:
1.浅耕翻:群落地上生物量和密度组成中,禾本科占主导地位。最高时禾本科生物量占到总生物量的95%,总密度的90%以上,在整个恢复演替过程中一直保持在60%以上。以羊草为主的根茎禾草随着演替进程的延续一直处于优势地位。高丛生禾草在恢复演替的后期增加。群落总地上生物量与降水相关关系不显著,主要受改良措施造成的群落结构变化的影响。浅耕翻处理恢复过程中多样性指数、均匀性指数总变化趋势为开口向上的抛物线状。以生物量为指标的多样性指数(DI)与恢复年限(x)之间的关系可以用如下方程表示:DI=0.0004x~2+0.0113x+0.228(r=0.7777).浅耕翻处理恢复过程中群落主要植物的种群密度、生物量与多样性指数存在着显著的相关关系。浅耕翻改良处理,25年的恢复演替过程可划分为四个阶段:第一阶段,即处理初期(1983～1985),群落类型为羊草+一、二年生杂类草群落;第二阶段(1986～1991年)的群落类型为羊草群落;第三阶段(1992～1996)的群落类型为羊草+冰草+多年生杂类草的过渡型群落类型;第四阶段,即演替后期(1997～2007),群落类型为羊草+丛生禾草、多年生杂类草群落,此时羊草的重要值已基本上类似于原生群落的。
2.耙地处理:群落总地上生物量和密度由于耙地改良处理持续恢复,禾本科植物生物量作为生物量构成的主体,处于建群地位。根茎禾草作为耙地的受益者在恢复演替的前期呈现持续增加的趋势,中期呈现高位维持,而后有所下降。小半灌木和丛生禾草受耙地处理的抑制,多年生植物在演替后期增加,使群落更接近与天然草原群落。群落总地上生物量与降水的相关关系不显著,主要与改良措施造成的群落结构变化有关。耙地处理后,均匀性指数、多样性指数的变化随着恢复演替的进程前期呈现波动下降的趋势,恢复中后期有所升高。以生物量为指标的多样性指数(DI)与恢复年限(x)之间的关系可以用如下方程表示:DI=0.001x~2-0.042x+0.896(r=0.7169)耙地处理后羊草的密度和生物量的变化与多样性指数具有极显著的负相关关系。耙地改良处理25年的恢复演替过程可划分为四个阶段:第一阶段,即处理初期(1983～1986),群落类型为羊草+一、二年生杂类草+冷蒿、冰草群落;第二阶段(1987～1996年),群落类型为羊草+小丛生禾草+冰草、杂类草群落;第三阶段(1997～2000),群落类型为羊草+冰草+丛生禾草、多年生杂类草的过渡型群落类型;第四阶段(2001～2007),群落类型为羊草+大针茅+冰草、丛生禾草群落,此时羊草的重要值已基本上接近原生群落,而随着针茅等丛生禾草的增加,群落更趋近于天然群落。
3.围栏封育自然恢复:围栏封育自然恢复群落的地上生物量和密度就功能群组成来看,根茎禾草在恢复演替的前9年呈现快速增加、缓慢增加并维持的趋势,丛生禾草后期增加,在退化群落中占主导地位的以冷蒿为主的小半灌木持续减少并维持较低的比例。与前两种改良措施相比,围栏封育自然恢复处理下群落生物量的恢复较滞后。退化草原围栏封育自然恢复过程中植物群落的多样性指数和均匀性指数总体上基本稳定,变幅不大。退化草原围栏封育自然恢复过程中羊草、冰草、冷蒿的生物量和密度的变化与多样性指数具有显著的负相关关系,星毛委陵菜的生物量和密度的变化与多样性指数具有显著的正相关关系。围封自然恢复,25年的恢复演替过程可划分为四个阶段:第一阶段,为封育后1—3年(1983～1985),群落类型为冷蒿+丛生小禾草+羊草群落;第二阶段(1986～1988)为由冷蒿群落向羊草群落过渡的演替阶段;第三阶段(1989～1991)群落类型为羊草+冰草、冷蒿+丛生禾草群落;第四阶段(1992～2007)群落已演变成羊草+针茅+杂类草群落,已基本接近原生羊草草原群落。随着恢复年限的延续,针茅种群的群落重要性将继续增加。
4.环境差异变化:经过24年的恢复改良,土壤理化性质明显改善,但仍有差别。耕翻、耙地、自然恢复处理后的土壤容重分别比围栏外降低22.30%、21.62%、11.49%,且浅耕翻和耙地处理后的土壤容重显著低于自然恢复处理后的土壤容重;土壤含水量分别比围栏外土壤含水量提高93.71%、91.36%、86.39%,改良措施之间差异不显著;土壤L1层细菌的数量分别比围栏外增加了91.56%、89.66%、73.56%,改良措施之间并无显著差异;真菌的数量分别比围栏外提高了53.06%、50.34%、22.45%,自然恢复显著低于另外两种改良方法;放线菌的数量分别比围栏外提高了41.72%、38.89%、22.63%,浅耕翻、耙地处理后的显著高于自然恢复的。因此,浅耕翻和耙地更有利于土壤微生物数量的增加。
5.几种措施的长期效果评价
就与天然羊草草原的相似度来讲,对于恢复A.frigida草原这种广泛分布的退化草原,除去了食草动物的自然恢复要优于两种机械改良的方法。对于那些不可能通过自然恢复的严重退化或沙漠化的草原而言,人类的干预也许是可行的。
Objectives:Grazing-induced grassland degradation in Inner Mongolia is the primary impediment to the sustainable socioeconomic development of the region. Understanding the natural succession processes of the degraded grasslands is necessary for managing these prestigious natural resources.Practical techniques are urgently needed to add or accelerate the restoration of the degraded grasslands. Effects of two mechanical intervention techniques,shallow plowing and harrowing,in facilitating restoration of a degraded Leymus chinensis grassland were assessed,in comparison with grassland natural recovery,based on a long-term motoring research.
Methods:The research was conducted in the Inner Mongolia Grassland Ecosystem Research Station of the Chinese Academy of Sciences,which is located in the Xilingol League of Inner Mongolia(43°20′N-44°00′N,116°06′E-117°05′E).The degraded L chinensis grassland was fenced in 1983(foe excluding grazing animals), when grassland community was dominated by Artemisia frigida,Cleistogenes squarrosa and Agropyron michnoi,etc.Shallow plowing and harrowing were applied on the grassland in designated areas in the same year.The plant species composition and biomass production of these treated grasslands and that in natural recovery were monitored following the treatments at plant growing season.The integrative analysis of the long-term data set resulted in following main findings:
Results:
1.Shallow plowing." The grasses were dominant in the whole restoration period, having higher herbage production and population density.Rhizomateous L.chinensis had extremely high biomass,especially in the early stage.Grassland production was not significantly related with rainfall in the season,but related with restoration years. This indicated that the changes in plant community structures over the period were related with restoration following the treatment.The species diversity(DI) and evenness indices of plant community(in terms of species biomass percentage) decreased first then increased during the observation period.DI had following relation with restoration years(x):DI=0.0004x~2 +0.0113x+0.228;r=0.7777.
Four restoration stages can be distinguished over the restoration period in grassland community succession:(1) L.chinensis + annual or biennial forbs community(1983-85);(2) L.chinensis predominant community(1986-91);(3) L. chinensis + Agropyron michnoi + perennial forbs community(1992-96);and(4) L. chinensis + bunchgrasses + perennial forbs community(1997-07),which were very similar to the climax community in the region.
2.Harrowing:Grasses restored following the harrowing,gradually becoming dominant in plant communities.Rhizomateous L.chinensis was facilitated by the treatment,which increased quickly at the early stage,but decreased later due to the increase of other species.Semi-shrub and bunchgrasses were inhibited by the treatment at the early stage.Species diversity(DI) and evenness indices also decreased in the early stage,but increased later.DI had following relation with restoration years(x):DI= 0.001x~2-0.042x + 0.896;r = 0.7169 The four stages following the treatment are(1) L.chinensis + annual and biennual forbs + Artemisia frigida community(1983-86);(2) L.chinensis + short bunchgrasses + forbs community(1987-96);(3) L.chinensis + Agropyron michnoi + bunchgrasses + forbs community(1997-2000);and(4) L.chinensis + Stipa grandis + short bunchgrasses community,in which L.chinensis had the same important value as it was in climax community,and community structure was very similar to natural climax community.
3.Natural recovery:The change in plant community was gradual.The biomass and population density of grasses increased while that of Artemisia frigida (semi-shrub) deceased in the early recovery stage,but the composition of species biomass maintained a relatively stable proportion in the later stage.The recovery of grassland biomass was slower than that of the shallow-plowed and harrowed grasslands.The species diversity and evenness indices had no significant change trend during the period.The four recovery stages were(1) Artemisia frigida + short bunchgrasses + L.chinensis community(1983-85);(2) Transitional stage from A. frigid to L chinensis dominant communities(1986-88);(3) L.chinensis + Agropyron michnoi + short bunchgrasses community(1989-91);and(4) L.chinensis + Stipa grandis + short bunchgrasses communities(1991-2007).
4.Environmental changes:Soil physical and chemical properties improved during the restoration for all the treatments,but the differences among the treatments last for the whole period.Soil bulk density of the plowed,harrowed grassland and in that of natural recovery in the exclosure(animal grazing excluded) were 22.3%,21.6%and 11.5%lower than that of public open grazing grassland at the end of observation period.Soil moisture content increased by 93.71%,91.36%and 86.39%respectively when compared with the soil moisture in grazed grassland.The soil bacteria number in soil layer L1 also increased by 91.56%,89.66%and 73.56%,and Actinomyces increased by 41.72%,38.89%and 22.63%,in comparison with that in the soil of grazed grassland.These results indicated that shallow plowing and harrowing benefited soil micro-organisms.
5.Natural recovery was superior to the two mechanical interventions for restoring the widely distributed A.frigida grassland,derived from L chinensis or other native grasslands.Human intervention might be justifiable for the severely-degraded or desertified grassland beyond the threshold of natural regeneration.
引文
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