封育对荒漠草原两种植物群落土壤种子库的影响
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摘要
土壤种子库作为植被的潜在种源,对荒漠草原植被恢复、群落演替具有重要的指示意义。以宁夏盐池县荒漠草原苦豆子群落和芨芨草群落为研究对象,在2种群落封育和未封育的草地上,选取地势相对平坦、环境条件较为一致的25m×30m的4个大的样方进行群落调查;并在4个大的样方内取0~2cm、2~5cm和5~10cm土层的土样,采取种子过筛直接辨认和种子萌发试验来进行土壤种子库的鉴定。分析了两种植物群落封育前后土壤种子库物种组成、种子密度、多样性及相似性等特征。结果表明:封育可提高两种植物群落土壤种子库中物种数目;而对两种植物群落土壤种子库种子密度的影响不同,苦豆子群落种子密度封育内外差异较小,芨芨草群落封育内种子密度显著提高。随着土层的加深,两种植物群落封育内外土壤种子库物种数和种子密度均呈递减趋势,封育后两种植物群落不同土层物种数均有增加,但封育对两种植物群落土壤种子库中种子密度的垂直分布影响不同。封育可提高两种植物群落土壤种子库的多样性、均匀度和丰富度,但两种植物群落土壤种子库的多样性特征对封育的响应不同,芨芨草群落在荒漠草原的稳定性更高。两种植物群落封育内外土壤种子库相似性相对较高,苦豆子群落封育后有向芨芨草群落演替的趋势。
The soil seed bank(SSB),as a potential source of vegetation,has significance for vegetation restoration and community succession on the desert steppe.In this study,we analyzed the SSB of two plant communities(Sophora alopecuroides and Achnatherum splendens communities),fenced and unfenced,on the desert steppe in Yanchi County,Ningxia Province.Four relatively even and flat quadrats(25 m×30 meach)of S.alopecuroides and A.splendens communities,fenced and unfenced,were selected for these analyses.The soil samples from the four quadrats were separated into three layers(0-2 cm,2-5 cm,and 5-10 cm)and sieved,and then the species of the seeds in the SSB were identified by direct observations and then by germination tests.We compared the characteristics of two communities(SSB species composition,seed density,diversity,and similarity)between fenced(enclosed)and unfenced conditions.The results showed that:1)Enclosure increased the number of species in the SSB in both communities,but its effects on seed density in the SSB differed between the two communities.The seed density of the S.alopecuroides community SSB was slightly different between the fenced and unfenced areas,while that of the A.splendens community SSB was markedly higher inside the fenced area.2)The number of species in the SSB and the seed density tended to decrease with increasing soil depth in both communities,inside and outside the fence.Although the number of species in different soil layers was greater in the fenced area for both communities,the effect of enclosure on the vertical distribution of seed density in the SSB differed between the two communities.3)The diversity,evenness,and abundance of the SSB were greater inside the fenced area than outside the fenced area for both communities,but the effects of enclosure on SSB diversity differed between the two communities.The S.alopecuroides community was more stable than the A.splendens community on the desert steppe.4)The SSBs of the two communities showed relatively high similarity,and there was a trend towards succession of the S.alopecuroides community to the A.splendens community after enclosure.
The soil seed bank(SSB),as a potential source of vegetation,has significance for vegetation restoration and community succession on the desert steppe.In this study,we analyzed the SSB of two plant communities(Sophora alopecuroides and Achnatherum splendens communities),fenced and unfenced,on the desert steppe in Yanchi County,Ningxia Province.Four relatively even and flat quadrats(25 m×30 meach)of S.alopecuroides and A.splendens communities,fenced and unfenced,were selected for these analyses.The soil samples from the four quadrats were separated into three layers(0-2 cm,2-5 cm,and 5-10 cm)and sieved,and then the species of the seeds in the SSB were identified by direct observations and then by germination tests.We compared the characteristics of two communities(SSB species composition,seed density,diversity,and similarity)between fenced(enclosed)and unfenced conditions.The results showed that:1)Enclosure increased the number of species in the SSB in both communities,but its effects on seed density in the SSB differed between the two communities.The seed density of the S.alopecuroides community SSB was slightly different between the fenced and unfenced areas,while that of the A.splendens community SSB was markedly higher inside the fenced area.2)The number of species in the SSB and the seed density tended to decrease with increasing soil depth in both communities,inside and outside the fence.Although the number of species in different soil layers was greater in the fenced area for both communities,the effect of enclosure on the vertical distribution of seed density in the SSB differed between the two communities.3)The diversity,evenness,and abundance of the SSB were greater inside the fenced area than outside the fenced area for both communities,but the effects of enclosure on SSB diversity differed between the two communities.The S.alopecuroides community was more stable than the A.splendens community on the desert steppe.4)The SSBs of the two communities showed relatively high similarity,and there was a trend towards succession of the S.alopecuroides community to the A.splendens community after enclosure.
引文
[1]Deng B,Ren G H,Liu Z Y,et al.Effect of 3-year fencing on soil seed banks of three alpine grassland communities.Acta Prataculturae Sinica,2012,21(5):23-31.邓斌,任国华,刘志云,等.封育三年对三种高寒草地群落土壤种子库的影响.草业学报,2012,21(5):23-31.
[2]Yang N,Fu M Y,Yang M Y,et al.Soil seed bank properties under different land used models on sloping land with purple soils in Hengyang of Hunan Province,south central China.Acta Botanica Boreali Occidentalia Sinica,2014,34(11):2324-2330.杨宁,付美云,杨满元,等.衡阳紫色土丘陵坡地不同土地利用模式下土壤种子库特征.西北植物学报,2014,34(11):2324-2330.
[3]Zhou H K,Zhou L,Liu W,et al.Influence of fencing on degraded Kobresia humilis meadows and nondegraded.Grassland of China,2003,25(5):15-22.周华坤,周立,刘伟,等.封育措施对退化与未退化矮嵩草草甸的影响.中国草地学报,2003,25(5):15-22.
[4]Yang L,Wang Y R,Yu J D.Review of research on seed banks of desert regions.Acta Prataculturae Sinica,2010,19(2):227-234.杨磊,王彦荣,余进德.干旱荒漠区土壤种子库研究进展.草业学报,2010,19(2):227-234.
[5]LüG F,Wu Y S,Li H,et al.Microorganisms,soil nutrient and enzyme activity changing with the proceeding of desert steppe degradation in Inner Mongolia.Journal of Desert Research,2010,30(1):104-109.吕桂芬,吴永胜,李浩,等.荒漠草原不同退化阶段土壤微生物、土壤养分及酶活性的研究.中国沙漠,2010,30(1):104-109.
[6]Sun H L.Chinese ecosystems.Beijing:Science Press,2005:550-665.孙鸿烈.中国生态系统.北京:科学出版社,2005:550-665.
[7]Wang G H,Wang J J,Tao L B,et al.Effects of enclosure on soil aggregate distribution and stability of desert steppe in Ningxia.Acta Agrestia Sinica,2017,25(1):76-81.王国会,王建军,陶利波,等.围封对宁夏荒漠草原土壤团聚体组成及其稳定性的影响.草地学报,2017,25(1):76-81.
[8]Shao W S,Li G Q,Chen K Y,et al.Comparative studies on soil enzymic activities of four kinds of common plant communities in desert steppe.Acta Botanica Boreali Occidentalia Sinica,2016,36(3):579-587.邵文山,李国旗,陈科元,等.荒漠草原4种常见植物群落土壤酶活性比较.西北植物学报,2016,36(3):579-587.
[9]Li F R,Zhao L Y,Wang S F,et al.Effects of enclosure management on the structure of soil seed bank and standing vegetation in degraded sandy grasslands of eastern Inner Mongolia.Acta Prataculturae Sinica,2003,12(4):90-99.李锋瑞,赵丽娅,王树芳,等.封育对退化沙质草地土壤种子库与地上群落结构的影响.草业学报,2003,12(4):90-99.
[10]Ma Q L,Lu Q,Wei L Y,et al.Varying characteristics of soil seed banks during the succession process of Nitraria tangutorum vegetation in an arid desert area.Acta Ecologica Sinica,2015,35(7):2285-2294.马全林,卢琦,魏林源,等.干旱荒漠白刺灌丛植被演替过程土壤种子库变化特征.生态学报,2015,35(7):2285-2294.
[11]Liu R X,Zhan J,Shi Z H,et al.Soil seed bank and its correlations with aboveground vegetation and environmental factors in water level fluctuating zone of Danjiangkou Reservoir,Central China.Chinese Journal of Applied Ecology,2013,24(3):801刘瑞雪,詹娟,史志华,等.丹江口水库消落带土壤种子库与地上植被和环境的关系.应用生态学报,2013,24(3):801-808.
[12]Zhang J T.Quantitative ecology.Beijing:Science Press,2004:77-92.张金屯.数量生态学.北京:科学出版社,2004:77-92.
[13]Yan R R,Wei Z J,Xin X P,et al.Effects of grazing systems on germinable soil seed bank of desert steppe.Journal of Desert Research,2011,31(3):703-709.闫瑞瑞,卫智军,辛晓平,等.放牧制度对荒漠草原可萌发土壤种子库的影响.中国沙漠,2011,31(3):703-709.
[14]Shang Z H,Long R J,Ma Y S,et al.Soil seed banks of degraded alpine grassland in headwater region of the Yellow River:quantities and dynamics of seed germination.Chinese Journal of Applied&Environmental Biology,2006,12(3):313-317.尚占环,龙瑞军,马玉寿,等.黄河源区退化高寒草地土壤种子库:种子萌发的数量和动态.应用与环境生物学报,2006,12(3):313-317.
[15]Silvertown J W.Introduction to plant population ecology.London and New York:Longman,1982:22-37.
[16]Ma M J,Zhou X H,LüZ W,et al.A comparison of the soil seed bank in an enclosed vs.a degraded alpine meadow in the eastern Tibetan Plateau.Acta Ecologica Sinica,2009,29(7):3658-3664.马妙君,周显辉,吕正文,等.青藏高原东缘封育和退化高寒草甸种子库差异.生态学报,2009,29(7):3658-3664.
[17]Li H Y,Mo X Q,Hao C.A review of study on soil seed bank in the past thirty years.Ecology and Environmental Sciences,2009,18(2):731-737.李洪远,莫训强,郝翠.近30年来土壤种子库研究的回顾与展望.生态环境学报,2009,18(2):731-737.
[18]Bekker R M,Bakker J P,Grandin U,et al.Seed size,shape and vertical distribution in the soil:indicators of seed longevity.Functional Ecology,1998,12(5):834-842.
[19]Ambrosiol,Doradoj,Delmonte J P.Assessment of the sample size to estimate the weed seed bank in soil.Weed Research,1997,37:129-137.
[20]Zeng Y J,Wang Y R,Nan Z B,et al.Soil seed banks of different grassland types of Alashan arid desert region,Inner Mongolia.Chinese Journal of Applied Ecology,2003,14(9):1457-1463.曾彦军,王彦荣,南志标,等.阿拉善干旱荒漠区不同植被类型土壤种子库研究.应用生态学报,2003,14(9):1457-1463.
[21]Ma M,Ma Z,Du G.Effects of water level on three wetlands soil seed banks on the Tibetan Plateau.Plos One,2014,9(7):e101458.
[22]Ma M,Zhou X,Du G.Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau.Plant and Soil,2013,369(1):283-295.
[23]Zhao L P,Cheng J M,Wan H E,et al.Characteristics of soil seed banks in grassland under fencing and grazing in Loess Plateau.Pratacultural Science,2008,25(10):78-83.赵凌平,程积民,万惠娥,等.黄土高原草地封育与放牧条件下土壤种子库特征.草业科学,2008,25(10):78-83.
[24]Huang X Y,Wang K,Wang Y T,et al.Soil seed bank dynamics during natural restoration by fence in a typical steppe of Inner Mongolia.Acta Agrestia Sinica,2011,19(1):38-42.黄欣颖,王堃,王宇通,等.典型草原封育过程中土壤种子库的变化特征.草地学报,2011,19(1):38-42.
[25]Shen Y,Liu C F,Ma H B,et al.Response of a soil seed bank to modes of grassland management on a desert steppe.Acta Ecologica Sinica,2015,35(14):4725-4732.沈艳,刘彩凤,马红彬,等.荒漠草原土壤种子库对草地管理方式的响应.生态学报,2015,35(14):4725-4732.
[26]Liu X K,Liu M S,Huang Z,et al.Pattern analysis of inter specific relationships in four arid communities in sand lake Ningxia Hui Autonomous region,China.Chinese Journal of Plant Ecology,2009,33(2):320-330.刘小恺,刘茂松,黄峥,等.宁夏沙湖4种干旱区群落中主要植物种间关系的格局分析.植物生态学报,2009,33(2):320-330.
[27]Chen L M G,Liu M S,Huang Z,et al.Distribution pattern of root biomass and inter specific relationship in Achnatherum splenoens-Sophora alopecuroides community in Northwest China.Chinese Journal of Plant Ecology,2009,33(4):748-754.晨乐木格,刘茂松,黄峥,等.我国西北地区芨芨草——苦豆子群落根系分布与种间关系.植物生态学报,2009,33(4):748-754.
[2]Yang N,Fu M Y,Yang M Y,et al.Soil seed bank properties under different land used models on sloping land with purple soils in Hengyang of Hunan Province,south central China.Acta Botanica Boreali Occidentalia Sinica,2014,34(11):2324-2330.杨宁,付美云,杨满元,等.衡阳紫色土丘陵坡地不同土地利用模式下土壤种子库特征.西北植物学报,2014,34(11):2324-2330.
[3]Zhou H K,Zhou L,Liu W,et al.Influence of fencing on degraded Kobresia humilis meadows and nondegraded.Grassland of China,2003,25(5):15-22.周华坤,周立,刘伟,等.封育措施对退化与未退化矮嵩草草甸的影响.中国草地学报,2003,25(5):15-22.
[4]Yang L,Wang Y R,Yu J D.Review of research on seed banks of desert regions.Acta Prataculturae Sinica,2010,19(2):227-234.杨磊,王彦荣,余进德.干旱荒漠区土壤种子库研究进展.草业学报,2010,19(2):227-234.
[5]LüG F,Wu Y S,Li H,et al.Microorganisms,soil nutrient and enzyme activity changing with the proceeding of desert steppe degradation in Inner Mongolia.Journal of Desert Research,2010,30(1):104-109.吕桂芬,吴永胜,李浩,等.荒漠草原不同退化阶段土壤微生物、土壤养分及酶活性的研究.中国沙漠,2010,30(1):104-109.
[6]Sun H L.Chinese ecosystems.Beijing:Science Press,2005:550-665.孙鸿烈.中国生态系统.北京:科学出版社,2005:550-665.
[7]Wang G H,Wang J J,Tao L B,et al.Effects of enclosure on soil aggregate distribution and stability of desert steppe in Ningxia.Acta Agrestia Sinica,2017,25(1):76-81.王国会,王建军,陶利波,等.围封对宁夏荒漠草原土壤团聚体组成及其稳定性的影响.草地学报,2017,25(1):76-81.
[8]Shao W S,Li G Q,Chen K Y,et al.Comparative studies on soil enzymic activities of four kinds of common plant communities in desert steppe.Acta Botanica Boreali Occidentalia Sinica,2016,36(3):579-587.邵文山,李国旗,陈科元,等.荒漠草原4种常见植物群落土壤酶活性比较.西北植物学报,2016,36(3):579-587.
[9]Li F R,Zhao L Y,Wang S F,et al.Effects of enclosure management on the structure of soil seed bank and standing vegetation in degraded sandy grasslands of eastern Inner Mongolia.Acta Prataculturae Sinica,2003,12(4):90-99.李锋瑞,赵丽娅,王树芳,等.封育对退化沙质草地土壤种子库与地上群落结构的影响.草业学报,2003,12(4):90-99.
[10]Ma Q L,Lu Q,Wei L Y,et al.Varying characteristics of soil seed banks during the succession process of Nitraria tangutorum vegetation in an arid desert area.Acta Ecologica Sinica,2015,35(7):2285-2294.马全林,卢琦,魏林源,等.干旱荒漠白刺灌丛植被演替过程土壤种子库变化特征.生态学报,2015,35(7):2285-2294.
[11]Liu R X,Zhan J,Shi Z H,et al.Soil seed bank and its correlations with aboveground vegetation and environmental factors in water level fluctuating zone of Danjiangkou Reservoir,Central China.Chinese Journal of Applied Ecology,2013,24(3):801刘瑞雪,詹娟,史志华,等.丹江口水库消落带土壤种子库与地上植被和环境的关系.应用生态学报,2013,24(3):801-808.
[12]Zhang J T.Quantitative ecology.Beijing:Science Press,2004:77-92.张金屯.数量生态学.北京:科学出版社,2004:77-92.
[13]Yan R R,Wei Z J,Xin X P,et al.Effects of grazing systems on germinable soil seed bank of desert steppe.Journal of Desert Research,2011,31(3):703-709.闫瑞瑞,卫智军,辛晓平,等.放牧制度对荒漠草原可萌发土壤种子库的影响.中国沙漠,2011,31(3):703-709.
[14]Shang Z H,Long R J,Ma Y S,et al.Soil seed banks of degraded alpine grassland in headwater region of the Yellow River:quantities and dynamics of seed germination.Chinese Journal of Applied&Environmental Biology,2006,12(3):313-317.尚占环,龙瑞军,马玉寿,等.黄河源区退化高寒草地土壤种子库:种子萌发的数量和动态.应用与环境生物学报,2006,12(3):313-317.
[15]Silvertown J W.Introduction to plant population ecology.London and New York:Longman,1982:22-37.
[16]Ma M J,Zhou X H,LüZ W,et al.A comparison of the soil seed bank in an enclosed vs.a degraded alpine meadow in the eastern Tibetan Plateau.Acta Ecologica Sinica,2009,29(7):3658-3664.马妙君,周显辉,吕正文,等.青藏高原东缘封育和退化高寒草甸种子库差异.生态学报,2009,29(7):3658-3664.
[17]Li H Y,Mo X Q,Hao C.A review of study on soil seed bank in the past thirty years.Ecology and Environmental Sciences,2009,18(2):731-737.李洪远,莫训强,郝翠.近30年来土壤种子库研究的回顾与展望.生态环境学报,2009,18(2):731-737.
[18]Bekker R M,Bakker J P,Grandin U,et al.Seed size,shape and vertical distribution in the soil:indicators of seed longevity.Functional Ecology,1998,12(5):834-842.
[19]Ambrosiol,Doradoj,Delmonte J P.Assessment of the sample size to estimate the weed seed bank in soil.Weed Research,1997,37:129-137.
[20]Zeng Y J,Wang Y R,Nan Z B,et al.Soil seed banks of different grassland types of Alashan arid desert region,Inner Mongolia.Chinese Journal of Applied Ecology,2003,14(9):1457-1463.曾彦军,王彦荣,南志标,等.阿拉善干旱荒漠区不同植被类型土壤种子库研究.应用生态学报,2003,14(9):1457-1463.
[21]Ma M,Ma Z,Du G.Effects of water level on three wetlands soil seed banks on the Tibetan Plateau.Plos One,2014,9(7):e101458.
[22]Ma M,Zhou X,Du G.Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau.Plant and Soil,2013,369(1):283-295.
[23]Zhao L P,Cheng J M,Wan H E,et al.Characteristics of soil seed banks in grassland under fencing and grazing in Loess Plateau.Pratacultural Science,2008,25(10):78-83.赵凌平,程积民,万惠娥,等.黄土高原草地封育与放牧条件下土壤种子库特征.草业科学,2008,25(10):78-83.
[24]Huang X Y,Wang K,Wang Y T,et al.Soil seed bank dynamics during natural restoration by fence in a typical steppe of Inner Mongolia.Acta Agrestia Sinica,2011,19(1):38-42.黄欣颖,王堃,王宇通,等.典型草原封育过程中土壤种子库的变化特征.草地学报,2011,19(1):38-42.
[25]Shen Y,Liu C F,Ma H B,et al.Response of a soil seed bank to modes of grassland management on a desert steppe.Acta Ecologica Sinica,2015,35(14):4725-4732.沈艳,刘彩凤,马红彬,等.荒漠草原土壤种子库对草地管理方式的响应.生态学报,2015,35(14):4725-4732.
[26]Liu X K,Liu M S,Huang Z,et al.Pattern analysis of inter specific relationships in four arid communities in sand lake Ningxia Hui Autonomous region,China.Chinese Journal of Plant Ecology,2009,33(2):320-330.刘小恺,刘茂松,黄峥,等.宁夏沙湖4种干旱区群落中主要植物种间关系的格局分析.植物生态学报,2009,33(2):320-330.
[27]Chen L M G,Liu M S,Huang Z,et al.Distribution pattern of root biomass and inter specific relationship in Achnatherum splenoens-Sophora alopecuroides community in Northwest China.Chinese Journal of Plant Ecology,2009,33(4):748-754.晨乐木格,刘茂松,黄峥,等.我国西北地区芨芨草——苦豆子群落根系分布与种间关系.植物生态学报,2009,33(4):748-754.