生物土壤结皮演替对高寒草原植被结构和土壤养分的影响
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摘要
生物土壤结皮(生物结皮)是退化高寒草原常见的地表覆盖物,具有调控土壤养分循环和影响植物群落结构和特征的作用。以青藏高原地区典型的高寒草原生态系统为研究对象,采用野外样方调查和室内分析相结合的方法,根据样方内生物结皮的发育程度将所有样方分为4个类型,并分析了青藏高原典型高寒草原生物结皮的发育演替对草本植物群落的结构、特征以及0-10 cm土壤理化性质的影响。结果表明,(1)在生物结皮由藻结皮向苔藓结皮发育的过程中,草本植物群落结构发生了变化,其中禾本科植物占比上升,莎草科植物和杂类草占比下降,而豆科植物占比无显著变化。(2)在草本植物群落中,禾本科和莎草科植物的重要值最高,溚草(Koeleria Pers.)、异针茅(Stipa aliena Keng)、高山嵩草(Kobresia pygmaea)、矮嵩草(Kobresia humilis)和矮火绒(Leontopodium nanum)是主要的优势物种。(3)生物结皮盖度与植被盖度呈极显著的负相关关系(P<0.01)。(4)生物结皮的发育演替增加了禾本科植物比例,减少了杂类草比例,增加了植物群落的多样性、高度和生物量。(5)无结皮、地衣结皮和苔藓结皮覆盖下0-10 cm土壤的全氮含量差异显著(P<0.05),而土壤全磷、有机碳含量、碳氮比和氮磷比无显著变化。由此可见,高寒草原生态系统下,生物结皮对植被和土壤的影响具有一定的复杂性和特殊性,需进一步探明其生态作用,以发挥其在退化草原治理中的重要作用。
Biological soil crust(biological crust) is a common surface covering in degraded alpine steppe, which has the function of regulating soil nutrient cycling and affecting plant community structure and traits. This study took typical alpine steppe ecosystem in Qinghai-Tibet Plateau as the research object and used the method of field quadrat sample and indoor analysis. All samples were divided into four types based on the development degree of biological crust to analyze the effects of biological crust development succession on vegetation community structure and traits and soil physical and chemical properties in 0-10 cm soil layer in typical alpine steppe of Qinghai-Tibet Plateau. Results showed that: plant community structure changed during the development process of biological crust from algal crust to moss crust development, among which the proportion of gramineous plants increased, cyperaceae plants and miscellaneous grass decreased, and leguminous plants had no significant change. In plant communities, Gramineous and Cyperaceae plants had the highest important values, and the main dominant species were Scylla Scylla., Stipa aliena Keng, Kobresia pygmaea, Kobresia humilis and Leontopodium nanum. There was a significant negative correlation between biological crust coverage and vegetation coverage(P<0.01). The development succession of biological crust increased the proportion of Gramineae plants,decreased the proportion of forb, and increased the diversity, height and biomass of plant communities. Below no crust, lichen crust and moss crust, total nitrogen content in 0-10 cm soil had significant difference(P<0.05), while there were no significant change in total phosphorus, organic carbon content, carbon and nitrogen ratio and nitrogen and phosphorus ratio. Therefore, in the alpine steppe ecosystem, the effects of biological crust on vegetation and soil has certain complexity and particularity, and its ecological role needs to be further explored in order to play an important role in the restoration of degraded grassland.
Biological soil crust(biological crust) is a common surface covering in degraded alpine steppe, which has the function of regulating soil nutrient cycling and affecting plant community structure and traits. This study took typical alpine steppe ecosystem in Qinghai-Tibet Plateau as the research object and used the method of field quadrat sample and indoor analysis. All samples were divided into four types based on the development degree of biological crust to analyze the effects of biological crust development succession on vegetation community structure and traits and soil physical and chemical properties in 0-10 cm soil layer in typical alpine steppe of Qinghai-Tibet Plateau. Results showed that: plant community structure changed during the development process of biological crust from algal crust to moss crust development, among which the proportion of gramineous plants increased, cyperaceae plants and miscellaneous grass decreased, and leguminous plants had no significant change. In plant communities, Gramineous and Cyperaceae plants had the highest important values, and the main dominant species were Scylla Scylla., Stipa aliena Keng, Kobresia pygmaea, Kobresia humilis and Leontopodium nanum. There was a significant negative correlation between biological crust coverage and vegetation coverage(P<0.01). The development succession of biological crust increased the proportion of Gramineae plants,decreased the proportion of forb, and increased the diversity, height and biomass of plant communities. Below no crust, lichen crust and moss crust, total nitrogen content in 0-10 cm soil had significant difference(P<0.05), while there were no significant change in total phosphorus, organic carbon content, carbon and nitrogen ratio and nitrogen and phosphorus ratio. Therefore, in the alpine steppe ecosystem, the effects of biological crust on vegetation and soil has certain complexity and particularity, and its ecological role needs to be further explored in order to play an important role in the restoration of degraded grassland.
引文
COXSON D S,2002.Biological soil crusts:structure,function,and management[J].Biological Conservation,108(1):129-130.
BELNAP J,MILLER D M,BEDFORD D R,et al.,2014.Pedological and geological relationships with soil lichen and moss distribution in the eastern Mojave Desert,CA,USA[J].Journal of Arid Environments,106:45-57.
BELNAP J,PRASSE R,HARPER K T,2001.Influence of biological soil crusts on soil environments and vascular plants[M].Berlin:Springer,281-300.
BELNAP J,2002.Nitrogen fixation in biological soil crusts from southeast Utah,USA[J].Biology and Fertility of Soils,35(2):128-135.
BELNAP,JAYNE,2003.Biological soil crusts in deserts:a short review of their role in soil fertility,stabilization,and water relations[J].Algological Studies,109(1):113-126.
BLISS LCGOLD W G,1999.Vascular plant reproduction,establishment,and growth and the effects of cryptogamic crusts within a polar desert ecosystem,Devon Island,N.W.T.Canada[J].Canadian Journal of Botany,77(5):623-636.
BRIGGS A,MORGAN J W,2008.Morphological diversity and abundance of biological soil crusts differ in relation to landscape setting and vegetation type[J].Australian Journal of Botany,56(3):246-253.
COPPOLA A,BASILE A,WANG X,et al.,2011.Hydrological behaviour of microbiotic crusts on sand dunes:Example from NW China comparing infiltration in crusted and crust-removed soil[J].Soil&Tillage Research,117:34-43.
DEINES L,ROSENTRETER R,ELDRIDGE D J,et al.,2007.Germination and seedling establishment of two annual grasses on lichen-dominated biological soil crusts[J].Plant&Soil,295(1-2):23-35.
ELBERT W,ANDREAE M O,2009.Microbiotic crusts on soil,rock and plants:Neglected major players in the global cycles of carbon and nitrogen[J].Biogeo Sciences Discuss,6:6983-7015.
PRASSE R,BORNKAMM R,2000.Effect of microbiotic soil surface crusts on emergence of vascular plants[J].Plant Ecology,150(1-2):65-75.
RIVERAAGUILAR V,GODINEZALVAREZ H,MANUELLCACHEUX I,et al.,2005.Physical effects of biological soil crusts on seed germination of two desert plants under laboratory conditions[J].Journal of Arid Environments,63(1):344-352.
ZHAO Y,XU M,BELNAP J,2010.Potential nitrogen fixation activity of different aged biological soil crusts from rehabilitated grasslands of the hilly Loess Plateau,China[J].Journal of Arid Environments,74(10):1186-1191.
陈兰周,刘永定,李敦海,等,2003.荒漠藻类及其结皮的研究[J].中国科学基金,17(2):90-93.CHEN L Z,LIU Y D,LI J H,et al.,2003.The research process/progress of desert algae and crus[J].Bulletin of National Science Foundation of China,17(2):90-93.
陈孟晨,张景光,冯丽,等,2017.沙坡头地区生物结皮覆盖区土壤种子库组成及垂直分布特征[J].生态学报,37(22):243-252.CHEN M C,ZHANG J G,FENG L,et al.,2017.The composition and vertical distribution characteristics of soil seed banks in soil coverage with biocrusts in the Shapotou Region[J].Acta Ecological Sinica,37(22):243-252.
贺郝钰,侯春梅,迟秀丽,等,2014.国际生物土壤结皮研究发展态势文献计量分析[J].生态学报,34(4):1035-1041.HE H Y,HOU C M,CHI X L,et al.,2014.Bibliometric analysis of the development trend of international biological soil crust research[J].Acta Ecological Sinica,34(4):1035-1041.
李国栋,张元明,2014.生物土壤结皮与种子附属物对4种荒漠植物种子萌发的交互影响[J].中国沙漠,34(3):725-731.LI G D,ZHANG Y M,2014.Interactive Effects of Biological Soil Crusts and Seed Appendages on Seed Germination of Four Desert Species[J].Journal of Desert Research,34(3):725-731.
李新荣,张景光,2000.干旱沙漠区土壤微生物结皮及其对固沙植被影响的研究[J].植物学报:英文版,42(9):965-970.LI X R,ZHANG J G,2000.Study on soil microbiotic crust and its influences on sand-fixing vegetation in arid desert region[J].Acta Botanica Sinica,42(9):965-970.
刘利霞,张宇清,吴斌,2007.生物结皮对荒漠地区土壤及植物的影响研究述评[J].中国水土保持科学,5(6):106-112.LIU L X,ZHANG Y Q,WU B,2007.Review on the influences of biological soil crusts on soil and vegetation in desertified areas[J].Science of Soil and Water Conservation,5(6):106-112.
聂华丽,张元明,吴楠,等,2009.生物结皮对5种不同形态的荒漠植物种子萌发的影响[J].植物生态学报,33(1):161-170.NIE H L,ZHANG Y M,WU N,et al,2009.Effects of biological crusts on the germination of five desert vascular plants with different seed morphologies[J].Chinese Journal of Plant Ecology,33(1):161-170.
宋阳,严平,张宏,等,2004.荒漠生物结皮研究中的几个问题[J].干旱区研究,21(4):439-443.SONG Y,YAN P,ZHANG H,et al.,2004.Recent progress and some existing problems in the study on the desert microbiotic crusts[J].Arid Zone Research,21(4):439-443.
王华磊,贾长虹,李潞滨,2013.土壤结皮及研究进展[J].河北联合大学学报(自然科学版),35(2):107-112.WANG H L,JIA C H,LI L B,2013.Soil Crusts and Research Progress[J].Journal of Hebei United University(Natural Science Edition),35(2):107-112.
李以康,欧阳经政,林丽,等,2015.高寒草甸植被退化过程中生物土壤结皮演变特征[J].生态学杂志,34(8):2238-2244.LI Y K,OUYANG J Z,LIN L,et al.,2015.Evolution characteristics of biological soil crusts(BSCs)during alpine meadow degrada-tion[J].Chinese Journal of Ecology,34(8):2238-2244.
王蕊,朱清科,赵磊磊,等,2011.黄土高原土壤生物结皮对植物种子出苗和生长的影响[J].干旱区研究,28(5):800-807.WANG R,ZHU Q K,ZHAO L L,et al.,2011.Effects of biological soil crusts on seed emergence and seedling growth in Loess Plateau,North Shaanxi province[J].Arid Zone Research,28(5):800-807.
肖波,赵允格,邵明安,2007.陕北水蚀风蚀交错区两种生物结皮对土壤理化性质的影响[J].生态学报,27(11):4662-4670.XIAO B,ZHAO Y G,SHAO M A,2007.Effects of biological soil crust on soil physicochemical properties in water-wind erosion crisscross region,Northern Shaanxi province,China[J].Acta Ecologica Sinica,27(11):4662-4670.
张国秀,赵允格,许明祥,等,2012.黄土丘陵区生物结皮对土壤磷素有效性及碱性磷酸酶活性的影响[J].植物营养与肥料学报,18(3):621-628.ZHANG G X,ZHAO Y G,XUN M X,et al.,2012.Impacts of biological soil crust on availability of phosphorus and phosphatase activity in hilly regions of the Loess Plateau,China[J].Plant Nutrition and Fertilizer Science,18(3):621-628.
张侃侃,卜崇峰,高国雄,2011.黄土高原生物结皮对土壤水分入渗的影响[J].干旱区研究,28(5):808-812.ZHANG K K,PU C F,GAO G X,2011.Effect of microbiotic crust on soil water infiltration in the Loess Plateau[J].Arid Zone Research,28(5):808-812.
赵允格,许明祥,王全九,等,2006.黄土丘陵区退耕地生物结皮对土壤理化性状的影响[J].自然资源学报,21(3):441-448.ZHAO Y G,XU M X,WANG Q J,et al.,2012.Impact of biological soil crust on soil physical and chemical properties of Rebabilitated grassland in Hilly Loess Plateau,China[J].Journal of Natural Resources,21(3):441-448.
赵允格,许明祥,王全九,等,2006.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,17(8):1429-1434.ZHAO Y G,XU M X,WANG Q J,et al.,2006.Physicaland chem ical properties of soilbio-crust on rehabilitated grassland in hilly LoessPlateau of China[J].Chinese Journal of Applied Ecology,17(8):1429-1434.
庄伟伟,张元明,2017.生物结皮对荒漠草本植物群落结构的影响[J].干旱区研究,34(6):1338-1344.
ZHUANG W W,ZHANG Y M,2017.Effect of soil microbiotic crust on plant community in the Gurbantunggut desert[J].Arid Zone Research,34(6):1338-1344.
BELNAP J,MILLER D M,BEDFORD D R,et al.,2014.Pedological and geological relationships with soil lichen and moss distribution in the eastern Mojave Desert,CA,USA[J].Journal of Arid Environments,106:45-57.
BELNAP J,PRASSE R,HARPER K T,2001.Influence of biological soil crusts on soil environments and vascular plants[M].Berlin:Springer,281-300.
BELNAP J,2002.Nitrogen fixation in biological soil crusts from southeast Utah,USA[J].Biology and Fertility of Soils,35(2):128-135.
BELNAP,JAYNE,2003.Biological soil crusts in deserts:a short review of their role in soil fertility,stabilization,and water relations[J].Algological Studies,109(1):113-126.
BLISS LCGOLD W G,1999.Vascular plant reproduction,establishment,and growth and the effects of cryptogamic crusts within a polar desert ecosystem,Devon Island,N.W.T.Canada[J].Canadian Journal of Botany,77(5):623-636.
BRIGGS A,MORGAN J W,2008.Morphological diversity and abundance of biological soil crusts differ in relation to landscape setting and vegetation type[J].Australian Journal of Botany,56(3):246-253.
COPPOLA A,BASILE A,WANG X,et al.,2011.Hydrological behaviour of microbiotic crusts on sand dunes:Example from NW China comparing infiltration in crusted and crust-removed soil[J].Soil&Tillage Research,117:34-43.
DEINES L,ROSENTRETER R,ELDRIDGE D J,et al.,2007.Germination and seedling establishment of two annual grasses on lichen-dominated biological soil crusts[J].Plant&Soil,295(1-2):23-35.
ELBERT W,ANDREAE M O,2009.Microbiotic crusts on soil,rock and plants:Neglected major players in the global cycles of carbon and nitrogen[J].Biogeo Sciences Discuss,6:6983-7015.
PRASSE R,BORNKAMM R,2000.Effect of microbiotic soil surface crusts on emergence of vascular plants[J].Plant Ecology,150(1-2):65-75.
RIVERAAGUILAR V,GODINEZALVAREZ H,MANUELLCACHEUX I,et al.,2005.Physical effects of biological soil crusts on seed germination of two desert plants under laboratory conditions[J].Journal of Arid Environments,63(1):344-352.
ZHAO Y,XU M,BELNAP J,2010.Potential nitrogen fixation activity of different aged biological soil crusts from rehabilitated grasslands of the hilly Loess Plateau,China[J].Journal of Arid Environments,74(10):1186-1191.
陈兰周,刘永定,李敦海,等,2003.荒漠藻类及其结皮的研究[J].中国科学基金,17(2):90-93.CHEN L Z,LIU Y D,LI J H,et al.,2003.The research process/progress of desert algae and crus[J].Bulletin of National Science Foundation of China,17(2):90-93.
陈孟晨,张景光,冯丽,等,2017.沙坡头地区生物结皮覆盖区土壤种子库组成及垂直分布特征[J].生态学报,37(22):243-252.CHEN M C,ZHANG J G,FENG L,et al.,2017.The composition and vertical distribution characteristics of soil seed banks in soil coverage with biocrusts in the Shapotou Region[J].Acta Ecological Sinica,37(22):243-252.
贺郝钰,侯春梅,迟秀丽,等,2014.国际生物土壤结皮研究发展态势文献计量分析[J].生态学报,34(4):1035-1041.HE H Y,HOU C M,CHI X L,et al.,2014.Bibliometric analysis of the development trend of international biological soil crust research[J].Acta Ecological Sinica,34(4):1035-1041.
李国栋,张元明,2014.生物土壤结皮与种子附属物对4种荒漠植物种子萌发的交互影响[J].中国沙漠,34(3):725-731.LI G D,ZHANG Y M,2014.Interactive Effects of Biological Soil Crusts and Seed Appendages on Seed Germination of Four Desert Species[J].Journal of Desert Research,34(3):725-731.
李新荣,张景光,2000.干旱沙漠区土壤微生物结皮及其对固沙植被影响的研究[J].植物学报:英文版,42(9):965-970.LI X R,ZHANG J G,2000.Study on soil microbiotic crust and its influences on sand-fixing vegetation in arid desert region[J].Acta Botanica Sinica,42(9):965-970.
刘利霞,张宇清,吴斌,2007.生物结皮对荒漠地区土壤及植物的影响研究述评[J].中国水土保持科学,5(6):106-112.LIU L X,ZHANG Y Q,WU B,2007.Review on the influences of biological soil crusts on soil and vegetation in desertified areas[J].Science of Soil and Water Conservation,5(6):106-112.
聂华丽,张元明,吴楠,等,2009.生物结皮对5种不同形态的荒漠植物种子萌发的影响[J].植物生态学报,33(1):161-170.NIE H L,ZHANG Y M,WU N,et al,2009.Effects of biological crusts on the germination of five desert vascular plants with different seed morphologies[J].Chinese Journal of Plant Ecology,33(1):161-170.
宋阳,严平,张宏,等,2004.荒漠生物结皮研究中的几个问题[J].干旱区研究,21(4):439-443.SONG Y,YAN P,ZHANG H,et al.,2004.Recent progress and some existing problems in the study on the desert microbiotic crusts[J].Arid Zone Research,21(4):439-443.
王华磊,贾长虹,李潞滨,2013.土壤结皮及研究进展[J].河北联合大学学报(自然科学版),35(2):107-112.WANG H L,JIA C H,LI L B,2013.Soil Crusts and Research Progress[J].Journal of Hebei United University(Natural Science Edition),35(2):107-112.
李以康,欧阳经政,林丽,等,2015.高寒草甸植被退化过程中生物土壤结皮演变特征[J].生态学杂志,34(8):2238-2244.LI Y K,OUYANG J Z,LIN L,et al.,2015.Evolution characteristics of biological soil crusts(BSCs)during alpine meadow degrada-tion[J].Chinese Journal of Ecology,34(8):2238-2244.
王蕊,朱清科,赵磊磊,等,2011.黄土高原土壤生物结皮对植物种子出苗和生长的影响[J].干旱区研究,28(5):800-807.WANG R,ZHU Q K,ZHAO L L,et al.,2011.Effects of biological soil crusts on seed emergence and seedling growth in Loess Plateau,North Shaanxi province[J].Arid Zone Research,28(5):800-807.
肖波,赵允格,邵明安,2007.陕北水蚀风蚀交错区两种生物结皮对土壤理化性质的影响[J].生态学报,27(11):4662-4670.XIAO B,ZHAO Y G,SHAO M A,2007.Effects of biological soil crust on soil physicochemical properties in water-wind erosion crisscross region,Northern Shaanxi province,China[J].Acta Ecologica Sinica,27(11):4662-4670.
张国秀,赵允格,许明祥,等,2012.黄土丘陵区生物结皮对土壤磷素有效性及碱性磷酸酶活性的影响[J].植物营养与肥料学报,18(3):621-628.ZHANG G X,ZHAO Y G,XUN M X,et al.,2012.Impacts of biological soil crust on availability of phosphorus and phosphatase activity in hilly regions of the Loess Plateau,China[J].Plant Nutrition and Fertilizer Science,18(3):621-628.
张侃侃,卜崇峰,高国雄,2011.黄土高原生物结皮对土壤水分入渗的影响[J].干旱区研究,28(5):808-812.ZHANG K K,PU C F,GAO G X,2011.Effect of microbiotic crust on soil water infiltration in the Loess Plateau[J].Arid Zone Research,28(5):808-812.
赵允格,许明祥,王全九,等,2006.黄土丘陵区退耕地生物结皮对土壤理化性状的影响[J].自然资源学报,21(3):441-448.ZHAO Y G,XU M X,WANG Q J,et al.,2012.Impact of biological soil crust on soil physical and chemical properties of Rebabilitated grassland in Hilly Loess Plateau,China[J].Journal of Natural Resources,21(3):441-448.
赵允格,许明祥,王全九,等,2006.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,17(8):1429-1434.ZHAO Y G,XU M X,WANG Q J,et al.,2006.Physicaland chem ical properties of soilbio-crust on rehabilitated grassland in hilly LoessPlateau of China[J].Chinese Journal of Applied Ecology,17(8):1429-1434.
庄伟伟,张元明,2017.生物结皮对荒漠草本植物群落结构的影响[J].干旱区研究,34(6):1338-1344.
ZHUANG W W,ZHANG Y M,2017.Effect of soil microbiotic crust on plant community in the Gurbantunggut desert[J].Arid Zone Research,34(6):1338-1344.
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