内蒙古典型草原群落内部植物和土壤空间异质性
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摘要
植物群落空间异质性及其动态是认识生态系统生物多样性和稳定性维持机制的基础,而目前对蒙古高原典型草原植物群落空间异质性及其维持机制尚没有专门的研究。本研究在内蒙古温带典型草原区选定针茅(Stipa grandis)和羊草(Leymus chinensis)为优势种的典型草原群落,测定了216个样点(在面积为13.5 hm~2的草地上规则排列12行×18列,样点间距25 m)的植物群落和土壤性质,探讨植物群落的斑块结构及其与土壤性质的关系。结果表明,草原群落具有明显的斑块结构,不同斑块间植物组成的差异促进了群落的生物多样性;草原群落不同斑块之间没有观测到明显的微地形差异且土壤含水量差异不显著(P> 0.05),但土壤容重和养分与不同斑块间植物养分含量的变化具有对应性;草原放牧会降低土壤的养分,增大土壤紧实度和容重,可以推论草原植物群落斑块尺度上的空间异质性可能与放牧影响的空间异质性相关;多年生丛生禾草针茅为优势的群落斑块具有聚集养分的作用,土壤有机碳和氮、磷元素相对丰富,而猪毛菜(Salsola collina)为优势的群落土壤养分低,限制了原生草原优势植物的生长及其对群落伴生植物的抑制,为一年生植物夏雨后快速生长提供了条件。本研究加深了对草原空间异质性的认识,有益于草原生物多样性保护和可持续管理。
Spatial heterogeneity in a grassland ecosystem is associated with grassland species diversity and production stability. Understanding the drivers and mechanisms of grassland spatial heterogeneity is essential for the development of best practices in grassland management. To our knowledge, patch-scale spatial heterogeneity of the grassland communities on the steppes on the Mongolian Plateau has not been investigated to date. Here, we investigated the spatial variation of plant communities and soil properties in 216 sites in a rectangular area of 13.5 hm~2 in a typical steppe dominated by Stipa grandis and Leymus chinensis in Inner Mongolia. Our results showed that plant community patches at these sites could be clustered into four groups, dominated by S. grandis, L. chinensis, Salsola collina, or jointly by S. grandis, L. chinensis, and Cleistogenes squarrosa(mixed patch). The differences in the species compositions among these patches indicated an enhancement of species diversity due to spatial heterogeneity. Soil bulk density, organic carbon(SOC), and nutrients(total nitrogen, TN; total phosphorous, TP) and pH differed among the 4 patch types(P < 0.05), but soil moisture did not(P >0.05), which suggests that soil nutrients and bulk density are associated with the spatial heterogeneity of plant communities.The differences in soil bulk density and nutrients among these patch types are most likely related to the heterogeneity of grazing impacts at the patch scale. The grassland patches dominated by perennial S. grandis had higher SOC, TN, and TP,and lower soil bulk density than those of patches dominated by annual S. collina. The low nutrient content of the S. collina patches suggests that the growth of dominant species, S. grandis and L. chinensis, was limited on these patches, which provided a favorable condition for the fast growth of annuals such as S. collina using the pulse of water and nutrients following the summer rainfall. Our research provides new insights into the spatial heterogeneity of steppe grasslands, and is helpful for conservation and sustainable management of the steppes.
Spatial heterogeneity in a grassland ecosystem is associated with grassland species diversity and production stability. Understanding the drivers and mechanisms of grassland spatial heterogeneity is essential for the development of best practices in grassland management. To our knowledge, patch-scale spatial heterogeneity of the grassland communities on the steppes on the Mongolian Plateau has not been investigated to date. Here, we investigated the spatial variation of plant communities and soil properties in 216 sites in a rectangular area of 13.5 hm~2 in a typical steppe dominated by Stipa grandis and Leymus chinensis in Inner Mongolia. Our results showed that plant community patches at these sites could be clustered into four groups, dominated by S. grandis, L. chinensis, Salsola collina, or jointly by S. grandis, L. chinensis, and Cleistogenes squarrosa(mixed patch). The differences in the species compositions among these patches indicated an enhancement of species diversity due to spatial heterogeneity. Soil bulk density, organic carbon(SOC), and nutrients(total nitrogen, TN; total phosphorous, TP) and pH differed among the 4 patch types(P < 0.05), but soil moisture did not(P >0.05), which suggests that soil nutrients and bulk density are associated with the spatial heterogeneity of plant communities.The differences in soil bulk density and nutrients among these patch types are most likely related to the heterogeneity of grazing impacts at the patch scale. The grassland patches dominated by perennial S. grandis had higher SOC, TN, and TP,and lower soil bulk density than those of patches dominated by annual S. collina. The low nutrient content of the S. collina patches suggests that the growth of dominant species, S. grandis and L. chinensis, was limited on these patches, which provided a favorable condition for the fast growth of annuals such as S. collina using the pulse of water and nutrients following the summer rainfall. Our research provides new insights into the spatial heterogeneity of steppe grasslands, and is helpful for conservation and sustainable management of the steppes.
引文
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[21]任京辰,张平究,潘根兴,宋林华.岩溶土壤的生态地球化学特征及其指示意义:以贵州贞丰-关岭岩溶石山地区为例.地球科学进展,2006,21(5):504-512.REN J C,ZHANG P J,PAN G X,SONG L H.Indices of eco-geochemical characteristics in a degradation-reclamation sequence of soils in mountainous Karst area:A case study in Guanling-Zhenfeng region,Guizhou,China.Advances in Earth Science,2006,21(5):504-512.
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[2]KOERNER S E,COLLINS S L.Small-scale patch structure in north American and south African grasslands responds differently to fire and grazing.Landscape Ecology,2013,28:1293-1306.
[3]LACA E A,SOKOLOW S,GALLI J R,CANGIANO C A.Allometry and spatial scales of foraging in mammalian herbivores.Ecology Letters,2010,13(3):311-20.
[4]张卫国,黄文冰,杨振宇.草地微斑块与草地退化关系的研究.草业学报,2003,12(3):44-50.ZHANG W G,HUANG W B,YANG Z Y.The study on the relationship between mini-patch and degradation of pasture.Acta Pratacultural Science,2003,12(3):44-50.
[5]王岭.大型草食动物采食对植物多样性与空间格局的响应及行为适应机制.长春:东北师范大学博士学位论文,2010.WANG L.Behavioral adaptive mechanisms of large herbivore foraging in response to diversity and spatial pattern of plants.PhDThesis.Changchun:Northeast Normal University,2010.
[6]陈俊华,刘兴良,何飞,刘世荣.卧龙巴朗山川滇高山栎灌丛主要木本植物种群生态位特征.林业科学,2010,46(3):23-28.CHEN J H,LIU X L,HE F,LIU S R.Niche characteristics of dominant woody populations in Quercus aquifoliodes shrub community in Balangshan Mountain in Wolong Nature Reserve.Scientia Silvae Sinicae,2010,46(3):23-28.
[7]孙磊,刘玉,武高林,魏学红.藏北退化草地群落生物量与土壤养分的关系.草业科学,2016,33(6):1062-1069.SUN L,LIU Y,WU G L,WEI X H.The relationships between community biomass and soil nutrients in the northern Tibet degradation grassland.Pratacultural Science,2016,33(6):1062-1069.
[8]JANSSENS F,PEETERS A,TALLOWIN J R B,BAKKER J P,BEKKER R M,FILLAT F,OOMES M J M.Relationship between soil chemical factors and grassland diversity.Plant and Soil,1998,202(1):69-78.
[9]胡忠良,潘根兴,李恋卿,杜有新,王新洲.贵州喀斯特山区不同植被下土壤C、N、P含量和空间异质性.生态学报,2009,29(8):4187-4195.HU Z L,PAN G X,LI L Q,DU Y X,WANG X Z.Changes in pools and heterogeneity of soil organic carbon,nitrogen and phosphorus under different vegetation types in Karst mountainous area of central Guizhou Province.Acta Ecologica Sinica,2009,29(8):4187-4195.
[10]CHEN J,STARK J M.Plant species effects and carbon and nitrogen cycling in a sagebrush crested wheatgrass soil.Soil Biology&Biochemistry,2000,32(1):47-57.
[11]李丹,康萨如拉,赵梦颖,张庆,任海娟,任婧,周俊梅,王珍,吴仁吉.内蒙古羊草草原不同退化阶段土壤养分与植物功能性状的关系.植物生态学报,2016,40(10):991-1002.LI D,Kangsarula,ZHAO M Y,ZHANG Q,REN H J,REN J,ZHOU J M,WANG Z,WU R J.Relationships between soil nutrients and plant functional traits in different degradation stages of Leymus chinensis steppe in Inner Mogolia,China.Acta Phytoecologica Sinica,2016,40(10):991-1002.
[12]PUTTEN W H V D,BARDGETT R D,BEVER J D,BEZEMER T M,CASPER B B,FUKAMI T,KARDOL P,KLIRONOMOS JN,KULMATISKI A,SCHWEITZER J A,SUDING K N,VOORDE T F J V D,WARDLE D A.Plant-soil feedbacks:The past,the present and future challenges.Journal of Ecology,2013,101:265-276.
[13]许岳飞,益西措姆,付娟娟,陈浩,苗彦军,陈俊,呼天明,镡建国.青藏高原高山嵩草草甸植物多样性和土壤养分对放牧的响应机制.草地学报,2012,20(6):1026-1032.XU Y F,Yixicuomu,FU J J,CHEN H,MIAO Y J,CHEN J,HU T M,TAN J G.Response of plant diversity and soil nutrient to grazing intensity in Kobresia pygmaea meadow of Qinghai-Tibet Plateau.Acta Agrectir Sinica,2012,20(6):1026-1032.
[14]刘振国,李镇清.植物群落中物种小尺度空间结构研究.植物生态学报,2005,29(6):1020-1028.LIU Z G,LI Z Q.Perspectives on small-scale spatial structure of plant species in plant communities.Acta Phytoecologica Sinica,2005,29(6):1020-1028.
[15]郭正刚,王根绪,沈禹颖,程国栋.青藏高原北部多年冻土区草地植物多样性.生态学报,2004,24(1):149-155.GUO Z G,WANG G X,SHEN Y Y,CHENG G D.Plant species diversity of grassland plant communities in permafrost regions of the northern Qinghai-Tibet Plateau.Acta Ecologica Sinica,2004,24(1):149-155.
[16]杨晓霞,任飞,周华坤,贺金生.青藏高原高寒草甸植物群落生物量对氮、磷添加的响应.植物生态学报,2014,38(2):159-166.YANG X X,REN F,ZHOU H K,HE J S.Responses of plant community biomass to nitrogen and phosphorus additions in an alpine meadow on the Qinghai-Xizang Plateau.Chinese Journal of Plant Ecology,2014,38(2):159-166.
[17]ZHANG X Z,SHEN Z X,FU G.A meta-analysis of the effects of experimental warming on soil carbon and nitrogen dynamics on the Tibetan Plateau.Applied Soil Ecology,2015,87(87):32-38.
[18]KOTLIAR N B,WIENS J A.Multiple scales of patchiness and patch structure:A hierarchical framework for the study of heterogeneity.Oikos,1990,59:253-260.
[19]MURPHY J,RILEY J P.A modified single solution method for the determination of phosphate in natural waters.Analytica Chimica Acta,1962,27:31-36.
[20]李芳.K-Means算法的k值自适应优化方法研究.合肥:安徽大学硕士学位论文,2015.LI F.Research of the adaptive optimization method about k value of K-Means algorithm.Master Thesis.Hefei:Anhui University,2015.
[21]任京辰,张平究,潘根兴,宋林华.岩溶土壤的生态地球化学特征及其指示意义:以贵州贞丰-关岭岩溶石山地区为例.地球科学进展,2006,21(5):504-512.REN J C,ZHANG P J,PAN G X,SONG L H.Indices of eco-geochemical characteristics in a degradation-reclamation sequence of soils in mountainous Karst area:A case study in Guanling-Zhenfeng region,Guizhou,China.Advances in Earth Science,2006,21(5):504-512.
[22]SEARLE K R,HOBBS N T,SHIPLEY L A.Should I stay or should I go?Patch departure decisions by herbivores at multiple scales.Oikos,2005,111:417-424.
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