关帝山云杉次生林不同生活型物种与生境相关性
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摘要
以关帝山庞泉沟自然保护区4 hm~2云杉次生林固定监测样地调查数据为基础,采用方差分解(RDA)分析了地形和土壤与不同生活型物种分布的关系,并对两生境因子进行主成分分析(PCA),通过广义可加模型(GAM)拟合了乔木、小乔木、灌木中优势物种分布与主成分分析中各主分量的关系。结果表明:(1)地形和土壤共解释了乔木、小乔木、灌木分布的53.34%、55.65%和45.83%,其中地形因子独立解释了8.36%、5.06%和5.48%,土壤因子独立解释了31.12%、44.22%和32.04%,两因子共同解释的部分为13.86%、6.37%和8.31%;(2)对13个生境指标进行主成分分析表明,前4个主分量分别代表了38.76%、25.73%、19.41%和8.93%,累计贡献率达92.56%;(3)GAM拟合结果显示,不同生活型及同一生活型不同物种分布与生境4个主分量相关程度均不同。模型可解释的偏差分别为:乔木层介于0.76%~29.00%,小乔木层介于0~20.30%,灌木层介于0.10%~23.50%。各物种及各生活型物种普遍与PC1相关程度较大,海拔、坡向、有效K、有效N、有效Mg对物种分布具有重要作用。研究结果表明,地形、土壤等生境因子对不同物种及不同生活型物种分布的影响有差异,这种差异有利于关帝山云杉次生林物种共存和多样性维持。
Based on the survey data of a 4-hm~2 fixed monitoring plot in Pangquangou Nature Reserve on Guandi Mountain,we analyzed correlation between different life-form species distribution and two habitat factors-terrain and soil in a secondary Picea forest using variance decomposition( RDA) and fitted relationships between the dominant species of trees,small trees and shrubs and principal components of habitat factors by generalized additive model( GAM). The results showed that:( 1) Terrain and soil together explained the distribution of trees( 53. 34%),small trees( 55.65%) and shrubs( 45. 83%),including an independent explanation by terrain( 8.36%,5. 06% and 5. 48%,respectively),an independent explanation by soil( 31. 12%,44.22% and 32.04%,respectively),and a joint part explanation( 13.86%,6.37% and 8.31%,respectively);( 2) The principal component analysis of 13 habitat indexes showed that the first four principal components represented 38. 76%,25. 73%,19. 41% and 8. 93%,respectively,with a cumulative contribution rate of 92.83%;( 3) For different species and life-form species,the degree of correlation between their distribution and the four principal components was different by GAM fitting. The deviations explained by the model were 0.76%-29.00% for tree layer,0-20.30% for small tree layer and 0.10%-23.50% for shrub layer. Each species and life form were better associated with PC1,and altitude,aspect,available K,N and Mg played an important role in species distribution. The result indicated that the effects of habitat factors such as terrain and soil on the different life-form species distribution were different,which promoted the coexistence and biodiversity maintaining of species in the secondary Picea forest on Guandi Mountain.
Based on the survey data of a 4-hm~2 fixed monitoring plot in Pangquangou Nature Reserve on Guandi Mountain,we analyzed correlation between different life-form species distribution and two habitat factors-terrain and soil in a secondary Picea forest using variance decomposition( RDA) and fitted relationships between the dominant species of trees,small trees and shrubs and principal components of habitat factors by generalized additive model( GAM). The results showed that:( 1) Terrain and soil together explained the distribution of trees( 53. 34%),small trees( 55.65%) and shrubs( 45. 83%),including an independent explanation by terrain( 8.36%,5. 06% and 5. 48%,respectively),an independent explanation by soil( 31. 12%,44.22% and 32.04%,respectively),and a joint part explanation( 13.86%,6.37% and 8.31%,respectively);( 2) The principal component analysis of 13 habitat indexes showed that the first four principal components represented 38. 76%,25. 73%,19. 41% and 8. 93%,respectively,with a cumulative contribution rate of 92.83%;( 3) For different species and life-form species,the degree of correlation between their distribution and the four principal components was different by GAM fitting. The deviations explained by the model were 0.76%-29.00% for tree layer,0-20.30% for small tree layer and 0.10%-23.50% for shrub layer. Each species and life form were better associated with PC1,and altitude,aspect,available K,N and Mg played an important role in species distribution. The result indicated that the effects of habitat factors such as terrain and soil on the different life-form species distribution were different,which promoted the coexistence and biodiversity maintaining of species in the secondary Picea forest on Guandi Mountain.
引文
鲍士旦.2005.土壤农化分析.第3版.北京:中国农业出版社.
陈云,袁志良,任思远,等.2014.宝天曼自然保护区不同生活型物种与土壤相关性分析.科学通报,59(24):2367-2376.
刘海丰,桑卫国,薛达元.2013.暖温带森林优势种群的地形生境变异性.生态学杂志,32(4):795-801.
王盛萍,张志强,张建军,等.2010.黄土残塬沟壑区流域次生植被物种分布的地形响应.生态学报,30(22):6102-6112.
王伟,骆争荣,周荣飞,等.2011.百山祖常绿阔叶林木本植物的生境相关性分析.生物多样性,19(2):134-142.
吴昊.2015.秦岭山地松栎混交林土壤养分空间变异及其与地形因子的关系.自然资源学报,30(5):858-869.
谢玉彬,马遵平,杨庆松,等.2012.基于地形因子的天童地区常绿树种和落叶树种共存机制研究.生物多样性,20(2):159-167.
闫海冰,韩有志,杨秀清,等.2010.关帝山云杉天然更新与土壤有效氮素异质性的空间关联性.应用生态学报,21(3):533-540.
杨秀清,韩有志,杨秀清,等.2009.华北山地典型天然次生林土壤氮素空间异质性对落叶松幼苗更新的影响.生态学报,29(9):4656-4664.
袁志良,陈云,韦博良,等.2013.暖温带-北亚热带生态过渡区物种生境相关性分析.生态学报,33(24):7819-7826.
张俪文.2010.环境空间异质性对物种空间分布和群落结构的影响(博士学位论文).北京:中国科学院植物研究所.
张娜,王希华,郑泽梅,等.2012.浙江天童常绿阔叶林土壤的空间异质性及其与地形的关系.应用生态学报,23(9):2361-2369.
Borcard D,Legendre P,Drapeau P.1992.Partialling out the spatial component of ecological variation.Ecology,73:1045-1055.
Chang LW,ZelenD,Li CF,et al.2013.Better environmental data may reverse conclusions about niche-and dispersalbased processes in community assembly.Ecology,94:2145-2151.
Condit R.1998.Tropical forest census plots:Methods and results from Barro Colorado Island,Panama and a comparison with other plots[EB/OL].[2017-2-10].http://www.springer.com.
Debski I,Burslem DFRP,Palmiotto PA,et al.2002.Habitat preferences of Aporosa in two Malaysian forests:Implications for abundance and coexistence.Ecology,83:2005-2018.
Gunatilleke CVS,Gunatilleke IAUN,Esufali S,et al.2006.Species-habitat associations in a Sri Lankan dipterocarp forest.Journal of Tropical Ecology,22:371-384.
Harms KE,Condit R,Hubbell SP,et al.2001.Habitat associations of trees and shrubs in a 50-ha neotropical forest plot.Journal of Ecology,89:947-959.
Hubbell SP,Foster RB,O'Brien ST,et al.1999.Light-gap disturbances,recruitment limitation,and tree diversity in a neotropical forest.Science,283:554-557.
Hubbell SP,Foster RB.1983.Diversity of canopy trees in a neotropical forest and implications for conservation//Sutton SL,Whitmore TC,Chadwick AC,eds.Tropical Rain Forest:Ecology and Management.Oxford:Blackwell Scientific Publications:25-41.
Hubbell SP,Foster RB.1986.Biology,Chance and History and the Structure of Tropical Rain Forest Tree Communities//Diamond JM and Case TJ,Eds.Community Ecology.Harper&Row,New York:314-329.
Itoh A,Yamakura T,Ohkubo T,et al.2003.Importance of topography and soil texture in the spatial distribution of two sympatric dipterocarp trees in a Bornean rain forest.Ecological Research,18:307-320.
John R,Dalling JW,Harms KE,et al.2007.Soil nutrients influence spatial distributions of tropical tree species.Proceedings of the National Academy of Science of the United States of America,104:864-869.
Kikuchi T,Miura O.1993.Vegetation patterns in relation to micro-scale landforms in hilly land regions.Vegetatio,106:147-154.
Lai J,Mi X,Ren H,et al.2009.Species-habitat associations change in a subtropical forest of China.Journal of Vegetation Science,20:415-423.
Nakashizuka T.2001.Species coexistence in temperate,mixed deciduous forests.Trends in Ecology&Evolution,16:205-210.
Nee S.2005.The neutral theory of biodiversity:Do the numbers add up?Functional Ecology,19:173-176.
Oksanen J,Blanchet FG,Kindt R,et al.2010.Vegan:Community Ecology Package.R package version 1.
Potts MD,Ashton PS,Kaufman LS,et al.2002.Habitat patterns in tropical rain forests:A comparison of 105 plots in northwest Borneo.Ecology,83:2782-2797.
Sakai A,Ohsawa M.1994.Topographical pattern of the forest vegetation on a river basin in a warm-temperate hilly region,central Japan.Ecological Research,9:269-280.
Silvertown JW,Charlesworth D.2001.Introduction to Plant Population Biology.Oxford:Blackwell Science:201-223.
Smith TW,Lundholm JT.2010.Variation partitioning as a tool to distinguish between niche and neutral processes.Ecography,33:648-655.
Svenning JC.1999.Microhabitat specialization in a species-rich palm community in Amazonian Ecuador.Journal of Ecology,87:55-65.
Tateno R,Hishi T,Takeda H.2004.Above-and belowground biomass and net primary production in a cool-temperate deciduous forest in relation to topographical changes in soil nitrogen.Forest Ecology and Management,193:297-306.
Tilman D,Pacala S.1993.The maintenance of species richness in plant communities//Ricklefs RE,Schluter D,eds.Species Diversity in Ecological Communities.Chicago:University of Chicago Press:13-25.
Trevor H.2013.Package“gam”[EB/OL].[2017-2-10].http://web.stanford.edu/~hastie/sw Data.htm.
Valencia R,Foster RB,Villa G,et al.2004.Tree species distributions and local habitat variation in the Amazon:Alarge plot in eastern Ecuador.Journal of Ecology,92:214-229.
Yamakura T,Kanzake M,Itoh A,et al.1995.Topography of a large-scale research plot established within a tropical rain forest at Lambir,Sarawak.Tropics,5:41-56.
Zhang L,Mi X,Shao H,et al.2011.Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest.Plant and Soil,347:211-220.
陈云,袁志良,任思远,等.2014.宝天曼自然保护区不同生活型物种与土壤相关性分析.科学通报,59(24):2367-2376.
刘海丰,桑卫国,薛达元.2013.暖温带森林优势种群的地形生境变异性.生态学杂志,32(4):795-801.
王盛萍,张志强,张建军,等.2010.黄土残塬沟壑区流域次生植被物种分布的地形响应.生态学报,30(22):6102-6112.
王伟,骆争荣,周荣飞,等.2011.百山祖常绿阔叶林木本植物的生境相关性分析.生物多样性,19(2):134-142.
吴昊.2015.秦岭山地松栎混交林土壤养分空间变异及其与地形因子的关系.自然资源学报,30(5):858-869.
谢玉彬,马遵平,杨庆松,等.2012.基于地形因子的天童地区常绿树种和落叶树种共存机制研究.生物多样性,20(2):159-167.
闫海冰,韩有志,杨秀清,等.2010.关帝山云杉天然更新与土壤有效氮素异质性的空间关联性.应用生态学报,21(3):533-540.
杨秀清,韩有志,杨秀清,等.2009.华北山地典型天然次生林土壤氮素空间异质性对落叶松幼苗更新的影响.生态学报,29(9):4656-4664.
袁志良,陈云,韦博良,等.2013.暖温带-北亚热带生态过渡区物种生境相关性分析.生态学报,33(24):7819-7826.
张俪文.2010.环境空间异质性对物种空间分布和群落结构的影响(博士学位论文).北京:中国科学院植物研究所.
张娜,王希华,郑泽梅,等.2012.浙江天童常绿阔叶林土壤的空间异质性及其与地形的关系.应用生态学报,23(9):2361-2369.
Borcard D,Legendre P,Drapeau P.1992.Partialling out the spatial component of ecological variation.Ecology,73:1045-1055.
Chang LW,ZelenD,Li CF,et al.2013.Better environmental data may reverse conclusions about niche-and dispersalbased processes in community assembly.Ecology,94:2145-2151.
Condit R.1998.Tropical forest census plots:Methods and results from Barro Colorado Island,Panama and a comparison with other plots[EB/OL].[2017-2-10].http://www.springer.com.
Debski I,Burslem DFRP,Palmiotto PA,et al.2002.Habitat preferences of Aporosa in two Malaysian forests:Implications for abundance and coexistence.Ecology,83:2005-2018.
Gunatilleke CVS,Gunatilleke IAUN,Esufali S,et al.2006.Species-habitat associations in a Sri Lankan dipterocarp forest.Journal of Tropical Ecology,22:371-384.
Harms KE,Condit R,Hubbell SP,et al.2001.Habitat associations of trees and shrubs in a 50-ha neotropical forest plot.Journal of Ecology,89:947-959.
Hubbell SP,Foster RB,O'Brien ST,et al.1999.Light-gap disturbances,recruitment limitation,and tree diversity in a neotropical forest.Science,283:554-557.
Hubbell SP,Foster RB.1983.Diversity of canopy trees in a neotropical forest and implications for conservation//Sutton SL,Whitmore TC,Chadwick AC,eds.Tropical Rain Forest:Ecology and Management.Oxford:Blackwell Scientific Publications:25-41.
Hubbell SP,Foster RB.1986.Biology,Chance and History and the Structure of Tropical Rain Forest Tree Communities//Diamond JM and Case TJ,Eds.Community Ecology.Harper&Row,New York:314-329.
Itoh A,Yamakura T,Ohkubo T,et al.2003.Importance of topography and soil texture in the spatial distribution of two sympatric dipterocarp trees in a Bornean rain forest.Ecological Research,18:307-320.
John R,Dalling JW,Harms KE,et al.2007.Soil nutrients influence spatial distributions of tropical tree species.Proceedings of the National Academy of Science of the United States of America,104:864-869.
Kikuchi T,Miura O.1993.Vegetation patterns in relation to micro-scale landforms in hilly land regions.Vegetatio,106:147-154.
Lai J,Mi X,Ren H,et al.2009.Species-habitat associations change in a subtropical forest of China.Journal of Vegetation Science,20:415-423.
Nakashizuka T.2001.Species coexistence in temperate,mixed deciduous forests.Trends in Ecology&Evolution,16:205-210.
Nee S.2005.The neutral theory of biodiversity:Do the numbers add up?Functional Ecology,19:173-176.
Oksanen J,Blanchet FG,Kindt R,et al.2010.Vegan:Community Ecology Package.R package version 1.
Potts MD,Ashton PS,Kaufman LS,et al.2002.Habitat patterns in tropical rain forests:A comparison of 105 plots in northwest Borneo.Ecology,83:2782-2797.
Sakai A,Ohsawa M.1994.Topographical pattern of the forest vegetation on a river basin in a warm-temperate hilly region,central Japan.Ecological Research,9:269-280.
Silvertown JW,Charlesworth D.2001.Introduction to Plant Population Biology.Oxford:Blackwell Science:201-223.
Smith TW,Lundholm JT.2010.Variation partitioning as a tool to distinguish between niche and neutral processes.Ecography,33:648-655.
Svenning JC.1999.Microhabitat specialization in a species-rich palm community in Amazonian Ecuador.Journal of Ecology,87:55-65.
Tateno R,Hishi T,Takeda H.2004.Above-and belowground biomass and net primary production in a cool-temperate deciduous forest in relation to topographical changes in soil nitrogen.Forest Ecology and Management,193:297-306.
Tilman D,Pacala S.1993.The maintenance of species richness in plant communities//Ricklefs RE,Schluter D,eds.Species Diversity in Ecological Communities.Chicago:University of Chicago Press:13-25.
Trevor H.2013.Package“gam”[EB/OL].[2017-2-10].http://web.stanford.edu/~hastie/sw Data.htm.
Valencia R,Foster RB,Villa G,et al.2004.Tree species distributions and local habitat variation in the Amazon:Alarge plot in eastern Ecuador.Journal of Ecology,92:214-229.
Yamakura T,Kanzake M,Itoh A,et al.1995.Topography of a large-scale research plot established within a tropical rain forest at Lambir,Sarawak.Tropics,5:41-56.
Zhang L,Mi X,Shao H,et al.2011.Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest.Plant and Soil,347:211-220.