地形对阔叶红松林物种多样性的影响
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摘要
为了更好地了解阔叶红松林(Pinus koraiensis)物种多样性的维持机制,对不同地形下的阔叶红松林乔木和灌木种,进行了物种多样性指数比较、物种多度分布模型拟合及物种多样性指数与环境因子的排序分析。结果表明,乔木在阴坡、半阴坡的丰富度显著高于半阳坡;灌木在坡度>25°处的均匀度显著高于坡度<6°处,在谷地处的4个多样性指数显著低于其他坡位,在阴坡、半阴坡处的4个多样性指数显著高于平地。统计模型、生态位模型、中性模型在不同地形下均能够通过拟合优度检验,物种多度分布在不同地形间差异不明显;地形对乔木的多样性指数的影响:海拔>凹凸度>坡度>坡向>坡位,对灌木的多样性指数的影响:坡位>海拔>坡向>坡度>凹凸度,土壤对乔木的物种多样性指数的影响:土壤pH>土壤容重>土壤有效N含量>土壤速效K含量,对灌木的物种多样性指数的影响:土壤pH>土壤有机质含量>土壤速效P含量>土壤全N含量>土壤全P含量。总体而言,物种多样性随着坡度的增加而增加,阴坡处最高,谷地处最低。
To better understand the mechanism regulating the species diversity of mixed broadleaved-Korean pine( Pinus koraiensis) forest,the species diversity index,goodness-of-fit test of six models for species-abundance distribution,and the significance test between environmental factors and ordination axis of arbor and shrub in the different topographies of mixed broadleaved-Korean pine forest were analyzed. The results showed that the abundance of arbor at the shady and semi-shady slopes is significantly higher than that in the semi-sunny slopes. The shrubs at slope gradient greater than 25 degrees were significantly higher than that at slope gradient less than 6 degrees. The slope position exhibited a significant effect on the four diversity indices of shrub,and valley was significantly lower than those of other slope positions. The four diversity indices of arbor at the shady and semi-shady slopes are significantly higher than those at flats. Statistical models,niche models,and neutral models can be tested by fitting goodness under different topographies,but species distribution in different topographies was not obvious. The effects of topographic factors on species diversity of arbor were in the order:elevation > concavity > slope aspect > slope aspect > slope position. The effects of topographic factors on species diversity of shrub were in the order: slope position > elevation > slope position > slope gradient > concavity. The effects of soil factors on species diversity of arbor were in the order: pH > BD > SHN > SAK. The effects of soil factors on species diversity of shrub were in the order: pH > SOC > SAP > STN > STP. In general,the species diversity increased with the increase in slope,and the species diversity at shady slope was the highest,and that at the valley was the lowest.
To better understand the mechanism regulating the species diversity of mixed broadleaved-Korean pine( Pinus koraiensis) forest,the species diversity index,goodness-of-fit test of six models for species-abundance distribution,and the significance test between environmental factors and ordination axis of arbor and shrub in the different topographies of mixed broadleaved-Korean pine forest were analyzed. The results showed that the abundance of arbor at the shady and semi-shady slopes is significantly higher than that in the semi-sunny slopes. The shrubs at slope gradient greater than 25 degrees were significantly higher than that at slope gradient less than 6 degrees. The slope position exhibited a significant effect on the four diversity indices of shrub,and valley was significantly lower than those of other slope positions. The four diversity indices of arbor at the shady and semi-shady slopes are significantly higher than those at flats. Statistical models,niche models,and neutral models can be tested by fitting goodness under different topographies,but species distribution in different topographies was not obvious. The effects of topographic factors on species diversity of arbor were in the order:elevation > concavity > slope aspect > slope aspect > slope position. The effects of topographic factors on species diversity of shrub were in the order: slope position > elevation > slope position > slope gradient > concavity. The effects of soil factors on species diversity of arbor were in the order: pH > BD > SHN > SAK. The effects of soil factors on species diversity of shrub were in the order: pH > SOC > SAP > STN > STP. In general,the species diversity increased with the increase in slope,and the species diversity at shady slope was the highest,and that at the valley was the lowest.
引文
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[26]赵雪,刘妍妍,金光泽.地形对阔叶红松林幼苗更新的影响.应用生态学报,2013,24(11):3035-3042.
[27]赵雪,徐丽娜,金光泽.地形对典型阔叶红松林灌木更新的影响.生物多样性,2015,23(6):767-774.
[28]徐丽娜,金光泽.小兴安岭凉水典型阔叶红松林动态监测样地:物种组成与群落结构.生物多样性,2012,20(4):470-481.
[29]韩大校,金光泽.地形和竞争对典型阔叶红松林不同生长阶段树木胸径生长的影响.北京林业大学学报,2017,39(1):9-19.
[30]Zhu Y,Cai H Y,Jiang F,Jin G Z.Variation of the biotic neighbourhood and topographic effects on tree survival in an old-growth temperate forest.Journal of Vegetation Science,2017,28(6):1166-1177.
[31]刘妍妍,金光泽.地形对小兴安岭阔叶红松(Pinus koraiensis)林粗木质残体分布的影响.生态学报,2009,29(3):1398-1407.
[32]Shi B K,Gao W F,Cai H Y,Jin G Z.Spatial variation of soil respiration is linked to the forest structure and soil parameters in an old-growth mixed broadleaved-Korean pine forest in northeastern China.Plant and Soil,2016,400(1/2):263-274.
[33]Klute A.Methods of Soil Analysis,Part1.Physical and Mineralogical Methods.2nd ed.Madison:Soil Science Society of America,Inc.,American Society of Agronomy,Inc.,1986:463.
[34]张金屯.数量生态学(第二版).北京:科学出版社,2011:52-75.
[35]Fisher R A,Corbet S,Williams C B.The relation between the number of species and the number of individuals in a random sample of an animal population.Journal of Animal Ecology,1943,12(1):42-58.
[36]Preston F W.The commonness,and rarity,of species.Ecology,1948,29(3):254-283.
[37]周淑荣,张大勇.群落生态学的中性理论.植物生态学报,2006,30(5):868-877.
[38]张姗,蔺菲,原作强,匡旭,贾仕宏,王芸芸,索炎炎,房帅,王绪高,叶吉,郝占庆.长白山阔叶红松林草本层物种多度分布格局及其季节动态.生物多样性,2015,23(5):641-648.
[39]Akaike H.Akaike's information criterion//Lovric M,ed.International Encyclopedia of Statistical Science.Berlin:Springer,2011:104.
[40]Slocomb J,Stauffer B,Dickson K L.On fitting the truncated lognormal distribution to species-abundance data using maximum likelihood estimation.Ecology,1977,58(3):693-696.
[41]Sekhon J S.Multivariate and propensity score matching software with automated balance optimization:the matching package for R.Journal of Statistical Software,2011,42(7):127-210.
[42]Dixon P.VEGAN,a package of R functions for community ecology.Journal of Vegetation Science,2003,14(6):927-930.
[43]May R M.Patterns of species abundance and diversity//Cody M L,Diamond J M.Ecology and Evolution of Communities.Cambridge:Belknap Press of Harvard University,1975:81-120.
[44]Nummelin M.Log-normal distribution of species abundances is not a universal indicator of rain forest disturbance.Journal of Applied Ecology,1998,35(3):454-457.
[45]Lebrija-Trejos E,Pérez-García E A,Meave J A,Bongers F,Poorter L.Functional traits and environmental filtering drive community assembly in a species-rich tropical system.Ecology,2010,91(2):386-398.
[46]OpedalΦH,Armbruster W S,Graae B J.Linking small-scale topography with microclimate,plant species diversity and intra-specific trait variation in an alpine landscape.Plant Ecology&Diversity,2015,8(3):305-315.
[2]Swenson N G,Anglada-Cordero P,Barone J A.Deterministic tropical tree community turnover:evidence from patterns of functional beta diversity along an elevational gradient.Proceedings.Biological Sciences,2011,278(1707):877-884.
[3]Mccool D K,Brown L C,Foster G R,Mutchler C K,Meyer L D.Revised slope steepness factor for the universal soil loss equation.Transactions of the ASAE,1987,30(5):1387-1396.
[4]李俊清.森林生态学(第2版).北京:高等教育出版社,2010:91-96.
[5]Liu J J,Tan Y H,Slik J W F.Topography related habitat associations of tree species traits,composition and diversity in a Chinese tropical forest.Forest Ecology and Management,2014,330(7):75-81.
[6]汪殿蓓,暨淑仪,陈飞鹏.植物群落物种多样性研究综述.生态学杂志,2001,20(4):55-60.
[7]叶万辉.物种多样性与植物群落的维持机制.生物多样性,2000,8(1):17-24.
[8]Mcintosh R P.An index of diversity and the relation of certain concepts to diversity.Ecology,1967,48(3):392-404.
[9]Belcher P R.Measurement of myocardial contractility.Journal of Cardiothoracic and Vascular Anesthesia,1997,11(6):812.
[10]Whittaker R H.Dominance and diversity in land plant communities.Science,1965,147(3655):250-260.
[11]Magurran A E.Ecological Diversity and Its Measurement.Princeton:Princeton University Press,1988:7-45.
[12]Tokeshi M.Niche apportionment or random assortment:species abundance patterns revisted.Journal of Animal Ecology,1990,59(3):1129-1146.
[13]Hughes R G.A model of the structure and dynamics of benthic marine invertebrate communities.Marine Ecology Progress Series,1984,15(1/2):1-11.
[14]Caswell H.Community structure:a neutral model analysis.Ecological Monographs,1976,46(3):327-354.
[15]周红章,于晓东,罗天宏,何君舰.物种多样性变化格局与时空尺度.生物多样性,2000,8(3):325-336.
[16]牛克昌,刘怿宁,沈泽昊,何芳良,方精云.群落构建的中性理论和生态位理论.生物多样性,2009,17(6):579-593.
[17]Young B,Yarie J,Verbyla D,Huettmann F,Herrick K,Chapin F S.Modeling and mapping forest diversity in the boreal forest of interior Alaska.Landscape Ecology,2017,32(2):397-413.
[18]Katovai E,Katovai D D,Edwards W,Laurance W F.Forest structure,plant diversity and local endemism in a highly varied New Guinea landscape.Tropical Conservation Science,2015,8(2):284-300.
[19]Currie D J,Paquin V.Large-scale biogeographical patterns of species richness of trees.Nature,1987,329(6137):326-327.
[20]Lomolino M V.Elevation gradients of species-density:historical and prospective views.Global Ecology and Biogeography,2001,10(1):3-13.
[21]Hosseinzadeh R,Soosani J,Alijani V,Khosravi S,Karimikia H.Diversity of woody plant species and their relationship to physiographic factors in central Zagros forests(Case study:Perc forest,Khorramabad,Iran).Journal of Forestry Research,2016,27(5):1137-1141.
[22]区余端,苏志尧,李镇魁,林义辉.地形因子对粤北山地森林不同生长型地表植物分布格局的影响.应用生态学报,2011,22(5):1107-1113.
[23]黄金燕,周世强,谭迎春,周小平,王鹏彦,张和民.卧龙自然保护区大熊猫栖息地植物群落多样性研究:丰富度、物种多样性指数和均匀度.林业科学,2007,43(3):73-78.
[24]Zhou T,Chen B M,Liu G,Huang F F,Liu J G,Liao W B,Wang Y Y,Ren S J,Chen C Q,Peng S L.Biodiversity of Jinggangshan Mountain:the importance of topography and geographical location in supporting higher biodiversity.PLoS One,2015,10(3):e0120208.
[25]王业蘧.阔叶红松林.哈尔滨:东北林业大学出版社,1995:72-74.
[26]赵雪,刘妍妍,金光泽.地形对阔叶红松林幼苗更新的影响.应用生态学报,2013,24(11):3035-3042.
[27]赵雪,徐丽娜,金光泽.地形对典型阔叶红松林灌木更新的影响.生物多样性,2015,23(6):767-774.
[28]徐丽娜,金光泽.小兴安岭凉水典型阔叶红松林动态监测样地:物种组成与群落结构.生物多样性,2012,20(4):470-481.
[29]韩大校,金光泽.地形和竞争对典型阔叶红松林不同生长阶段树木胸径生长的影响.北京林业大学学报,2017,39(1):9-19.
[30]Zhu Y,Cai H Y,Jiang F,Jin G Z.Variation of the biotic neighbourhood and topographic effects on tree survival in an old-growth temperate forest.Journal of Vegetation Science,2017,28(6):1166-1177.
[31]刘妍妍,金光泽.地形对小兴安岭阔叶红松(Pinus koraiensis)林粗木质残体分布的影响.生态学报,2009,29(3):1398-1407.
[32]Shi B K,Gao W F,Cai H Y,Jin G Z.Spatial variation of soil respiration is linked to the forest structure and soil parameters in an old-growth mixed broadleaved-Korean pine forest in northeastern China.Plant and Soil,2016,400(1/2):263-274.
[33]Klute A.Methods of Soil Analysis,Part1.Physical and Mineralogical Methods.2nd ed.Madison:Soil Science Society of America,Inc.,American Society of Agronomy,Inc.,1986:463.
[34]张金屯.数量生态学(第二版).北京:科学出版社,2011:52-75.
[35]Fisher R A,Corbet S,Williams C B.The relation between the number of species and the number of individuals in a random sample of an animal population.Journal of Animal Ecology,1943,12(1):42-58.
[36]Preston F W.The commonness,and rarity,of species.Ecology,1948,29(3):254-283.
[37]周淑荣,张大勇.群落生态学的中性理论.植物生态学报,2006,30(5):868-877.
[38]张姗,蔺菲,原作强,匡旭,贾仕宏,王芸芸,索炎炎,房帅,王绪高,叶吉,郝占庆.长白山阔叶红松林草本层物种多度分布格局及其季节动态.生物多样性,2015,23(5):641-648.
[39]Akaike H.Akaike's information criterion//Lovric M,ed.International Encyclopedia of Statistical Science.Berlin:Springer,2011:104.
[40]Slocomb J,Stauffer B,Dickson K L.On fitting the truncated lognormal distribution to species-abundance data using maximum likelihood estimation.Ecology,1977,58(3):693-696.
[41]Sekhon J S.Multivariate and propensity score matching software with automated balance optimization:the matching package for R.Journal of Statistical Software,2011,42(7):127-210.
[42]Dixon P.VEGAN,a package of R functions for community ecology.Journal of Vegetation Science,2003,14(6):927-930.
[43]May R M.Patterns of species abundance and diversity//Cody M L,Diamond J M.Ecology and Evolution of Communities.Cambridge:Belknap Press of Harvard University,1975:81-120.
[44]Nummelin M.Log-normal distribution of species abundances is not a universal indicator of rain forest disturbance.Journal of Applied Ecology,1998,35(3):454-457.
[45]Lebrija-Trejos E,Pérez-García E A,Meave J A,Bongers F,Poorter L.Functional traits and environmental filtering drive community assembly in a species-rich tropical system.Ecology,2010,91(2):386-398.
[46]OpedalΦH,Armbruster W S,Graae B J.Linking small-scale topography with microclimate,plant species diversity and intra-specific trait variation in an alpine landscape.Plant Ecology&Diversity,2015,8(3):305-315.