功能多样性对典型阔叶红松林生产力的影响
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摘要
为比较生物量比率假说与生态位互补假说在解释生产力变异的相对重要性,探讨生物多样性和生产力之间的关系是否受到生物和非生物因素的影响,该研究依托小兴安岭9 hm~2阔叶红松(Pinus koraiensis)林动态监测样地,通过计算群落初始生物量、物种多样性、功能多样性、植物性状的群落加权平均值和测定环境因子,运用线性回归模型、结构方程模型,比较了物种多样性和功能多样性与生产力的相关性。结果表明:(1)物种多样性和功能多样性均对生产力有显著作用,功能多样性比物种多样性与生产力的关系更为密切;(2)功能多样性指数比群落加权平均值能更好地解释生产力变异,说明生态位互补假说更适用于解释阔叶红松林群落内生产力的变异;(3)生物多样性与生产力的关系受生物因素与非生物因素的共同作用,相较于多样性和功能性状组成(植被质量),初始林分生物量(植被数量)能更有效地解释生产力的变异。生物多样性与生产力关系的研究应从植被质量与植被数量同时出发,评估生态系统过程的多种非生物和生物驱动因素,同时维护森林功能多样性,加强植物与土壤环境的保护,对有效增加生产力和维持生物多样性具有重要意义。
Aims Exploring the relationship between biodiversity and ecosystem productivity has become a hot topic in ecological research. The results support the mass ratio hypothesis and niche complementarity hypothesis, but their relative importance is still controversial. Our aim is to test the relative importance of these two hypotheses in explaining the variability of productivity, and to explore whether the relationship between biodiversity and productivity is influenced by both biotic and abiotic factors.Methods We used the data of woody plants in a 9 hm2 typical mixed broadleaved-Korean pine forest. By calculating the initial biomass, species diversity, functional diversity, community-weighted mean functional traits and measuring environmental factors, we analyzed the relationship between diversity and productivity by the linear regression model and structural equation model.Important findings The results showed that:(1) Both species diversity and functional diversity played a significant role in productivity, and functional diversity was more closely related to productivity than species diversity;(2) the functional diversity index could better explain the variation of productivity than community-weighted mean functional traits. It suggested that the niche complementarity hypothesis was more suitable for explaining the variation of productivity in the mixed broadleaved-Korean pine forest;(3) the relationship between biodiversity and productivity was affected by biotic and abiotic factors, and compared with diversity and functional character composition(vegetation quality), initial stand biomass(vegetation quantity) could explain the variation of productivity more effectively. Our study suggests that, it is important to maintain forest functional diversity and strengthen the protection of plants and soil environments to increase productivity and biodiversity effectively.
Aims Exploring the relationship between biodiversity and ecosystem productivity has become a hot topic in ecological research. The results support the mass ratio hypothesis and niche complementarity hypothesis, but their relative importance is still controversial. Our aim is to test the relative importance of these two hypotheses in explaining the variability of productivity, and to explore whether the relationship between biodiversity and productivity is influenced by both biotic and abiotic factors.Methods We used the data of woody plants in a 9 hm2 typical mixed broadleaved-Korean pine forest. By calculating the initial biomass, species diversity, functional diversity, community-weighted mean functional traits and measuring environmental factors, we analyzed the relationship between diversity and productivity by the linear regression model and structural equation model.Important findings The results showed that:(1) Both species diversity and functional diversity played a significant role in productivity, and functional diversity was more closely related to productivity than species diversity;(2) the functional diversity index could better explain the variation of productivity than community-weighted mean functional traits. It suggested that the niche complementarity hypothesis was more suitable for explaining the variation of productivity in the mixed broadleaved-Korean pine forest;(3) the relationship between biodiversity and productivity was affected by biotic and abiotic factors, and compared with diversity and functional character composition(vegetation quality), initial stand biomass(vegetation quantity) could explain the variation of productivity more effectively. Our study suggests that, it is important to maintain forest functional diversity and strengthen the protection of plants and soil environments to increase productivity and biodiversity effectively.
引文
Ali A,Yan ER,Chang SX,Cheng JY,Liu XY(2017).Community-weighted mean of leaf traits and divergence of wood traits predict aboveground biomass in secondary subtropical forests.Science of the Total Environment,574,654-662.
BartońK(2018).Package‘MuMIn’.https://cran.r-project.org/web/packages/MuMIn/MuMIn.pdf.Cited:2018-07-21.
Cardinale BJ,Duffy JE,Gonzalez A,Hooper DU,Perrings C,Venail P(2012).Biodiversity loss and its impact on humanity.Nature,489,59-67.
Cardinale BJ,Gonzalez A(2011).The functional role of producer diversity in ecosystems.American Journal of Botany,98,572-592.
Chang LW,Zeleny D,Li CF,Chiu ST,Hsieh CF(2013).Better environmental data may reverse conclusions about nicheand dispersal-based processes in community assembly.Ecology,94,2145-2151.
Chave J,Condit R,Lao S,Caspersen JP,Foster RB,Hubbell SP(2003).Spatial and temporal variation of biomass in a tropical forest:Results from a large census plot in Panama.Journal of Ecology,91,240-252.
Chen CG,Zhu JF(1989).A Handbook for Main Tree Species Biomass in Northeast China.China Forestry Publishing House,Beijing.[陈传国,朱俊凤(1989).东北主要林木生物量手册.中国林业出版社,北京.]
Chen DM,Cheng JH,Chu PF,Mi J,Hu SJ,Xie YC,Tuvshintogtokh I,Bai YF(2016).Effect of diversity on biomass across grasslands on the Mongolian Plateau:Contrasting effects between plants and soil nematodes.Journal of Biogeography,43,955-966.
Cornelissen JHC,Lavorel S,Garnier E,Diaz S,Buchmann N,Gurvich DE,Reich PB,ter Steege H,Morgan HD,van der Heijden MGA,Pausas JG,Poorter H(2003).A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,51,335-380.
Díaz S,Lavorel S,Bello FD,Quétier F,Grigulis K,Robson TM(2007).Incorporating plant functional diversity effects in ecosystem service assessments.Proceedings of the National Academy of Sciences of the United States of America,104,20684-20689.
D?nescu A,Albrecht AT,Bauhus J(2016).Structural diversity promotes productivity of mixed,uneven-aged forests in southwestern Germany.Oecologia,182,319-333.
Edwards JR,Lambert LS(2007).Methods for integrating moderation and mediation:A general analytical framework using moderated path analysis.Psychological Methods,12,1-22.
Finegan B,Pe?a-Claros M,de Oliveira A,Ascarrunz N,Bret-Harte MS,Carre?o-Rocabado G,Casanoves F,Díaz S,Eguiguren Velepucha P,Fernandez F,Licona JC,Lorenzo L,Salgado Negret B,Vaz M,Poorter L,Canham C(2015).Does functional trait diversity predict above-ground biomass and productivity of tropical forests?Testing three alternative hypotheses.Journal of Ecology,103,191-201.
Fox J(2008).Applied Regression Analysis and Generalized Linear Models.2nd edn.Sage Publications,Thousand Oaks,USA.
Fox J,Monette G(1992).Generalized collinearity diagnostics.Journal of the American Statistical Association,87,178-183.
Gazol A,Camarero JJ(2016).Functional diversity enhances silver fir growth resilience to an extreme drought.Journal of Ecology,104,1063-1075.
Grace JB,Anderson TM,Seabloom EW,Borer ET,Adler PB,Harpole WS,Hautier Y,Hillebrand H,Lind EM,Partel M,Bakker JD,Buckley YM,Crawley MJ,Damschen EI,Davies KF,Fay PA,Firn J,Gruner DS,Hector A,Knops JM,MacDougall AS,Melbourne BA,Morgan JW,Orrock JL,Prober SM,Smith MD(2016).Integrative modelling reveals mechanisms linking productivity and plant species richness.Nature,529,390-393.
Grime JP(1998).Benefits of plant diversity to ecosystems:Immediate,filter and founder effects.Journal of Ecology,86,902-910.
Howard AL(2013).Handbook of structural equation modeling.Structural Equation Modeling-a Multidisciplinary Journal,20,354-360.
Huang XR(2018).Relationship between plant functional diversity and productivity of Pinus massoniana plantations in Guangxi.Biodiversity Science,26,690-700.[黄小荣(2018).广西马尾松林植物功能多样性与生产力的关系.生物多样性,26,690-700.]
LalibertéE,Legendre P(2010).A distance-based framework for measuring functional diversity from multiple traits.Ecology,91,299-305.
Lin DM,Lai JS,Muller-Landau HC,Mi XC,Ma KP(2012).Topographic variation in aboveground biomass in a subtropical evergreen broad-leaved forest in China.PLOSONE,DOI:10.1371/journal.pone.0048244.
Lohbeck M,Poorter L,Martinez-Ramos M,Bongers F(2015).Biomass is the main driver of changes in ecosystem process rates during tropical forest succession.Ecology,96,1242-1252.
LüTT,Wang P,Yan H,Zhang W,Liao GX,Jiang HB,Zou CL,Sheng LX(2014).Relationship between functional diversity and productivity in meadow and marsh plant communities.Chinese Journal of Plant Ecology,38,405-416.[吕亭亭,王平,燕红,张稳,廖桂项,姜海波,邹畅林,盛连喜(2014).草甸和沼泽植物群落功能多样性与生产力的关系.植物生态学报,38,405-416.]
McGill BJ,Enquist BJ,Weiher E,Westoby M(2006).Rebuilding community ecology from functional traits.Trends in Ecology and Evolution,21,178-185.
Mensah S,Veldtman R,Assogbadjo AE,GlèlèKR,Seifert T(2016).Tree species diversity promotes aboveground carbon storage through functional diversity and functional dominance.Ecology and Evolution,6,7546-7557.
Micheli F,Halpern BS(2010).Low functional redundancy in coastal marine assemblages.Ecology Letters,8,391-400.
Mokany K,Ash J,Roxburgh S(2008).Functional identity is more important than diversity in influencing ecosystem processes in a temperate native grassland.Journal of Ecology,96,884-893.
Mouchet MA,Villeger S,Mason NWH,Mouillot D(2010).Functional diversity measures:An overview of their redundancy and their ability to discriminate community assembly rules.Functional Ecology,24,867-876.
Oksanen J,Kindt R,Legendre P,O’Hara B,Simpson GL,Slymos P,Stevens MHH,Wagner H(2009).The vegan Package.http://cran.r-project.org/web/packages/vegan/vegan.pdf.Cited:2019-02-04.
Ouyang S,Xiang W,Wang X,Zeng Y,Lei P,Deng X,Peng C(2016).Significant effects of biodiversity on forest biomass during the succession of subtropical forest in south China.Forest Ecology and Management,372,291-302.
Petchey OL,Hector A,Gaston KJ(2004).How do different measures of functional diversity perform?Ecology,85,847-857.
Ratcliffe S,Liebergesell M,Ruiz-Benito P,Madrigal González J,Mu?oz Casta?eda JM,K?ndler G,Lehtonen A,Dahlgren J,Kattge J,Pe?uelas J,Zavala MA,Wirth C(2016).Modes of functional biodiversity control on tree productivity across the European continent.Global Ecology and Biogeography,25,251-262.
Rosseel Y(2012).lavaan:An R package for structural equation modeling.Journal of Statistical Software,48,1-36.
Ruiz-Benito P,Gomez-Aparicio L,Paquette A,Messier C,Kattge J,Zavala MA(2014).Diversity increases carbon storage and tree productivity in Spanish forests.Global Ecology and Biogeography,23,311-322.
Schellberg J,Pontes LDS(2012).Plant functional traits and nutrient gradients on grassland.Grass and Forage Science,67,305-319.
Sun RY,Li QF,Niu CJ,Lou AR(2002).Basic Ecology.Higher Education Press,Beijing.[孙儒泳,李庆芬,牛翠娟,娄安如(2002).基础生态学.高等教育出版社,北京.]
Tilman D(1997).Distinguishing between the effects of species diversity and species composition.Oikos,80,185.
Tobner CM,Paquette A,Gravel D,Reich PB,Williams LJ,Messier C(2016).Functional identity is the main driver of diversity effects in young tree communities.Ecology Letters,19,638-647.
Van MT,Pe?a-Claros M,Ascarrunz N,Arets EJMM,Licona JC,Toledo M,Poorter L(2017).Abiotic and biotic drivers of biomass change in a neotropical forest.Journal of Ecology,105,1223-1234.
Wu Z,Raven P(1994-2009).Flora of China.Science Press and Missouri Botanical Garden Press,Beijing and St Louis.
Zhang Y,Chen HYH(2015).Individual size inequality links forest diversity and above-ground biomass.Journal of Ecology,103,1245-1252.
Zhang Y,Chen HYH,Reich PB(2012).Forest productivity increases with evenness,species richness and trait variation:A global meta-analysis.Journal of Ecology,100,742-749.
BartońK(2018).Package‘MuMIn’.https://cran.r-project.org/web/packages/MuMIn/MuMIn.pdf.Cited:2018-07-21.
Cardinale BJ,Duffy JE,Gonzalez A,Hooper DU,Perrings C,Venail P(2012).Biodiversity loss and its impact on humanity.Nature,489,59-67.
Cardinale BJ,Gonzalez A(2011).The functional role of producer diversity in ecosystems.American Journal of Botany,98,572-592.
Chang LW,Zeleny D,Li CF,Chiu ST,Hsieh CF(2013).Better environmental data may reverse conclusions about nicheand dispersal-based processes in community assembly.Ecology,94,2145-2151.
Chave J,Condit R,Lao S,Caspersen JP,Foster RB,Hubbell SP(2003).Spatial and temporal variation of biomass in a tropical forest:Results from a large census plot in Panama.Journal of Ecology,91,240-252.
Chen CG,Zhu JF(1989).A Handbook for Main Tree Species Biomass in Northeast China.China Forestry Publishing House,Beijing.[陈传国,朱俊凤(1989).东北主要林木生物量手册.中国林业出版社,北京.]
Chen DM,Cheng JH,Chu PF,Mi J,Hu SJ,Xie YC,Tuvshintogtokh I,Bai YF(2016).Effect of diversity on biomass across grasslands on the Mongolian Plateau:Contrasting effects between plants and soil nematodes.Journal of Biogeography,43,955-966.
Cornelissen JHC,Lavorel S,Garnier E,Diaz S,Buchmann N,Gurvich DE,Reich PB,ter Steege H,Morgan HD,van der Heijden MGA,Pausas JG,Poorter H(2003).A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,51,335-380.
Díaz S,Lavorel S,Bello FD,Quétier F,Grigulis K,Robson TM(2007).Incorporating plant functional diversity effects in ecosystem service assessments.Proceedings of the National Academy of Sciences of the United States of America,104,20684-20689.
D?nescu A,Albrecht AT,Bauhus J(2016).Structural diversity promotes productivity of mixed,uneven-aged forests in southwestern Germany.Oecologia,182,319-333.
Edwards JR,Lambert LS(2007).Methods for integrating moderation and mediation:A general analytical framework using moderated path analysis.Psychological Methods,12,1-22.
Finegan B,Pe?a-Claros M,de Oliveira A,Ascarrunz N,Bret-Harte MS,Carre?o-Rocabado G,Casanoves F,Díaz S,Eguiguren Velepucha P,Fernandez F,Licona JC,Lorenzo L,Salgado Negret B,Vaz M,Poorter L,Canham C(2015).Does functional trait diversity predict above-ground biomass and productivity of tropical forests?Testing three alternative hypotheses.Journal of Ecology,103,191-201.
Fox J(2008).Applied Regression Analysis and Generalized Linear Models.2nd edn.Sage Publications,Thousand Oaks,USA.
Fox J,Monette G(1992).Generalized collinearity diagnostics.Journal of the American Statistical Association,87,178-183.
Gazol A,Camarero JJ(2016).Functional diversity enhances silver fir growth resilience to an extreme drought.Journal of Ecology,104,1063-1075.
Grace JB,Anderson TM,Seabloom EW,Borer ET,Adler PB,Harpole WS,Hautier Y,Hillebrand H,Lind EM,Partel M,Bakker JD,Buckley YM,Crawley MJ,Damschen EI,Davies KF,Fay PA,Firn J,Gruner DS,Hector A,Knops JM,MacDougall AS,Melbourne BA,Morgan JW,Orrock JL,Prober SM,Smith MD(2016).Integrative modelling reveals mechanisms linking productivity and plant species richness.Nature,529,390-393.
Grime JP(1998).Benefits of plant diversity to ecosystems:Immediate,filter and founder effects.Journal of Ecology,86,902-910.
Howard AL(2013).Handbook of structural equation modeling.Structural Equation Modeling-a Multidisciplinary Journal,20,354-360.
Huang XR(2018).Relationship between plant functional diversity and productivity of Pinus massoniana plantations in Guangxi.Biodiversity Science,26,690-700.[黄小荣(2018).广西马尾松林植物功能多样性与生产力的关系.生物多样性,26,690-700.]
LalibertéE,Legendre P(2010).A distance-based framework for measuring functional diversity from multiple traits.Ecology,91,299-305.
Lin DM,Lai JS,Muller-Landau HC,Mi XC,Ma KP(2012).Topographic variation in aboveground biomass in a subtropical evergreen broad-leaved forest in China.PLOSONE,DOI:10.1371/journal.pone.0048244.
Lohbeck M,Poorter L,Martinez-Ramos M,Bongers F(2015).Biomass is the main driver of changes in ecosystem process rates during tropical forest succession.Ecology,96,1242-1252.
LüTT,Wang P,Yan H,Zhang W,Liao GX,Jiang HB,Zou CL,Sheng LX(2014).Relationship between functional diversity and productivity in meadow and marsh plant communities.Chinese Journal of Plant Ecology,38,405-416.[吕亭亭,王平,燕红,张稳,廖桂项,姜海波,邹畅林,盛连喜(2014).草甸和沼泽植物群落功能多样性与生产力的关系.植物生态学报,38,405-416.]
McGill BJ,Enquist BJ,Weiher E,Westoby M(2006).Rebuilding community ecology from functional traits.Trends in Ecology and Evolution,21,178-185.
Mensah S,Veldtman R,Assogbadjo AE,GlèlèKR,Seifert T(2016).Tree species diversity promotes aboveground carbon storage through functional diversity and functional dominance.Ecology and Evolution,6,7546-7557.
Micheli F,Halpern BS(2010).Low functional redundancy in coastal marine assemblages.Ecology Letters,8,391-400.
Mokany K,Ash J,Roxburgh S(2008).Functional identity is more important than diversity in influencing ecosystem processes in a temperate native grassland.Journal of Ecology,96,884-893.
Mouchet MA,Villeger S,Mason NWH,Mouillot D(2010).Functional diversity measures:An overview of their redundancy and their ability to discriminate community assembly rules.Functional Ecology,24,867-876.
Oksanen J,Kindt R,Legendre P,O’Hara B,Simpson GL,Slymos P,Stevens MHH,Wagner H(2009).The vegan Package.http://cran.r-project.org/web/packages/vegan/vegan.pdf.Cited:2019-02-04.
Ouyang S,Xiang W,Wang X,Zeng Y,Lei P,Deng X,Peng C(2016).Significant effects of biodiversity on forest biomass during the succession of subtropical forest in south China.Forest Ecology and Management,372,291-302.
Petchey OL,Hector A,Gaston KJ(2004).How do different measures of functional diversity perform?Ecology,85,847-857.
Ratcliffe S,Liebergesell M,Ruiz-Benito P,Madrigal González J,Mu?oz Casta?eda JM,K?ndler G,Lehtonen A,Dahlgren J,Kattge J,Pe?uelas J,Zavala MA,Wirth C(2016).Modes of functional biodiversity control on tree productivity across the European continent.Global Ecology and Biogeography,25,251-262.
Rosseel Y(2012).lavaan:An R package for structural equation modeling.Journal of Statistical Software,48,1-36.
Ruiz-Benito P,Gomez-Aparicio L,Paquette A,Messier C,Kattge J,Zavala MA(2014).Diversity increases carbon storage and tree productivity in Spanish forests.Global Ecology and Biogeography,23,311-322.
Schellberg J,Pontes LDS(2012).Plant functional traits and nutrient gradients on grassland.Grass and Forage Science,67,305-319.
Sun RY,Li QF,Niu CJ,Lou AR(2002).Basic Ecology.Higher Education Press,Beijing.[孙儒泳,李庆芬,牛翠娟,娄安如(2002).基础生态学.高等教育出版社,北京.]
Tilman D(1997).Distinguishing between the effects of species diversity and species composition.Oikos,80,185.
Tobner CM,Paquette A,Gravel D,Reich PB,Williams LJ,Messier C(2016).Functional identity is the main driver of diversity effects in young tree communities.Ecology Letters,19,638-647.
Van MT,Pe?a-Claros M,Ascarrunz N,Arets EJMM,Licona JC,Toledo M,Poorter L(2017).Abiotic and biotic drivers of biomass change in a neotropical forest.Journal of Ecology,105,1223-1234.
Wu Z,Raven P(1994-2009).Flora of China.Science Press and Missouri Botanical Garden Press,Beijing and St Louis.
Zhang Y,Chen HYH(2015).Individual size inequality links forest diversity and above-ground biomass.Journal of Ecology,103,1245-1252.
Zhang Y,Chen HYH,Reich PB(2012).Forest productivity increases with evenness,species richness and trait variation:A global meta-analysis.Journal of Ecology,100,742-749.