功能多样性和谱系多样性对热带云雾林群落构建的影响
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摘要
近年来,功能多样性和谱系多样性为探究群落构建机制提供了新方法。为了更准确地了解海南岛高海拔热带云雾林群落构建机制,该研究以海南岛霸王岭热带云雾林为对象,测定7个环境因子和13个植物功能性状。利用主成分分析(PCA)筛选环境因子,以霸王岭、尖峰岭和黎母山热带云雾林分布物种建立区域物种库,结合模型,分析Rao二次熵(RaoQ)和平均成对谱系距离(MPD)变化对植物群落构建的影响。结果表明:林冠开阔度、土壤全磷含量和坡度是影响植物群落构建的关键环境因子。多数功能性状的系统发育信号很低且不显著,说明热带云雾林群落的系统发育关系与功能性状随历史进程变化不一致。RaoQ和MPD的实际观测值都显著低于期望值,且其标准效应值与土壤磷含量显著相关,说明生境过滤是驱动热带云雾林群落构建的关键因子,土壤磷含量是群落构建的关键环境筛。
Aims Phylogenetic and functional diversity are important components of species biodiversity, and are thought to play key roles in the mechanisms of community assembly. In this study, we explore the mechanisms of community assembly in tropical cloud forest plant community in Hainan Island, in southern China, using phylogenetic and functional diversity based methods.Methods We constructed a species pool of 186 woody plant species from three tropical cloud forest sites in Hainan Island. For these species, we measured 13 functional traits and assessed their phylogenetic signals. In addition, we measured seven environmental factors and assessed their relationships using Principal component analysis(PCA). Then we chose Rao's quadratic entropy(RaoQ) and mean pairwise distance(MPD) indices to examine the effects of functional diversity and phylogenetic diversity on tropical cloud forest community assembly. To do this we compared these indices to expectations under null models that assume neutral community assembly. We used standard effect sizes to evaluate the influence of environmental factors on community assembly.Important findings Canopy openness, soil total phosphorus content and slope were significant environmental predictors in tropical cloud forest. The phylogenetic signals of most functional traits were very low and not significant, indicating that the phylogenetic relationship and functional traits were not consistent with the change of historical process. The observed values of RaoQ and MPD were significantly lower than expected, and their standard effect sizes were significantly correlated with soil phosphorus content, which suggested that habitat filtering driven by soil phosphorus was the key factor driving the community assembly in tropical cloud forest.
Aims Phylogenetic and functional diversity are important components of species biodiversity, and are thought to play key roles in the mechanisms of community assembly. In this study, we explore the mechanisms of community assembly in tropical cloud forest plant community in Hainan Island, in southern China, using phylogenetic and functional diversity based methods.Methods We constructed a species pool of 186 woody plant species from three tropical cloud forest sites in Hainan Island. For these species, we measured 13 functional traits and assessed their phylogenetic signals. In addition, we measured seven environmental factors and assessed their relationships using Principal component analysis(PCA). Then we chose Rao's quadratic entropy(RaoQ) and mean pairwise distance(MPD) indices to examine the effects of functional diversity and phylogenetic diversity on tropical cloud forest community assembly. To do this we compared these indices to expectations under null models that assume neutral community assembly. We used standard effect sizes to evaluate the influence of environmental factors on community assembly.Important findings Canopy openness, soil total phosphorus content and slope were significant environmental predictors in tropical cloud forest. The phylogenetic signals of most functional traits were very low and not significant, indicating that the phylogenetic relationship and functional traits were not consistent with the change of historical process. The observed values of RaoQ and MPD were significantly lower than expected, and their standard effect sizes were significantly correlated with soil phosphorus content, which suggested that habitat filtering driven by soil phosphorus was the key factor driving the community assembly in tropical cloud forest.
引文
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Andersen KM,Endara MJ,Turner BL,Dalling JW(2012).Trait-based community assembly of understory palms along a soil nutrient gradient in a lower montane tropical forest.Oecologia,168,519-531.
Blomberg SP,Garland T,Ives AR(2003).Testing for phylogenetic signal in comparative data:Behavioral traits are more labile.Evolution,57,717-745.
Botta-Dukát Z,Czúcz B(2016).Testing the ability of functional diversity indices to detect trait convergence and divergence using individual-based simulation.Methods in Ecology and Evolution,7,114-126.
Bu WS,Zang RG,Ding Y(2014).Field observed relationships between biodiversity and ecosystem functioning during secondary succession in a tropical lowland rainforest.Acta Oecologica,55,1-7.
Cadotte MW,Cardinale BJ,Oakley TH(2008).Evolutionary history and the effect of biodiversity on plant productivity.Proceedings of the National Academy of Sciences of the United States of America,105,17012-17017.
Cavender-Bares J,Kozak KH,Fine PVA,Kembel SW(2009).The merging of community ecology and phylogenetic biology.Ecology Letters,12,693-715.
CBOL Plant Wording Group(2009).A DNA barcode for land plants.Proceedings of the National Academy of Sciences of the United States of America,106,12794-12797.
Cornelissen JHC,Lavorel S,Garnier E,Diaz S,Buchmann N,Gurvich DE,Reich PB,Steege H,Morgan HD,Heijden A,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.
Cornell HV(1985).Local and regional richness of cynipine gall wasps on California oaks.Ecology,66,1247-1260.
Cornell HV,Harrison SP(2014).What are species pools and when are they important?Annual Review of Ecology,Evolution,and Systematics,45,45-67.
de Bello F,Price JN,Münkemüller T,Liira J,Zobel M,Thuiller W,Gerhold P,G?tzenberger L,Lavergne S,Lep?S,Zobel K,P?rtel M(2012).Functional species pool framework to test for biotic effects on community assembly.Ecology,93,2263-2273.
Eller CB,Burgess SSO,Oliveira RS(2015).Environmental controls in the water use patterns of a tropical cloud forest tree species,Drimys brasiliensis(Winteraceae).Tree Physiology,35,387-399.
Eriksson O(1993).The species-pool hypothesis and plant community diversity.Oikos,68,371-374.
Goldsmith GR,Matzke NJ,Dawson TE(2013).The incidence and implications of clouds for cloud forest plant water relations.Ecology Letters,16,307-314.
Gonzalez-Caro S,Umana MN,Alvarez E,Stevenson PR,Swenson NG(2014).Phylogenetic alpha and beta diversity in tropical tree assemblages along regional-scale environmental gradients in northwest South America.Journal of Plant Ecology,7,145-153.
Hardy OJ(2008).Testing the spatial phylogenetic structure of local communities:Statistical performances of different null models and test statistics on a locally neutral community.Journal of Ecology,96,914-926.
Harrison S,Cornell H(2008).Toward a better understanding of the regional causes of local community richness.Ecology Letters,11,969-979.
He JS,Wang X,Flynn DFB,Wang L,Schmid B,Fang J(2009).Taxonomic,phylogenetic,and environmental trade-offs between leaf productivity and persistence.Ecology,90,2779-2791.
Kang Y,Deng Z,Zang R,Long W(2017).DNA barcoding analysis and phylogenetic relationships of tree species in tropical cloud forests.Scientific Reports,7,12564.DOI:10.1038/s41598-017-13057-0.
Kraft NJB,Ackerly DD(2010).Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest.Ecological Monographs,80,401-422.
Kraft NJB,Valencia R,Ackerly DD(2008).Functional traits and niche-based tree community assembly in an Amazonia forest.Science,322,580-582.
Long WX,Schamp BS,Zang RG,Ding Y,Huang YF,Xiang YZ(2015a).Community assembly in a tropical cloud forest related to specific leaf area and maximum species height.Journal of Vegetation Science,47,416-423.
Long WX,Xiong MH,Zang RG,Schamp BS,Yang XB,Ding Y,Huang YF,Xiang YZ(2015b).Changes in patterns of species co-occurrence across two tropical cloud forests differing in soil nutrients and air temperature.Biotropica,47,416-423.
Long WX,Zang RG,Ding Y(2011a).Air temperature and soil phosphorus availability correlate with trait differences between two types of tropical cloud forests.Flora,206,896-903.
Long WX,Zang RG,Schamp BS,Ding Y(2011b).Within-and among-species variation in specific leaf area drive community assembly in a tropical cloud forest.Oecologia,167,1103-1113.
Losos JB(2008).Phylogenetic niche conservatism,phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species.Ecology Letters,11,995-1007.
Luna-Vega I,Magallón S(2010).Phylogenetic composition of angiosperm diversity in the cloud forests of Mexico.Biotropica,42,444-454.
Mason NWH,Richardson SJ,Peltzer DA,Bello FD,Wardle DA,Allen RB(2012).Changes in coexistence mechanisms along a long-term soil chronosequence revealed by functional trait diversity.Journal of Ecology,100,678-689.
McGill BJ,Enquist BJ,Weiher E,Westoby M(2006).Rebuilding community ecology from functional traits.Trends in Ecology&Evolution,21,178-185.
Mc Intire EJB,Fajardo A(2014).Facilitation as a ubiquitous driver of biodiversity.New Phytologist,201,403-416.
Mori AS,Shiono T,Koide D,Kitagawa R,Ota AT,Mizumachi E(2013).Community assembly processes shape an altitudinal gradient of forest biodiversity.Global Ecology and Biogeography,22,878-888.
Naeem S,Wright JP(2003).Disentangling biodiversity effects on ecosystem functioning:deriving solutions to a seemingly insurmountable problem.Ecology Letters,6,567-579.
Niu KC,Liu YN,Shen ZH,He FL,Fang JY(2009).Community assemble:The relative important of neutral theory and niche theory.Biodiversity Science,17,579-593.[牛克昌,刘怿宁,沈泽昊,何芳良,方精云(2009).群落构建的中性理论和生态位理论.生物多样性,17,579-593.]
Paine CET,Baraloto C,Chave J,Hérault B(2011).Functional traits of individual trees reveal ecological constraints on community assembly in tropical rain forests.Oikos,120,720-727.
P?rtel M,Szava-Kovats R,Zobel M(2011).Dark diversity:Shedding light on absent species.Trends in Ecology&Evolution,26,124-128.
Pearse WD,Jones A,Purvis A(2013).Barro Colorado Island’s phylogenetic assemblage structure across fine spatial scales and among clades of different ages.Ecology,94,2861-2872.
Ricklefs RE(1987).Community diversity:Relative roles of local and regional processes.Science,235,206-207.
Spasojevic MJ,Suding KN(2012).Inferring community assembly mechanisms from functional diversity patterns:The importance of multiple assembly processes.Journal of Ecology,100,652-661.
Srivastava DS(1999).Using local-regional richness plots to test for species saturation:Pitfalls and potentials.Journal of Animal Ecology,68,1-16
Swenson NG(2011).Phylogenetic beta diversity metrics,trait evolution and inferring the functional beta diversity of communities.PLOS ONE,6,e21264.DOI:10.137/journal.pone.0021264.
Swenson NG(2013).The assembly of tropical tree communitiesthe advances and shortcomings of phylogenetic and functional trait analyses.Ecography,36,264-276.
Swenson NG,Enquist BJ(2009).Opposing assembly mechanisms in a Neotropical dry forest:Implications for phylogenetic and functional community ecology.Ecology,90,2161-2170.
Swenson NG,Enquist BJ,Pither J,Thompson J,Zimmerman JK(2006).The problem and promise of scale dependency in community phylogenetics.Ecology,87,2418-2424.
Swenson NG,Erickson DL,Mi X,Bourg NA,Forero-Monta?a J,Ge XJ,Howe R,Lake JK,Liu XJ,Ma KP,Pei NC,Thomson J,Uriarte M,Wolf A,Wright SJ,Ye WH,Zhang JL,Zimmerman JK,Kress WJ(2012).Phylogenetic and functional alpha and beta diversity in temperate and tropical tree communities.Ecology,93,S112-S125.
Vamosi SM,Heard SB,Vamosi JC,Webb CO(2009).Emerging patterns in the comparative analysis of phylogenetic community structure.Molecular Ecology,18,572-592.
Wang XX,Long WX,Yang XB,Xiong MH,Kang Y,Huang J,Wang X,Hong XJ,Zhou ZL,Lu YQ,Fang J,Li SX(2016).Patterns of plant diversity within and among three tropical cloud forest communities in Hainan Island.Chinese Journal of Plant Ecology,40,469-479.[王茜茜,龙文兴,杨小波,熊梦辉,康勇,黄瑾,王旭,洪小江,周照骊,陆雍泉,方精,李时兴(2016).海南岛3个林区热带云雾林植物多样性变化.植物生态学报,40,469-479.]
Webb CO,Ackerly DD,Kembel SW(2008).Phylocom:Software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics,24,2098-2100.
Webb CO,Ackerly DD,Mcpeek MA,Donoghue MJ(2002).Phylogenies and community ecology.Annual Review of Ecology and Systematics,33,475-505.
Weiher E,Keddy CPA(1998).Community assembly rules,morphological dispersion,and the coexistence of plant species.Oikos,81,309-322.
Yang J,Zhang G,Ci X,Swenson NG,Cao M,Sha LQ,Li J,Baskin CC,Slik JWF,Lin LX(2014).Functional and phylogenetic assembly in a Chinese tropical tree community across size classes,spatial scales and habitats.Functional Ecology,28,520-529.
Zobel M,Scheiner S(2016).The species pool concept as a framework for studying patterns of plant diversity:Official organ of the International Association for Vegetation Science.Journal of Vegetation Science,27,8-18.
Andersen KM,Endara MJ,Turner BL,Dalling JW(2012).Trait-based community assembly of understory palms along a soil nutrient gradient in a lower montane tropical forest.Oecologia,168,519-531.
Blomberg SP,Garland T,Ives AR(2003).Testing for phylogenetic signal in comparative data:Behavioral traits are more labile.Evolution,57,717-745.
Botta-Dukát Z,Czúcz B(2016).Testing the ability of functional diversity indices to detect trait convergence and divergence using individual-based simulation.Methods in Ecology and Evolution,7,114-126.
Bu WS,Zang RG,Ding Y(2014).Field observed relationships between biodiversity and ecosystem functioning during secondary succession in a tropical lowland rainforest.Acta Oecologica,55,1-7.
Cadotte MW,Cardinale BJ,Oakley TH(2008).Evolutionary history and the effect of biodiversity on plant productivity.Proceedings of the National Academy of Sciences of the United States of America,105,17012-17017.
Cavender-Bares J,Kozak KH,Fine PVA,Kembel SW(2009).The merging of community ecology and phylogenetic biology.Ecology Letters,12,693-715.
CBOL Plant Wording Group(2009).A DNA barcode for land plants.Proceedings of the National Academy of Sciences of the United States of America,106,12794-12797.
Cornelissen JHC,Lavorel S,Garnier E,Diaz S,Buchmann N,Gurvich DE,Reich PB,Steege H,Morgan HD,Heijden A,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.
Cornell HV(1985).Local and regional richness of cynipine gall wasps on California oaks.Ecology,66,1247-1260.
Cornell HV,Harrison SP(2014).What are species pools and when are they important?Annual Review of Ecology,Evolution,and Systematics,45,45-67.
de Bello F,Price JN,Münkemüller T,Liira J,Zobel M,Thuiller W,Gerhold P,G?tzenberger L,Lavergne S,Lep?S,Zobel K,P?rtel M(2012).Functional species pool framework to test for biotic effects on community assembly.Ecology,93,2263-2273.
Eller CB,Burgess SSO,Oliveira RS(2015).Environmental controls in the water use patterns of a tropical cloud forest tree species,Drimys brasiliensis(Winteraceae).Tree Physiology,35,387-399.
Eriksson O(1993).The species-pool hypothesis and plant community diversity.Oikos,68,371-374.
Goldsmith GR,Matzke NJ,Dawson TE(2013).The incidence and implications of clouds for cloud forest plant water relations.Ecology Letters,16,307-314.
Gonzalez-Caro S,Umana MN,Alvarez E,Stevenson PR,Swenson NG(2014).Phylogenetic alpha and beta diversity in tropical tree assemblages along regional-scale environmental gradients in northwest South America.Journal of Plant Ecology,7,145-153.
Hardy OJ(2008).Testing the spatial phylogenetic structure of local communities:Statistical performances of different null models and test statistics on a locally neutral community.Journal of Ecology,96,914-926.
Harrison S,Cornell H(2008).Toward a better understanding of the regional causes of local community richness.Ecology Letters,11,969-979.
He JS,Wang X,Flynn DFB,Wang L,Schmid B,Fang J(2009).Taxonomic,phylogenetic,and environmental trade-offs between leaf productivity and persistence.Ecology,90,2779-2791.
Kang Y,Deng Z,Zang R,Long W(2017).DNA barcoding analysis and phylogenetic relationships of tree species in tropical cloud forests.Scientific Reports,7,12564.DOI:10.1038/s41598-017-13057-0.
Kraft NJB,Ackerly DD(2010).Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest.Ecological Monographs,80,401-422.
Kraft NJB,Valencia R,Ackerly DD(2008).Functional traits and niche-based tree community assembly in an Amazonia forest.Science,322,580-582.
Long WX,Schamp BS,Zang RG,Ding Y,Huang YF,Xiang YZ(2015a).Community assembly in a tropical cloud forest related to specific leaf area and maximum species height.Journal of Vegetation Science,47,416-423.
Long WX,Xiong MH,Zang RG,Schamp BS,Yang XB,Ding Y,Huang YF,Xiang YZ(2015b).Changes in patterns of species co-occurrence across two tropical cloud forests differing in soil nutrients and air temperature.Biotropica,47,416-423.
Long WX,Zang RG,Ding Y(2011a).Air temperature and soil phosphorus availability correlate with trait differences between two types of tropical cloud forests.Flora,206,896-903.
Long WX,Zang RG,Schamp BS,Ding Y(2011b).Within-and among-species variation in specific leaf area drive community assembly in a tropical cloud forest.Oecologia,167,1103-1113.
Losos JB(2008).Phylogenetic niche conservatism,phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species.Ecology Letters,11,995-1007.
Luna-Vega I,Magallón S(2010).Phylogenetic composition of angiosperm diversity in the cloud forests of Mexico.Biotropica,42,444-454.
Mason NWH,Richardson SJ,Peltzer DA,Bello FD,Wardle DA,Allen RB(2012).Changes in coexistence mechanisms along a long-term soil chronosequence revealed by functional trait diversity.Journal of Ecology,100,678-689.
McGill BJ,Enquist BJ,Weiher E,Westoby M(2006).Rebuilding community ecology from functional traits.Trends in Ecology&Evolution,21,178-185.
Mc Intire EJB,Fajardo A(2014).Facilitation as a ubiquitous driver of biodiversity.New Phytologist,201,403-416.
Mori AS,Shiono T,Koide D,Kitagawa R,Ota AT,Mizumachi E(2013).Community assembly processes shape an altitudinal gradient of forest biodiversity.Global Ecology and Biogeography,22,878-888.
Naeem S,Wright JP(2003).Disentangling biodiversity effects on ecosystem functioning:deriving solutions to a seemingly insurmountable problem.Ecology Letters,6,567-579.
Niu KC,Liu YN,Shen ZH,He FL,Fang JY(2009).Community assemble:The relative important of neutral theory and niche theory.Biodiversity Science,17,579-593.[牛克昌,刘怿宁,沈泽昊,何芳良,方精云(2009).群落构建的中性理论和生态位理论.生物多样性,17,579-593.]
Paine CET,Baraloto C,Chave J,Hérault B(2011).Functional traits of individual trees reveal ecological constraints on community assembly in tropical rain forests.Oikos,120,720-727.
P?rtel M,Szava-Kovats R,Zobel M(2011).Dark diversity:Shedding light on absent species.Trends in Ecology&Evolution,26,124-128.
Pearse WD,Jones A,Purvis A(2013).Barro Colorado Island’s phylogenetic assemblage structure across fine spatial scales and among clades of different ages.Ecology,94,2861-2872.
Ricklefs RE(1987).Community diversity:Relative roles of local and regional processes.Science,235,206-207.
Spasojevic MJ,Suding KN(2012).Inferring community assembly mechanisms from functional diversity patterns:The importance of multiple assembly processes.Journal of Ecology,100,652-661.
Srivastava DS(1999).Using local-regional richness plots to test for species saturation:Pitfalls and potentials.Journal of Animal Ecology,68,1-16
Swenson NG(2011).Phylogenetic beta diversity metrics,trait evolution and inferring the functional beta diversity of communities.PLOS ONE,6,e21264.DOI:10.137/journal.pone.0021264.
Swenson NG(2013).The assembly of tropical tree communitiesthe advances and shortcomings of phylogenetic and functional trait analyses.Ecography,36,264-276.
Swenson NG,Enquist BJ(2009).Opposing assembly mechanisms in a Neotropical dry forest:Implications for phylogenetic and functional community ecology.Ecology,90,2161-2170.
Swenson NG,Enquist BJ,Pither J,Thompson J,Zimmerman JK(2006).The problem and promise of scale dependency in community phylogenetics.Ecology,87,2418-2424.
Swenson NG,Erickson DL,Mi X,Bourg NA,Forero-Monta?a J,Ge XJ,Howe R,Lake JK,Liu XJ,Ma KP,Pei NC,Thomson J,Uriarte M,Wolf A,Wright SJ,Ye WH,Zhang JL,Zimmerman JK,Kress WJ(2012).Phylogenetic and functional alpha and beta diversity in temperate and tropical tree communities.Ecology,93,S112-S125.
Vamosi SM,Heard SB,Vamosi JC,Webb CO(2009).Emerging patterns in the comparative analysis of phylogenetic community structure.Molecular Ecology,18,572-592.
Wang XX,Long WX,Yang XB,Xiong MH,Kang Y,Huang J,Wang X,Hong XJ,Zhou ZL,Lu YQ,Fang J,Li SX(2016).Patterns of plant diversity within and among three tropical cloud forest communities in Hainan Island.Chinese Journal of Plant Ecology,40,469-479.[王茜茜,龙文兴,杨小波,熊梦辉,康勇,黄瑾,王旭,洪小江,周照骊,陆雍泉,方精,李时兴(2016).海南岛3个林区热带云雾林植物多样性变化.植物生态学报,40,469-479.]
Webb CO,Ackerly DD,Kembel SW(2008).Phylocom:Software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics,24,2098-2100.
Webb CO,Ackerly DD,Mcpeek MA,Donoghue MJ(2002).Phylogenies and community ecology.Annual Review of Ecology and Systematics,33,475-505.
Weiher E,Keddy CPA(1998).Community assembly rules,morphological dispersion,and the coexistence of plant species.Oikos,81,309-322.
Yang J,Zhang G,Ci X,Swenson NG,Cao M,Sha LQ,Li J,Baskin CC,Slik JWF,Lin LX(2014).Functional and phylogenetic assembly in a Chinese tropical tree community across size classes,spatial scales and habitats.Functional Ecology,28,520-529.
Zobel M,Scheiner S(2016).The species pool concept as a framework for studying patterns of plant diversity:Official organ of the International Association for Vegetation Science.Journal of Vegetation Science,27,8-18.