抚育采伐对蛟河次生针阔混交林功能结构和谱系结构的影响
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摘要
【目的】研究不同强度森林抚育采伐对温带针阔混交林群落结构、生物多样性及生态系统功能的影响机制,为温带针阔混交林经营管理提供科学依据和理论指导。【方法】以吉林蛟河4块1 hm2的针阔混交林采伐样地为研究对象,样地于2011年7月建立,2011年12月进行抚育采伐作业,并于2015年7月对保留木进行复测。采集、测定叶面积、比叶面积、叶碳含量、叶氮含量、叶片碳氮比和最大树高6类植物功能性状数据,根据物种之间的性状距离和谱系距离构建功能性状树和系统发育树。通过Blomberg’s K值法检验植物功能性状的系统发育信号;利用功能多样性指数和最近邻体性状距离指数计算2011年采伐前和2015采伐后不同空间尺度上(10 m×10 m、20 m×20 m和50 m×50 m)群落功能多样性和功能结构的变化情况;利用谱系多样性指数和最近亲缘关系指数计算群落谱系多样性和谱系结构的变化情况;通过树木4年间平均胸高断面积的增长量来量化保留木的生长差异。【结果】除叶面积外,其余5类功能性状均表现出显著的系统发育信号;抚育采伐前后,群落功能结构和谱系结构的变化具有尺度依赖性,在10 m×10 m尺度上,采伐后群落的离散度上升,聚集度下降,在20 m×20 m和50 m×50 m尺度上,采伐后群落的离散度下降,而聚集度上升;功能多样性和谱系多样性对采伐强度的响应同样存在尺度效应,中低强度抚育采伐对群落多样性的影响仅体现在小尺度上,而高强度抚育采伐即使在大尺度上也会对群落多样性产生显著影响;经历不同强度的抚育采伐后,样地内保留木的生长差异表现为中度采伐样地>重度采伐样地>轻度采伐样地>对照样地。【结论】中低强度的抚育采伐能够优化群落的功能结构和谱系结构,从而达到促进资源利用、加快保留木生长的目的;高强度的采伐会导致生态位空间过度释放,不利于资源的充分利用以及树木生长,也会对森林生物多样性产生严重的负面影响。因此,从调整群落结构、促进保留木生长以及保护生物多样性的角度出发,对温带针阔混交林的抚育采伐作业应控制在中等强度以内。
【Objective】 This paper aims to explore the influences of tending felling on forest structure,biodiversity and ecosystem function,and to provide a better guidance for the management of mixed broadleaf-conifer temperate forest.【Method】 In 2011,four 1 hm2 stem-mapped plots with different felling intensity were established in Jiaohe,Jilin Province. Six plant traits that have been suggested to have great functional significance for plant growth were measured,including leaf area,specific leaf area,leaf carbon,leaf nitrogen,leaf C/N ratio and maximum tree height. All functional traits were determined for 22 woody species. Traits tree was constructed using the distance matrix of functional traits,meanwhile,phylogenetic tree was obtained based on the distance matrix of phylogeny. Phylogenetic signals of functional traits were detected using the Blomberg's K statistics. Functional structure of forest was represented by the functional diversity index and the nearest neighbor trait distance index. Phylogenetic structure was estimated using the phylogenetic diversity index and the nearest taxon index. The potential effects of tending felling on the functional and phylogeneticstructures were examined at three different spatial scales. The influences of felling intensity on radial growth of the residual trees were also assessed.【Result】 Five studied traits( specific leaf area,leaf carbon,leaf nitrogen,leaf C/N ratio and maximum tree height) showed significant phylogenetic signal. The variations of functional and phylogenetic structures depended on the studied scales. At the small scale( 10 m × 10 m),tending felling enhanced the degree of dispersion but reduced the degree of aggregation of community structure. However,at the middle and large scales( 20 m × 20 m and50 m × 50 m),tending felling enhanced the degree of aggregation but reduced the degree of dispersion. The result showed that the influences of felling intensity on the functional and phylogenetic diversity also have scale-dependency. Low and moderate felling intensity showed significant effects only at small study scale,but high felling intensity at all study scales. The radial growths of residual trees were from top to bottom in the order of moderate felling intensity,high felling intensity,low felling intensity and the controls.【Conclusion】Tending felling with low or moderate intensity could improve forest functional and phylogenetic structure and promote the utilization of resources. However,felling with high intensity is unfavorable to make full use of resources,moreover,felling with high intensity has a significant negative impact on biodiversity conservation. Therefore,from the perspective of forest management,low or moderate felling intensity is the best way to regulate community spatial structure,conserve biodiversity and accelerate the growth of trees.
【Objective】 This paper aims to explore the influences of tending felling on forest structure,biodiversity and ecosystem function,and to provide a better guidance for the management of mixed broadleaf-conifer temperate forest.【Method】 In 2011,four 1 hm2 stem-mapped plots with different felling intensity were established in Jiaohe,Jilin Province. Six plant traits that have been suggested to have great functional significance for plant growth were measured,including leaf area,specific leaf area,leaf carbon,leaf nitrogen,leaf C/N ratio and maximum tree height. All functional traits were determined for 22 woody species. Traits tree was constructed using the distance matrix of functional traits,meanwhile,phylogenetic tree was obtained based on the distance matrix of phylogeny. Phylogenetic signals of functional traits were detected using the Blomberg's K statistics. Functional structure of forest was represented by the functional diversity index and the nearest neighbor trait distance index. Phylogenetic structure was estimated using the phylogenetic diversity index and the nearest taxon index. The potential effects of tending felling on the functional and phylogeneticstructures were examined at three different spatial scales. The influences of felling intensity on radial growth of the residual trees were also assessed.【Result】 Five studied traits( specific leaf area,leaf carbon,leaf nitrogen,leaf C/N ratio and maximum tree height) showed significant phylogenetic signal. The variations of functional and phylogenetic structures depended on the studied scales. At the small scale( 10 m × 10 m),tending felling enhanced the degree of dispersion but reduced the degree of aggregation of community structure. However,at the middle and large scales( 20 m × 20 m and50 m × 50 m),tending felling enhanced the degree of aggregation but reduced the degree of dispersion. The result showed that the influences of felling intensity on the functional and phylogenetic diversity also have scale-dependency. Low and moderate felling intensity showed significant effects only at small study scale,but high felling intensity at all study scales. The radial growths of residual trees were from top to bottom in the order of moderate felling intensity,high felling intensity,low felling intensity and the controls.【Conclusion】Tending felling with low or moderate intensity could improve forest functional and phylogenetic structure and promote the utilization of resources. However,felling with high intensity is unfavorable to make full use of resources,moreover,felling with high intensity has a significant negative impact on biodiversity conservation. Therefore,from the perspective of forest management,low or moderate felling intensity is the best way to regulate community spatial structure,conserve biodiversity and accelerate the growth of trees.
引文
曹科,饶米德,余建中,等.2013.古田山木本植物功能性状的系统发育信号及其对群落结构的影响.生物多样性,21(5):564-571.(Cao K,Rao M D,Yu J Z,et al.2013.The phylogenetic signal of functional traits and their effects on community structure in an evergreen broad-leaved forest.Biodiversity Science,21(5):564-571.[in Chinese])
房帅,原作强,蔺菲,等.2014.长白山阔叶红松林木本植物系统发育与功能性状结构.科学通报,59(24):2342-2348.(Fang S,Yuan Z Q,Lin F,et al.2014.Functional and phylogenetic structures of woody plants in broad-leaved korean pine mixed forest in Changbai Mountains,Jilin,China.Chinese Science Bulletin,59(24):2342-2348.[in Chinese])
刘晓娟,马克平.2015.植物功能性状研究进展.中国科学:生命科学,45(4):325-339.(Liu X J,Ma K P.2015.Plant functional traits-concepts,applications and future directions.Science in China:Life Sciences,45(4):325-339.[in Chinese])
宋凯,米湘成,贾琪,等.2011.不同程度人为干扰对古田山森林群落谱系结构的影响.生物多样性,19(2):190-196.(Song K,Mi X C,Jia Q,et al.2011.Variation in phylogenetic structure of forest communities along a human disturbance gradient in Gutianshan forest,China.Biodiversity Science,19(2):190-196.[in Chinese])
Asner G P,Knapp D E,Broadbent E N,et al.2005.Selective logging in the Brazilian Amazon.Science,310(5747):480-482.
Bladon K D,Silins U,Landhusser S M,et al.2006.Differential transpiration by three boreal tree species in response to increased evaporative demand after variable retention harvesting.Agricultural and Forest Meteorology,138(1):104-119.
Blomberg S P,Garland T,Ives A R.2003.Testing for phylogenetic signal in comparative data:behavioral traits are more labile.Evolution,57(4):717-745.
Cadotte M W,Cardinale B J,Oakley T H.2008.Evolutionary history and the effect of biodiversity on plant productivity.Proceedings of the National Academy of Sciences,105(44):17012-17017.
Cannon C H,Peart D R,Leighton M.1998.Tree species diversity in commercially logged Bornean rainforest.Science,281(5381):1366-1368.
Cavender-Bares J,Keen A,Miles B.2006.Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale.Ecology,87(sp7):S109-S122.
Chazdon R L.2003.Tropical forest recovery:legacies of human impact and natural disturbances.Perspectives in Plant Ecology,Evolution and Systematics,6(1):51-71.
Cornelissen J H C,Lavorel S,Garnier E,et al.2003.A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,51(4):335-380.
Curran L M,Caniago I,Paoli G D,et al.1999.Impact of El Ni1o and logging on canopy tree recruitment in Borneo.Science,286(5447):2184-2188.
Dalerum F,Cameron E Z,Kunkel K,et al.2012.Interactive effects of species richness and species traits on functional diversity and redundancy.Theoretical Ecology,5(1):129-139.
Feng G,Svenning J C,Mi X C,et al.2014.Anthropogenic disturbance shapes phylogenetic and functional tree community structure in a subtropical forest.Forest Ecology and Management,(313):188-198.
Flynn D F B,Mirotchnick N,Jain M,et al.2011.Functional and phylogenetic diversity as predictors of biodiversity-ecosystemfunction relationships.Ecology,92(8):1573-1581.
Gadow K V,Schmidt M.1998.Periodische inventuren und eingriffsinventuren.Forst und Holz,53(22):667-671.
Gunderson L H.2000.Ecological resilience-in theory and application.Annual Review of Ecology and Systematic,31(1):425-439.
He J S,Fang J,Wang Z,et al.2006.Stoichiometry and large-scale pattems of leaf carbon and nitrogen in the grassland biomes of China.Oecologia,149(1):115-122.
Kraft N J B,Ackerly D D.2010.Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest.Ecological Monographs,80(3):401-422.
Law R,Illian J,Burslem D F R P,et al.2009.Ecological information from spatial patterns of plants:insights from point process theory.Journal of Ecology,97(4):616-628.
Lebrija-Trejos E,Pérez-García E A,Meave J A,et al.2010.Functional traits and environmental filtering drive community assembly in a species-rich tropical system.Ecology,91(2):386-398.
Liu X J,Swenson N G,Zhang J L,et al.2013.The environment and space,not phylogeny,determine trait dispersion in a subtropical forest.Functional Ecology,27(1):264-272.
Loreau M.2004.Does functional redundancy exist?.Oikos,104(3):606-611.
Losos J B.2008.Phylogenetic niche conservatism,phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species.Ecology Letters,11(10):995-1003.
Meynard C N,Devictor V,Mouillot D,et al.2011.Beyond taxonomic diversity patterns:how doα,βandγcomponents of bird functional and phylogenetic diversity respond to environmental gradients across France?.Global Ecology and Biogeography,20(6):893-903.
Moles A T,Warton D I,Warman L,et al.2009.Global patterns in plant height.Journal of Ecology,97(5):923-932.
Mouchet M A,Villeger S,Mason N W H,et al.2010.Functional diversity measures:an overview of their redundancy and their ability to discriminate community assembly rules.Functional Ecology,24(4):867-876.
Norden N,Letcher S G,Boukili V,et al.2012.Demographic drivers of successional changes in phylogenetic structure across life-history changes in plant communities.Ecology,93(sp8):S70-S82.
Park D S,Potter D.2015.Why close relatives make bad neighbours:phylogenetic conservatism in niche preferences and dispersal disproves Darwin’s naturalization hypothesis in the thistle tribe.Molecular Ecology,24(12):3181-3193.
Petchey O L,Gaston K J.2002.Functional diversity(FD),species richness and community composition.Ecology Letters,5(3):402-411.
Prinzing A.2001.The niche of higher plants:evidence for phylogenetic conservatism.Proceedings of the Royal Society of London B:Biological Sciences,268(1483):2383-2389.
Purschke O,Schmid B C,Sykes M T,et al.2013.Contrasting changes in taxonomic,phylogenetic and functional diversity during a longterm succession:insights into assembly processes.Journal of Ecology,101(4):857-866.
Roscher C,Schumacher J,Gubsch M,et al.2012.Using plant functional traits to explain diversity-productivity relationships.PLo S One,7(5):e36760.
Russo S E,Brown P,Tan S,et al.2008.Interspecific demographic trade-offs and soil-related habitat associations of tree species along resource gradients.Journal of Ecology,96(1):192-203.
Srivastava D S,Cadotte M W,Mac Donald A A M,et al.2012.Phylogenetic diversity and the functioning of ecosystems.Ecology Letters,15(7):637-648.
Stoll P,Bergius E.2005.Pattern and process:competition causes regular spacing of individuals within plant populations.Journal of Ecology,93(2):395-403.
Thomson F J,Moles A T,Auld T D,et al.2011.Seed dispersal distance is more strongly correlated with plant height than with seed mass.Journal of Ecology,99(6):1299-1307.
Thorpe H C,Thomas S C,Caspersen J P.2007a.Residual tree growth responses to partial stand harvest in the black spruce(Picea mariana)boreal forest.Canadian Journal of Forest Research,37(9):1563-1571.
Thorpe H C,Thomas S C.2007b.Partial harvesting in the Canadian boreal:success will depend on stand dynamic responses.The Forestry Chronicle,83(3):319-325.
Uriarte M,Swenson N G,Chazdon R L,et al.2010.Trait similarity,shared ancestry and the structure of neighbourhood interactions in a subtropical wet forest:implications for community assembly.Ecology Letters,13(12):1503-1514.
Webb C O,Ackerly D D,Kembel S W.2008.Phylocom:software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics,24(18):2098-2100.
Webb C O,Ackerly D D,Mc Peek M A,et al.2002.Phylogenies and community ecology.Annual Review of Ecology and Systematics,33(1):475-505.
Webb C O,Donoghue M J.2005.Phylomatic:tree assembly for applied phylogenetics.Molecular Ecology Notes,5(1):181-183.
Wiens J J,Graham C H.2005.Niche conservatism:integrating evolution,ecology,and conservation biology.Annual Review of Ecology,Evolution,and Systematics,(36):519-539.
Wikstr9m N,Savolainen V,Chase M W.2001.Evolution of the angiosperms:calibrating the family tree.Proceedings of the Royal Society of London B:Biological Sciences,268(1482):2211-2220.
Wright I J,Reich P B,Westoby M,et al.2004.The worldwide leaf economics spectrum.Nature,428(6985):821-827.
Zhang C Y,Zhao X H,Gadow K V.2014.Analyzing selective harvest events in three large forest observational studies in north eastern China.Forest Ecology and Management,(316):100-109.
房帅,原作强,蔺菲,等.2014.长白山阔叶红松林木本植物系统发育与功能性状结构.科学通报,59(24):2342-2348.(Fang S,Yuan Z Q,Lin F,et al.2014.Functional and phylogenetic structures of woody plants in broad-leaved korean pine mixed forest in Changbai Mountains,Jilin,China.Chinese Science Bulletin,59(24):2342-2348.[in Chinese])
刘晓娟,马克平.2015.植物功能性状研究进展.中国科学:生命科学,45(4):325-339.(Liu X J,Ma K P.2015.Plant functional traits-concepts,applications and future directions.Science in China:Life Sciences,45(4):325-339.[in Chinese])
宋凯,米湘成,贾琪,等.2011.不同程度人为干扰对古田山森林群落谱系结构的影响.生物多样性,19(2):190-196.(Song K,Mi X C,Jia Q,et al.2011.Variation in phylogenetic structure of forest communities along a human disturbance gradient in Gutianshan forest,China.Biodiversity Science,19(2):190-196.[in Chinese])
Asner G P,Knapp D E,Broadbent E N,et al.2005.Selective logging in the Brazilian Amazon.Science,310(5747):480-482.
Bladon K D,Silins U,Landhusser S M,et al.2006.Differential transpiration by three boreal tree species in response to increased evaporative demand after variable retention harvesting.Agricultural and Forest Meteorology,138(1):104-119.
Blomberg S P,Garland T,Ives A R.2003.Testing for phylogenetic signal in comparative data:behavioral traits are more labile.Evolution,57(4):717-745.
Cadotte M W,Cardinale B J,Oakley T H.2008.Evolutionary history and the effect of biodiversity on plant productivity.Proceedings of the National Academy of Sciences,105(44):17012-17017.
Cannon C H,Peart D R,Leighton M.1998.Tree species diversity in commercially logged Bornean rainforest.Science,281(5381):1366-1368.
Cavender-Bares J,Keen A,Miles B.2006.Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale.Ecology,87(sp7):S109-S122.
Chazdon R L.2003.Tropical forest recovery:legacies of human impact and natural disturbances.Perspectives in Plant Ecology,Evolution and Systematics,6(1):51-71.
Cornelissen J H C,Lavorel S,Garnier E,et al.2003.A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,51(4):335-380.
Curran L M,Caniago I,Paoli G D,et al.1999.Impact of El Ni1o and logging on canopy tree recruitment in Borneo.Science,286(5447):2184-2188.
Dalerum F,Cameron E Z,Kunkel K,et al.2012.Interactive effects of species richness and species traits on functional diversity and redundancy.Theoretical Ecology,5(1):129-139.
Feng G,Svenning J C,Mi X C,et al.2014.Anthropogenic disturbance shapes phylogenetic and functional tree community structure in a subtropical forest.Forest Ecology and Management,(313):188-198.
Flynn D F B,Mirotchnick N,Jain M,et al.2011.Functional and phylogenetic diversity as predictors of biodiversity-ecosystemfunction relationships.Ecology,92(8):1573-1581.
Gadow K V,Schmidt M.1998.Periodische inventuren und eingriffsinventuren.Forst und Holz,53(22):667-671.
Gunderson L H.2000.Ecological resilience-in theory and application.Annual Review of Ecology and Systematic,31(1):425-439.
He J S,Fang J,Wang Z,et al.2006.Stoichiometry and large-scale pattems of leaf carbon and nitrogen in the grassland biomes of China.Oecologia,149(1):115-122.
Kraft N J B,Ackerly D D.2010.Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest.Ecological Monographs,80(3):401-422.
Law R,Illian J,Burslem D F R P,et al.2009.Ecological information from spatial patterns of plants:insights from point process theory.Journal of Ecology,97(4):616-628.
Lebrija-Trejos E,Pérez-García E A,Meave J A,et al.2010.Functional traits and environmental filtering drive community assembly in a species-rich tropical system.Ecology,91(2):386-398.
Liu X J,Swenson N G,Zhang J L,et al.2013.The environment and space,not phylogeny,determine trait dispersion in a subtropical forest.Functional Ecology,27(1):264-272.
Loreau M.2004.Does functional redundancy exist?.Oikos,104(3):606-611.
Losos J B.2008.Phylogenetic niche conservatism,phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species.Ecology Letters,11(10):995-1003.
Meynard C N,Devictor V,Mouillot D,et al.2011.Beyond taxonomic diversity patterns:how doα,βandγcomponents of bird functional and phylogenetic diversity respond to environmental gradients across France?.Global Ecology and Biogeography,20(6):893-903.
Moles A T,Warton D I,Warman L,et al.2009.Global patterns in plant height.Journal of Ecology,97(5):923-932.
Mouchet M A,Villeger S,Mason N W H,et al.2010.Functional diversity measures:an overview of their redundancy and their ability to discriminate community assembly rules.Functional Ecology,24(4):867-876.
Norden N,Letcher S G,Boukili V,et al.2012.Demographic drivers of successional changes in phylogenetic structure across life-history changes in plant communities.Ecology,93(sp8):S70-S82.
Park D S,Potter D.2015.Why close relatives make bad neighbours:phylogenetic conservatism in niche preferences and dispersal disproves Darwin’s naturalization hypothesis in the thistle tribe.Molecular Ecology,24(12):3181-3193.
Petchey O L,Gaston K J.2002.Functional diversity(FD),species richness and community composition.Ecology Letters,5(3):402-411.
Prinzing A.2001.The niche of higher plants:evidence for phylogenetic conservatism.Proceedings of the Royal Society of London B:Biological Sciences,268(1483):2383-2389.
Purschke O,Schmid B C,Sykes M T,et al.2013.Contrasting changes in taxonomic,phylogenetic and functional diversity during a longterm succession:insights into assembly processes.Journal of Ecology,101(4):857-866.
Roscher C,Schumacher J,Gubsch M,et al.2012.Using plant functional traits to explain diversity-productivity relationships.PLo S One,7(5):e36760.
Russo S E,Brown P,Tan S,et al.2008.Interspecific demographic trade-offs and soil-related habitat associations of tree species along resource gradients.Journal of Ecology,96(1):192-203.
Srivastava D S,Cadotte M W,Mac Donald A A M,et al.2012.Phylogenetic diversity and the functioning of ecosystems.Ecology Letters,15(7):637-648.
Stoll P,Bergius E.2005.Pattern and process:competition causes regular spacing of individuals within plant populations.Journal of Ecology,93(2):395-403.
Thomson F J,Moles A T,Auld T D,et al.2011.Seed dispersal distance is more strongly correlated with plant height than with seed mass.Journal of Ecology,99(6):1299-1307.
Thorpe H C,Thomas S C,Caspersen J P.2007a.Residual tree growth responses to partial stand harvest in the black spruce(Picea mariana)boreal forest.Canadian Journal of Forest Research,37(9):1563-1571.
Thorpe H C,Thomas S C.2007b.Partial harvesting in the Canadian boreal:success will depend on stand dynamic responses.The Forestry Chronicle,83(3):319-325.
Uriarte M,Swenson N G,Chazdon R L,et al.2010.Trait similarity,shared ancestry and the structure of neighbourhood interactions in a subtropical wet forest:implications for community assembly.Ecology Letters,13(12):1503-1514.
Webb C O,Ackerly D D,Kembel S W.2008.Phylocom:software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics,24(18):2098-2100.
Webb C O,Ackerly D D,Mc Peek M A,et al.2002.Phylogenies and community ecology.Annual Review of Ecology and Systematics,33(1):475-505.
Webb C O,Donoghue M J.2005.Phylomatic:tree assembly for applied phylogenetics.Molecular Ecology Notes,5(1):181-183.
Wiens J J,Graham C H.2005.Niche conservatism:integrating evolution,ecology,and conservation biology.Annual Review of Ecology,Evolution,and Systematics,(36):519-539.
Wikstr9m N,Savolainen V,Chase M W.2001.Evolution of the angiosperms:calibrating the family tree.Proceedings of the Royal Society of London B:Biological Sciences,268(1482):2211-2220.
Wright I J,Reich P B,Westoby M,et al.2004.The worldwide leaf economics spectrum.Nature,428(6985):821-827.
Zhang C Y,Zhao X H,Gadow K V.2014.Analyzing selective harvest events in three large forest observational studies in north eastern China.Forest Ecology and Management,(316):100-109.