邻域竞争对长白山阔叶红松林关键树种生长的影响
|
摘要
林木间的竞争是影响树木生长、形态和死亡的主要因素.单木邻域竞争分析能够反映个体间相互作用规律及其距离范围,对于减缓林木竞争、促进林木生长具有重要意义.为弄清竞争对阔叶红松林林木生长的影响,本研究基于Hegyi单木竞争指数和邻域分析方法,探讨了长白山原始阔叶红松林中的5个关键树种——红松、紫椴、水曲柳、蒙古栎和春榆(胸高断面积合计占80%)竞争的邻域半径,并分析了竞争对关键树种生长和死亡的影响.结果表明:红松、紫椴、水曲柳和蒙古栎4个树种单木竞争的邻域半径均为11 m,春榆为13 m.关键树种单木邻域竞争强度与其生长量的对数呈显著负相关,与树木个体的大小呈显著正相关;竞争强度对树木生长影响的相对重要性随着个体的生长而降低.邻域竞争显著增加了关键树种的死亡率.本研究表明长白山阔叶红松林中邻域竞争对关键树种的生长和存活有重要影响,研究结果对阔叶红松林关键树种竞争环境的调整和生产力的提升具有指导意义.
Competition is the main factor affecting the growth, morphology and death of trees in fore-sts. The analysis of individual competition can reflect the characteristics of interaction among individuals and their interaction ranges, which is important for reducing individual competition and promoting tree growth. To understand the effects of competition on tree growth in broad-leaved Korean pine forest, based on Hegyi single-tree competition index and neighborhood analysis method, we explored the neighborhood radius of competition for five key tree species, i.e. Pinus koraiensis, Tilia amurensis, Fraxinus mandshurica, Quercus mongolica and Ulmus japonica(80% of basal area at breast height in total), and analyzed the effects of competition on the growth and death of the key tree species. The results showed that the neighborhood radius of single-tree competition of four tree species, P. koraiensis, T. amurensis, F. mandshurica and Q. mongolica was 11 m, while that of U. pumila was 13 m. The single-tree competition intensity for all five key tree species was negatively correlated with the logarithm of its growth increment, and positively correlated with the size of individual trees. The relative importance of competition intensity on tree growth decreased with tree growth. Neighborhood competition significantly increased tree mortality. Our results revealed the effects of neighborhood competition on the growth and survival of the key tree species at different developmental stages in broad-leaved Korean pine forests in Changbai Mountain. The results are instructive to the adjustment of competitive environment and the improvement of productivity of key tree species in broad-leaved Korean pine forests.
Competition is the main factor affecting the growth, morphology and death of trees in fore-sts. The analysis of individual competition can reflect the characteristics of interaction among individuals and their interaction ranges, which is important for reducing individual competition and promoting tree growth. To understand the effects of competition on tree growth in broad-leaved Korean pine forest, based on Hegyi single-tree competition index and neighborhood analysis method, we explored the neighborhood radius of competition for five key tree species, i.e. Pinus koraiensis, Tilia amurensis, Fraxinus mandshurica, Quercus mongolica and Ulmus japonica(80% of basal area at breast height in total), and analyzed the effects of competition on the growth and death of the key tree species. The results showed that the neighborhood radius of single-tree competition of four tree species, P. koraiensis, T. amurensis, F. mandshurica and Q. mongolica was 11 m, while that of U. pumila was 13 m. The single-tree competition intensity for all five key tree species was negatively correlated with the logarithm of its growth increment, and positively correlated with the size of individual trees. The relative importance of competition intensity on tree growth decreased with tree growth. Neighborhood competition significantly increased tree mortality. Our results revealed the effects of neighborhood competition on the growth and survival of the key tree species at different developmental stages in broad-leaved Korean pine forests in Changbai Mountain. The results are instructive to the adjustment of competitive environment and the improvement of productivity of key tree species in broad-leaved Korean pine forests.
引文
[1] Luo Y,Chen H.Competition,species interaction and ageing control tree mortality in boreal forests.Journal of Ecology,2011,99:1470-1480
[2] Comita LS,Muller HC,Aguilar S,et al.Asymmetric density dependence shapes species abundances in a tropical tree community.Science,2010,329:330-332
[3] Condit R.The importance of demographic niches to tree diversity.Science,2006,313:98-101
[4] Das A,Battles J,Stephenson NL,et al.The contribution of competition to tree mortality in old-growth coni-ferous forests.Forest Ecology and Management,2011,261:1203-1213
[5] Enrique O,Vanclay JK.Competition and dominance between angiosperms and Araucaria angustifolia,(Bert.) O.Kuntze in the Atlantic Forest in southern Brazil.Forest Ecology and Management,2018,425:119-125
[6] Aleinikovas M,Linkevi■.The impact of competition for growing space on diameter,basal area and height growth in pine trees.Baltic Fores-try,2014,20:301-313
[7] Weiner J.Neighborhood interference amongst Pinus rigida individuals.Journal of Ecology,1984,72:183-195
[8] Philip JB.Some limitations inherent to static indices of plant competition.Canadian Journal of Forest Research,1993,23:2141-2152
[9] Fraver S,Anthony WD,Bradford JB,et al.Tree growth and competition in an old-growth Picea abies forest of boreal Sweden:Influence of tree spatial patterning.Journal of Vegetation Science,2014,25:374-385
[10] Thorpe HC,Astrup R,Trowbridge A,et al.Competition and tree crowns:A neighborhood analysis of three boreal tree species.Forest Ecology and Management,2010,259:1586-1596
[11] Curzon MT,Anthony WD,Fraver S,et al.Succession,climate,and neighborhood dynamics influence tree growth over time:An 87-year record of change in a Pinus resinosa-dominated forest,Minnesota,USA.Journal of Vegetation Science,2017,28:82-92
[12] Weiskittel AR,Hann DW,Kershaw JA,et al.Forest Growth and Yield Modeling.West Sussex:John Wiley & Sons Incorporated,2011
[13] Dai L-M (代力民),Gu H-Y (谷会岩),Shao G-F (邵国凡),et al.The Broadleaved-Korean Pine Mixed Forest on Changbai Mountain of China.Shenyang:Liao-ning Science and Technology Press,2004 (in Chinese)
[14] Du H-Y (杜洪业),Xu C-Y (徐程扬).Research process of the relationship between keystone species and the dynamic of community in forestry ecosystem.World Forestry Research (世界林业研究),2012,25(5):35-39 (in Chinese)
[15] Sun X-C (孙晓程),Zhang Y-Z (张亚洲),Fan X-N (范晓男).The status and operating management of broad leaved Pinus koriensis forests in Changbai Mountain.Journal of Jilin Normal University (Natural Science) (吉林师范大学学报:自然科学版),2008,29(2):105-107 (in Chinese)
[16] Yu D-P (于大炮),Zhou W-M (周旺明),Bao Y (包也),et al.Forest management of Korean pine and broadleaf mixed forest in Northeast China since the implementation of Natural Forest Protection Project.Acta Ecologica Sinica (生态学报),2015,35(1):10-17 (in Chinese)
[17] Hui GY,Wang Y,Zhang GQ,et al.A novel approach for assessing the neighborhood competition in two diffe-rent aged forests.Forest Ecology and Management,2018,422:49-58
[18] Wang L-W (王利伟),Li B-H (李步杭),Ye J (叶吉),et al.Dynamics of short-term tree mortality in broad-leaved Korean pine (Pinus koraiensis) mixed forest in the Changbai Mountains.Biodiversity Science (生物多样性),2011,19(2):260-270 (in Chinese)
[19] Zhang ZC,Papaik MJ,Wang XG,et al.The effect of tree size,neighborhood competition and environment on tree growth in an old-growth temperate forest.Journal of Plant Ecology,2016,10:970-980
[20] Hegyi F.A simulation model for managing jack-pine stands// Fries J,ed.Growth Models for Tree and Stand Simulation.Proceedings of IUFRO Meeting S4.01.04,Royal College of Forestry,Stockolm,1974:74-90
[21] Vanclay JK.Spatially-explicit competition indices and the analysis of mixed-species plantings with the Simile modelling environment.Forest Ecology and Management,2006,366:295-302
[22] Aho K,Derryberry DW,Peterson T.Model selection for ecologists:The worldviews of AIC and BIC.Ecology,2016,95:631-636
[23] Burnham KP,Anderson DR.Model Selection and Multimodal Inference:A Practical Information-theoretic Approach.2nd Ed.New York:Springer,2003
[24] Chao L (巢林),Liu Y-Y (刘艳艳),Zhang W-D (张伟东),et al.Diameter class structure,growth dynamic,intraspecific and interspecific competition of Betula platyphylla on swamp ecotone.Chinese Journal of Ecology (生态学杂志),2017,36(3):577-583 (in Chinese)
[25] Xu J (徐建),Wei X-L (韦新良),Wang J (王敬),et al.Intraspecific and interspecific competition of dominant species in a deciduous,broadleaf forest of Longwang Mountain.Journal of Zhejiang A&F University (浙江农林大学学报),2014,31(6):868-876 (in Chinese)
[26] Kang D,Deng J,Qin XW,et al.Effect of competition on spatial patterns of oak forests on the Chinese Loess Plateau.Journal of Arid Land,2017,9:122-131
[27] Zhang Z-C (张昭臣).Dynamics of Tree Mortality and Growth in Coniferous and Broad-leaved Mixed Forests in Changbai Mountain.PhD Thesis.Beijing:University of Chinese Academy of Sciences,2010 (in Chinese)
[28] Aleinikovas M,Edgaras L,Andrius K,et al.The impact of competition for growing space on diameter,basal area and height growth in pine trees.Baltic Fores-try,2014,20:301-313
[29] Huang JG,Stadt KJ,Dawson A,et al.Modelling growth-competition relationships in trembling aspen and white spruce mixed boreal forests of western Canada.PLoS One,2013,8(10):77607
[30] Han D-X (韩大校),Jin G-Z (金光泽).Influences of topography and competition on DBH growth in different growth stages in a typical mixed broadleaved-Korean pine forest,northeastern China.Journal of Beijing Forestry University (北京林业大学学报),2017,39(1):9-19 (in Chinese)
[31] Bella IE.A new competition model for individual trees.Forest Science,1971,17:364-372
[32] David SG,Zavala MA,Schalkwijk DB,et al.Rank reversals in tree growth along tree size,competition and climatic gradients for four forest canopy dominant species in Central Spain.Annals of Forest Science,2008,65:605
[33] Cai L-R (蔡礼荣),Kuang X (匡旭),Fang S (房帅),et al.Factors influencing tree radial growth of three common species in broad-leaved Korean pine mixed forests in Changbai Mountains,China.Chinese Journal of Applied ecology (应用生态学报),2017,28(5):1407-1413 (in Chinese)
[34] Macfarlane DW,Kobe RK.Selecting models for capturing tree-size effects on growth resource relationships.Canadian Journal of Forest Research,2006,36:1695-1704
[35] Pritchard JM,Comeau PG.Effects of opening size and stand characteristics on light transmittance and temperature under young trembling aspen stands.Forest Ecology and Management,2004,200:119-128
[36] Schwinning S,Weiner J.Mechanisms determining the degree of size asymmetry in competition among plants.Oecologia,1998,113:447-455
[37] Weiner J.Asymmetric competition in plant populations.Trends in Ecology & Evolution,1990,5:360-364
[38] Wyckoff PH,Clark JS.Tree growth prediction using size and exposed crown area.Canadian Journal of Forest Research,2005,35:13-20
[39] Gonzalez AE,Meakem V,Eng CY,et al.Patterns of tree mortality in a temperate deciduous forest derived from a large forest dynamics plot.Ecosphere,2016,7:e01595,doi:10.1002/ecs2.1595
[40] David AC,Robert BA.Mortality and tree-size distributions in natural mixed-age forests.Journal of Ecology,2007,95:27-40
[41] Tong X (童鑫).Exploring Community Assembly Through the Lens of Spatial Structure:From Population Genetics to Community Composition.PhD Thesis.Shanghai:East China Normal University,2015 (in Chinese)
[42] Zhang Z-C (张昭臣),Hao Z-Q (郝占庆),Ye J (叶吉),et al.Short-term death dynamics of trees in natural secondary poplar-birch forest in Changbai Mountains of Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2013,24(2):303-310 (in Chinese)
[43] Canham CD,Papaik MJ,Latty EF.Interspecific variation in susceptibility to windthrow as a function of tree size and storm severity for northern temperate tree species.Canadian Journal of Forest Research,2001,31:1-10
[44] Takahashi K.Effects of altitude and competition on growth and mortality of the conifer Abies sachalinensis.Ecological Research,2010,25:801-812
[2] Comita LS,Muller HC,Aguilar S,et al.Asymmetric density dependence shapes species abundances in a tropical tree community.Science,2010,329:330-332
[3] Condit R.The importance of demographic niches to tree diversity.Science,2006,313:98-101
[4] Das A,Battles J,Stephenson NL,et al.The contribution of competition to tree mortality in old-growth coni-ferous forests.Forest Ecology and Management,2011,261:1203-1213
[5] Enrique O,Vanclay JK.Competition and dominance between angiosperms and Araucaria angustifolia,(Bert.) O.Kuntze in the Atlantic Forest in southern Brazil.Forest Ecology and Management,2018,425:119-125
[6] Aleinikovas M,Linkevi■.The impact of competition for growing space on diameter,basal area and height growth in pine trees.Baltic Fores-try,2014,20:301-313
[7] Weiner J.Neighborhood interference amongst Pinus rigida individuals.Journal of Ecology,1984,72:183-195
[8] Philip JB.Some limitations inherent to static indices of plant competition.Canadian Journal of Forest Research,1993,23:2141-2152
[9] Fraver S,Anthony WD,Bradford JB,et al.Tree growth and competition in an old-growth Picea abies forest of boreal Sweden:Influence of tree spatial patterning.Journal of Vegetation Science,2014,25:374-385
[10] Thorpe HC,Astrup R,Trowbridge A,et al.Competition and tree crowns:A neighborhood analysis of three boreal tree species.Forest Ecology and Management,2010,259:1586-1596
[11] Curzon MT,Anthony WD,Fraver S,et al.Succession,climate,and neighborhood dynamics influence tree growth over time:An 87-year record of change in a Pinus resinosa-dominated forest,Minnesota,USA.Journal of Vegetation Science,2017,28:82-92
[12] Weiskittel AR,Hann DW,Kershaw JA,et al.Forest Growth and Yield Modeling.West Sussex:John Wiley & Sons Incorporated,2011
[13] Dai L-M (代力民),Gu H-Y (谷会岩),Shao G-F (邵国凡),et al.The Broadleaved-Korean Pine Mixed Forest on Changbai Mountain of China.Shenyang:Liao-ning Science and Technology Press,2004 (in Chinese)
[14] Du H-Y (杜洪业),Xu C-Y (徐程扬).Research process of the relationship between keystone species and the dynamic of community in forestry ecosystem.World Forestry Research (世界林业研究),2012,25(5):35-39 (in Chinese)
[15] Sun X-C (孙晓程),Zhang Y-Z (张亚洲),Fan X-N (范晓男).The status and operating management of broad leaved Pinus koriensis forests in Changbai Mountain.Journal of Jilin Normal University (Natural Science) (吉林师范大学学报:自然科学版),2008,29(2):105-107 (in Chinese)
[16] Yu D-P (于大炮),Zhou W-M (周旺明),Bao Y (包也),et al.Forest management of Korean pine and broadleaf mixed forest in Northeast China since the implementation of Natural Forest Protection Project.Acta Ecologica Sinica (生态学报),2015,35(1):10-17 (in Chinese)
[17] Hui GY,Wang Y,Zhang GQ,et al.A novel approach for assessing the neighborhood competition in two diffe-rent aged forests.Forest Ecology and Management,2018,422:49-58
[18] Wang L-W (王利伟),Li B-H (李步杭),Ye J (叶吉),et al.Dynamics of short-term tree mortality in broad-leaved Korean pine (Pinus koraiensis) mixed forest in the Changbai Mountains.Biodiversity Science (生物多样性),2011,19(2):260-270 (in Chinese)
[19] Zhang ZC,Papaik MJ,Wang XG,et al.The effect of tree size,neighborhood competition and environment on tree growth in an old-growth temperate forest.Journal of Plant Ecology,2016,10:970-980
[20] Hegyi F.A simulation model for managing jack-pine stands// Fries J,ed.Growth Models for Tree and Stand Simulation.Proceedings of IUFRO Meeting S4.01.04,Royal College of Forestry,Stockolm,1974:74-90
[21] Vanclay JK.Spatially-explicit competition indices and the analysis of mixed-species plantings with the Simile modelling environment.Forest Ecology and Management,2006,366:295-302
[22] Aho K,Derryberry DW,Peterson T.Model selection for ecologists:The worldviews of AIC and BIC.Ecology,2016,95:631-636
[23] Burnham KP,Anderson DR.Model Selection and Multimodal Inference:A Practical Information-theoretic Approach.2nd Ed.New York:Springer,2003
[24] Chao L (巢林),Liu Y-Y (刘艳艳),Zhang W-D (张伟东),et al.Diameter class structure,growth dynamic,intraspecific and interspecific competition of Betula platyphylla on swamp ecotone.Chinese Journal of Ecology (生态学杂志),2017,36(3):577-583 (in Chinese)
[25] Xu J (徐建),Wei X-L (韦新良),Wang J (王敬),et al.Intraspecific and interspecific competition of dominant species in a deciduous,broadleaf forest of Longwang Mountain.Journal of Zhejiang A&F University (浙江农林大学学报),2014,31(6):868-876 (in Chinese)
[26] Kang D,Deng J,Qin XW,et al.Effect of competition on spatial patterns of oak forests on the Chinese Loess Plateau.Journal of Arid Land,2017,9:122-131
[27] Zhang Z-C (张昭臣).Dynamics of Tree Mortality and Growth in Coniferous and Broad-leaved Mixed Forests in Changbai Mountain.PhD Thesis.Beijing:University of Chinese Academy of Sciences,2010 (in Chinese)
[28] Aleinikovas M,Edgaras L,Andrius K,et al.The impact of competition for growing space on diameter,basal area and height growth in pine trees.Baltic Fores-try,2014,20:301-313
[29] Huang JG,Stadt KJ,Dawson A,et al.Modelling growth-competition relationships in trembling aspen and white spruce mixed boreal forests of western Canada.PLoS One,2013,8(10):77607
[30] Han D-X (韩大校),Jin G-Z (金光泽).Influences of topography and competition on DBH growth in different growth stages in a typical mixed broadleaved-Korean pine forest,northeastern China.Journal of Beijing Forestry University (北京林业大学学报),2017,39(1):9-19 (in Chinese)
[31] Bella IE.A new competition model for individual trees.Forest Science,1971,17:364-372
[32] David SG,Zavala MA,Schalkwijk DB,et al.Rank reversals in tree growth along tree size,competition and climatic gradients for four forest canopy dominant species in Central Spain.Annals of Forest Science,2008,65:605
[33] Cai L-R (蔡礼荣),Kuang X (匡旭),Fang S (房帅),et al.Factors influencing tree radial growth of three common species in broad-leaved Korean pine mixed forests in Changbai Mountains,China.Chinese Journal of Applied ecology (应用生态学报),2017,28(5):1407-1413 (in Chinese)
[34] Macfarlane DW,Kobe RK.Selecting models for capturing tree-size effects on growth resource relationships.Canadian Journal of Forest Research,2006,36:1695-1704
[35] Pritchard JM,Comeau PG.Effects of opening size and stand characteristics on light transmittance and temperature under young trembling aspen stands.Forest Ecology and Management,2004,200:119-128
[36] Schwinning S,Weiner J.Mechanisms determining the degree of size asymmetry in competition among plants.Oecologia,1998,113:447-455
[37] Weiner J.Asymmetric competition in plant populations.Trends in Ecology & Evolution,1990,5:360-364
[38] Wyckoff PH,Clark JS.Tree growth prediction using size and exposed crown area.Canadian Journal of Forest Research,2005,35:13-20
[39] Gonzalez AE,Meakem V,Eng CY,et al.Patterns of tree mortality in a temperate deciduous forest derived from a large forest dynamics plot.Ecosphere,2016,7:e01595,doi:10.1002/ecs2.1595
[40] David AC,Robert BA.Mortality and tree-size distributions in natural mixed-age forests.Journal of Ecology,2007,95:27-40
[41] Tong X (童鑫).Exploring Community Assembly Through the Lens of Spatial Structure:From Population Genetics to Community Composition.PhD Thesis.Shanghai:East China Normal University,2015 (in Chinese)
[42] Zhang Z-C (张昭臣),Hao Z-Q (郝占庆),Ye J (叶吉),et al.Short-term death dynamics of trees in natural secondary poplar-birch forest in Changbai Mountains of Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2013,24(2):303-310 (in Chinese)
[43] Canham CD,Papaik MJ,Latty EF.Interspecific variation in susceptibility to windthrow as a function of tree size and storm severity for northern temperate tree species.Canadian Journal of Forest Research,2001,31:1-10
[44] Takahashi K.Effects of altitude and competition on growth and mortality of the conifer Abies sachalinensis.Ecological Research,2010,25:801-812