森林群落多样性与空间格局研究综述
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摘要
物种多样性和空间格局是过去自然过程和人类干扰的结果,也是群落未来发展的起点,是生态学理论研究的中心议题,也是应用研究的热点。文中对这2个方面进行了综述,指出存在的不足并提出了建议,以适应管理更复杂森林的需求。在群落物种多样性方面,针对物种的长期共存机制提出了各种不同的理论或假说,目前不同观点有整合趋势,并开始探讨不同机制在物种多样性维持方面的贡献;个体大小的多样性是多样性研究的一个新内容,最近开展了很多有关林木直径大小的研究。在空间格局方面,提出很多分析方法和检验判别统计量,在只需检验格局类型时扩散系数表现较好,而需探讨不同尺度格局类型时可以使用点格局分析。由于林木的高生长差异,多样性和格局在垂直方向有独特的表现,但是在当前研究中较少涉及垂直方向。今后应加强这方面研究,这对群落水平的天然林保护和森林近自然经营具有现实意义。
Species diversity and spatial pattern is the results of natural processes and human disturbances in the past,and also a starting point for its future community development. They are not only the central issue for ecology theory study,but also the focus of applied research. The paper reviewed the two aspects and also pointed out the existing problems with corresponding suggestions for meeting the needs to manage more complicated forest. Different theories or hypotheses have been put forward that concerning long-term coexistence of species in the community,and at present,some of them are to be integrated. In terms of species diversity of community,various theories or assumptions have been proposed for the long-term coexistence of species,but now different thoughts tends to be converged and the contributions by different mechanisms to species diversity maintenance began to be discussed. Size diversity is a new study content of the diversity of the community,and many studies have been conducted for tree diameter diversity. A number of analysis methods and statistics have been put forward for spatial pattern analysis,and diffusion coefficientis good for testing pattern types while point pattern analysis could be used to explore pattern types at different scales. Owing to the difference in height growth of trees,diversify and pattern have unique performance in vertical distribution. But there is little touched on vertical distribution in current studies,and therefore the future enhancement of the study on this aspect would be realistically significant for nature forest protection and close-to-nature forest management at community level.
Species diversity and spatial pattern is the results of natural processes and human disturbances in the past,and also a starting point for its future community development. They are not only the central issue for ecology theory study,but also the focus of applied research. The paper reviewed the two aspects and also pointed out the existing problems with corresponding suggestions for meeting the needs to manage more complicated forest. Different theories or hypotheses have been put forward that concerning long-term coexistence of species in the community,and at present,some of them are to be integrated. In terms of species diversity of community,various theories or assumptions have been proposed for the long-term coexistence of species,but now different thoughts tends to be converged and the contributions by different mechanisms to species diversity maintenance began to be discussed. Size diversity is a new study content of the diversity of the community,and many studies have been conducted for tree diameter diversity. A number of analysis methods and statistics have been put forward for spatial pattern analysis,and diffusion coefficientis good for testing pattern types while point pattern analysis could be used to explore pattern types at different scales. Owing to the difference in height growth of trees,diversify and pattern have unique performance in vertical distribution. But there is little touched on vertical distribution in current studies,and therefore the future enhancement of the study on this aspect would be realistically significant for nature forest protection and close-to-nature forest management at community level.
引文
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[22]BUONGIORNO J,DAHIR S,LU H C,et al.Tree size diversity and economic returns in uneven-aged forest stands[J].Forest Science,1994,40(1):83-103.
[23]LIANG J,BUONGIORNO J,MONSERUD R A,et al.Effects of diversity of tree species and size on forest basal area growth,recruitment,and mortality[J].Forest Ecology and Management,2007,243(1):116-127.
[24]YOUNG B,LIANG J J,CHAPIN III F S.Effects of species and tree size diversity on recruitment in the Alaskan boreal forest:a geospatial approach[J].Forest Ecology and Management,2011,262(8):1608-1617.
[25]LEXERD N L,EID T.An evaluation of different diameter diversity indices based on criteria related to forest management planning[J].Forest Ecology and Management,2006,222(1):17-28.
[26]雷相东,唐守正.林分结构多样性指标研究综述[J].林业科学,2002,38(3):140-146.
[27]HUI G,POMMERENING A.Analysing tree species and size diversity patterns in multi-species uneven-aged forests of Northern China[J].Forest Ecology and Management,2014,316(2):125-138.
[28]FERRIS-KAAN R,PATTERSON G S.Monitoring vegetation changes in conservation management of forests:Forestry Commission bulletin108[M].London,UK:HMSO,1992.
[29]MACARTHUR R H,MACARTHUR J W.On bird species diversity[J].Ecology,1961,42(3):594-598.
[30]MURDOCH W W,EVANS F C,PETERSON C H.Diversity and pattern in plants and insects[J].Ecology,1972,53(5):819-829.
[31]TANABE S I,TODA M J,VINOKUROVA A V.Tree shape,forest structure and diversity of drosophilid community:comparison between boreal and temperate birch forests[J].Ecological Research,2001,16(3):369-385.
[32]KUULUVAINEN T,PENTTINEN A,LEINONEN K,et al.Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests:an example from boreal spruce forest in southern Finland[J].Silva Fennica,1996,30(2/3):315-328.
[33]ZENNER E K,HIBBS D E.A new method for modeling the heterogeneity of forest structure[J].Forest Ecology and Management,2000,129(1):75-87.
[34]CONDIT R,ASHTON P S,BAKER P,et al.Spatial patterns in the distribution of tropical tree species[J].Science,2000,288(5470):1414-1418.
[35]惠刚盈,克劳斯·冯佳多.森林空间结构量化分析方法[M].北京:中国科学技术出版社,2003.
[36]马钦彦.负二项分布及种群格局检验分析[J].北京林业大学学报,2009,31(3):1-5.
[37]NEYMAN J.On a new class of"contagious"distributions,applicable in entomology and bacteriology[J].Annual Math Statistics,1939,10(1):35-57.
[38]张金屯.数量生态学[M].北京:科学出版社,2004:77-94.
[39]FORTIN M J,DALE M.Spatial analysis:a guide for ecologists[M].Cambridge,UK:Cambridge University,2005.
[40]BADDELEY A,GREGORI P,Mateu J,et al.Case studies in point process modeling[M].New York,USA:Springer Science+Business Media Inc.,2006:3-21,275.
[41]WIEGAND T,MOLONEY K A.Rings,circles,and null-models for point pattern analysis in ecology[J].Oikos,2004,104(2):209-229.
[42]PERRY G L W,MILLER B P,ENRIGHT N J.A comparison of methods for the statistical analysis of spatial point patterns in plant ecology[J].Plant Ecology,2006,187(1):59-82.
[43]MCGILL B J,ETIENNE R S,GRAY J S,et al.Species abundance distributions:moving beyond single prediction theories to integration within an ecological framework[J].Ecology letters,2007,10(10):995-1015.
[44]RIPLEY B D.The second-order analysis of stationary point processes[J].Journal of Applied Probability,1976,13(2):255-266.
[45]BESAG J.Contribution to the discussion of Dr.Ripley’s paper[J].Journal of the Royal Statistical Society(Series B),1977,39(2):193-195.
[46]DIXON P M.Ripley’s K function[M]//EL-SHAARAWI A H,PIEGORSCH W W(eds).Encyclopedia of environmetrics:vol.3.Chichester,UK:John Wiley&Sons,2002:1796-1803.
[47]ZHANG C Y,JIN W B,GAO L S,et al.Scale dependent structuring of spatial diversity in two temperate forest communities[J].Forest Ecology and Management,2014,316(2):110-116.
[48]魏彦波,程艳霞,李金功,等.植物多样性促进种支配局域空间多样性结构[J].北京林业大学学报,2014,36(6):66-72.
[49]范春雨,元正龙,赵秀海.吉林蛟河近熟林树种多样性格局尺度依赖性分析[J].北京林业大学学报,2014,36(6):73-79.
[2]方精云.群落生态学迎来新的辉煌时代[J].生物多样性,2009,17(6):531-532.
[3]Von GADOW K,ZHANG C Y,WEHENKEL C.Forest structure and diversity[M]//PUKKALA T,Von GADOW K(eds).Continuous cover forestry.2nd edition.London,UK:Springer Science&Business Media,2011:30-33.
[4]NOWAK M A,SIGMUND J.Evolutionary dynamics of biological games[J].Science,2004,303(5659):793-799.
[5]CHEN S Y,ZHANG J L,JIA P,et al.Effects of size variation and spatial structure on plastic response of plant height to light competition[J].Chinese Science Bulletin,2010,55(12):1135-1141.
[6]HUBBELL S P.Neutral theory and the evolution of ecological equivalence[J].Ecology,2006,87(6):1387-1398.
[7]CHASE J M.Towards a really unified theory for metacommunities[J].Functional Ecology,2005,19(1):182-186.
[8]周淑荣,张大勇.群落生态学的中性理论[J].植物生态学报,2006,30(5):868-877.
[9]WIEGAND T,GUNATILLEKE C V S,GUNATILLEKE I A U,et al.How individual species structure diversity in tropical forests[J].PNAS,2007,104(48):19029-19033.
[10]牛克昌,刘怿宁,沈泽昊,等.群落构建中的中性理论和生态位理论[J].生物多样性,2009,17(6):579-593.
[11]PAVOINE S,BONSALL M B.Measuring biodiversity to explain community assembly:a unified approach[J].Biological Reviews,2011,86(4):792-812.
[12]RICKLEFS R E.Disintegration of the ecological community[J].American Naturalist,2008,172(6):741-750.
[13]方精云,王襄平,唐志尧.局域和区域过程共同控制着群落的物种多样性:种库假说[J].生物多样性,2009,17(6):605-612.
[14]TILMAN D.Niche tradeoffs,neutrality,and community structure:a stochastic theory of resource competition,invasion and community assembly[J].PNAS,2004,101(30):10854-10861.
[15]GRAVEL D,CANHAM C D,BEAUDET M,et al.Reconciling niche and neutrality:the continuum hypothesis[J].Ecology Letters,2006,9(4):399-409.
[16]LANDE R.Statistics and partitioning of species diversity,and similarity among multiple communities[J].Oikos,1996,76(1):5
[17]徐洋,刘文治,刘贵华.生态位限制和物种库限制对湖滨湿地植物群落分布格局的影响[J].植物生态学报,2009,33(3):546-554.
[18]袁秀,马克明,王德.黄河三角洲植物生态位和生态幅度对物种分布—多度关系的解释[J].生物多样性,2012,20(2):159-167.
[19]KUNSTLER G,LAVERGNE S,COURBAUD B,et al.Competitive interactions between forest trees are driven by species'trait hierarchy,not phylogenetic or functional similarity:implications for forest community assembly[J].Ecology Letters,2012,15(8):831-840.
[20]GROSS K.Positive interactions among competitors can produce species-rich communities[J].Ecology Letters,2008,11(9):929-936.
[21]张炜平,潘莎,贾昕,等.植物间正相互作用对种群动态和群落结构的影响:基于个体模型的研究进展[J].植物生态学报,2013,37(6):571-582.
[22]BUONGIORNO J,DAHIR S,LU H C,et al.Tree size diversity and economic returns in uneven-aged forest stands[J].Forest Science,1994,40(1):83-103.
[23]LIANG J,BUONGIORNO J,MONSERUD R A,et al.Effects of diversity of tree species and size on forest basal area growth,recruitment,and mortality[J].Forest Ecology and Management,2007,243(1):116-127.
[24]YOUNG B,LIANG J J,CHAPIN III F S.Effects of species and tree size diversity on recruitment in the Alaskan boreal forest:a geospatial approach[J].Forest Ecology and Management,2011,262(8):1608-1617.
[25]LEXERD N L,EID T.An evaluation of different diameter diversity indices based on criteria related to forest management planning[J].Forest Ecology and Management,2006,222(1):17-28.
[26]雷相东,唐守正.林分结构多样性指标研究综述[J].林业科学,2002,38(3):140-146.
[27]HUI G,POMMERENING A.Analysing tree species and size diversity patterns in multi-species uneven-aged forests of Northern China[J].Forest Ecology and Management,2014,316(2):125-138.
[28]FERRIS-KAAN R,PATTERSON G S.Monitoring vegetation changes in conservation management of forests:Forestry Commission bulletin108[M].London,UK:HMSO,1992.
[29]MACARTHUR R H,MACARTHUR J W.On bird species diversity[J].Ecology,1961,42(3):594-598.
[30]MURDOCH W W,EVANS F C,PETERSON C H.Diversity and pattern in plants and insects[J].Ecology,1972,53(5):819-829.
[31]TANABE S I,TODA M J,VINOKUROVA A V.Tree shape,forest structure and diversity of drosophilid community:comparison between boreal and temperate birch forests[J].Ecological Research,2001,16(3):369-385.
[32]KUULUVAINEN T,PENTTINEN A,LEINONEN K,et al.Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests:an example from boreal spruce forest in southern Finland[J].Silva Fennica,1996,30(2/3):315-328.
[33]ZENNER E K,HIBBS D E.A new method for modeling the heterogeneity of forest structure[J].Forest Ecology and Management,2000,129(1):75-87.
[34]CONDIT R,ASHTON P S,BAKER P,et al.Spatial patterns in the distribution of tropical tree species[J].Science,2000,288(5470):1414-1418.
[35]惠刚盈,克劳斯·冯佳多.森林空间结构量化分析方法[M].北京:中国科学技术出版社,2003.
[36]马钦彦.负二项分布及种群格局检验分析[J].北京林业大学学报,2009,31(3):1-5.
[37]NEYMAN J.On a new class of"contagious"distributions,applicable in entomology and bacteriology[J].Annual Math Statistics,1939,10(1):35-57.
[38]张金屯.数量生态学[M].北京:科学出版社,2004:77-94.
[39]FORTIN M J,DALE M.Spatial analysis:a guide for ecologists[M].Cambridge,UK:Cambridge University,2005.
[40]BADDELEY A,GREGORI P,Mateu J,et al.Case studies in point process modeling[M].New York,USA:Springer Science+Business Media Inc.,2006:3-21,275.
[41]WIEGAND T,MOLONEY K A.Rings,circles,and null-models for point pattern analysis in ecology[J].Oikos,2004,104(2):209-229.
[42]PERRY G L W,MILLER B P,ENRIGHT N J.A comparison of methods for the statistical analysis of spatial point patterns in plant ecology[J].Plant Ecology,2006,187(1):59-82.
[43]MCGILL B J,ETIENNE R S,GRAY J S,et al.Species abundance distributions:moving beyond single prediction theories to integration within an ecological framework[J].Ecology letters,2007,10(10):995-1015.
[44]RIPLEY B D.The second-order analysis of stationary point processes[J].Journal of Applied Probability,1976,13(2):255-266.
[45]BESAG J.Contribution to the discussion of Dr.Ripley’s paper[J].Journal of the Royal Statistical Society(Series B),1977,39(2):193-195.
[46]DIXON P M.Ripley’s K function[M]//EL-SHAARAWI A H,PIEGORSCH W W(eds).Encyclopedia of environmetrics:vol.3.Chichester,UK:John Wiley&Sons,2002:1796-1803.
[47]ZHANG C Y,JIN W B,GAO L S,et al.Scale dependent structuring of spatial diversity in two temperate forest communities[J].Forest Ecology and Management,2014,316(2):110-116.
[48]魏彦波,程艳霞,李金功,等.植物多样性促进种支配局域空间多样性结构[J].北京林业大学学报,2014,36(6):66-72.
[49]范春雨,元正龙,赵秀海.吉林蛟河近熟林树种多样性格局尺度依赖性分析[J].北京林业大学学报,2014,36(6):73-79.