秦岭辛家山不同森林类型土壤养分及植物多样性研究
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摘要
森林是全球生态系统非常重要的组成部分,具有调节气候、涵养水源、净化空气、防风固沙等多种功能,对维护地球生态平衡和人类的生存、发展至关重要。
秦岭―黄河、长江两大水系的分水岭,也是中国南北气候的分界线。明显的垂直地带性植被带和土壤类型孕育了秦岭丰富、独特的植被类型。辛家山,位于秦岭西部南坡,嘉陵江上游,北依秦岭主脊,地形地貌波状起伏,具有既复杂又规律的特征。有利于进行植物群落多样性形成、森林结构类型特征以及土壤养分状况变化规律的相关研究。本研究于2008年6月至2010年8月,对秦岭辛家山林区9种不同森林类型(白桦林、水曲柳林、锐齿栎林、油松-水曲柳混交林、云杉-锐齿栎混交林、油松-千金榆混交林、油松林、华山松、云杉林)土壤及地上植被群落进行调查,在此基础上,定量分析了辛家山林区的土壤化学性质及其与不同森林类型之间的关系、不同深度的变化规律、群落物种多样性等。结果表明:
(1)辛家山林区不同森林类型之间土壤养分、pH值都存在显著差异。有机质含量都呈现表层土壤(0~20cm)含量高于中、下层,且随着土层加深有机质含量递减的变化趋势。9种森林类型中,锐齿栎和云杉林下土壤有机质含量较高;云杉林土壤全氮含量(2.97g/kg)、碱解氮含量(257.61mg/kg)均是9种森林中为最高;油松林土壤全磷含量最高,是华山松林的5.64倍,水曲柳林土壤有效磷含量最高;华山松林土壤各项指标均最低。
(2)9种不同森林类型土壤三个土层(0~20cm、20~40cm、40~60cm)在垂直变化上也存在差异。其中,油松-水曲柳混交林表层土壤有机质含量比下层的高出1.39倍,相差最大,相差最小的白桦林土壤表层有机质含量比下层高出了14%;锐齿栎林土壤表层全氮含量比下层高了44%;油松-千金榆混交林土壤有效磷含量递减幅度最大,达到了24%。
(3)该区9种森林类型土壤pH值介于4.75~6.22之间,土壤pH多表现为表层低于下层。
(4)相关分析表明:土壤全氮与有机质、碱解氮、全磷含量达到极显著正相关;碱解氮与有机质、全磷显著正相关;pH与全氮显著负相关;土壤有效磷含量与土壤全磷量相关性不显著。
(5)Pielou指数排序为:云杉+锐齿栎林>油松+水曲柳林>锐齿栎林>白桦林>油松林>云杉林>油松+千金榆>水曲柳林>华山松林;Simpson指数(D)的顺序为:云杉+锐齿栎林>油松+水曲柳林>锐齿栎林>白桦林>油松林>油松-千金榆混交林>云杉林>水曲柳林>华山松林;Shannon-Wiener指数(H)的顺序为:云杉+锐齿栎林>白桦林>云杉林>锐齿栎林>油松+水曲柳林>水曲柳林>油松+千金榆>油松林>华山松林。
(6)本研究所选取的6个土壤因子中,除了pH与植物多样性没有相关性外,其余多成显著相关。
The forest is the important constituent of global ecosystem which has the functions of adjusting the climate, keeping waterhead, purify air, wind-breaking and sand-fixation. It plays an important role in keeping the ecological balance of the earth. It has the vital significance on survival and development of mankind.
Qinling Mountain, the watershed of two major river systems: Yangtze River and Yellow River, is north-south clime boundary of China. The complicated features and varied weather bless this region with advantageous conditions of unique and rich vegetation.
Xinjiashan is located in the south slope of western Qinling and upstream of Jialing river. Next to Qinling major ridge, it has both complicated and regular features, which is helpful to understand forest structure characteristics, biodiversity formation and soil nutrient status. The research investigated 9 different typical forest communities and soil under them in Xinjiashan from June 2008 to September 2010. On this basis, quantitative analysis was done to study the relations between soil fertility and community type, soil depth, diversity of species. The results show that:
(1)The soil nutrient and pH among different forest types were significant different in Xinjiashan. The soil organic matter content had a certain vertical variation, but the maximal value occurs in the surface soil laye(r0~20cm)in most cases. Among 9 different typical forest communities, the organic matter content under Picea asperata and Quercus aliena were higher than others. The soil under Picea asperata had the most total nitrogen and alkali-hydro nitrogen. The total phosphorus content of Pinus tabulaeformis was highest, while the available phosphorus content of Fraxinus mandshurica was the most. Differently, the various indices of soil under Pinus armandii were relatively low.
(2) Soil nutrient was significantly different in three soil layer (0 ~ 20cm, 20 ~ 40cm, 40 ~ 60cm) under different forest types. For instance, the organic matter content of surface soil under Pinus tabulaeformis and Fraxiinus mandshurica mixed forest was 1.39 times higher than the lower. The total nitrogen content of surface soil under Quercus aliena was 44% higher than the lower. The Pinus tabulaeformis and Carpinus cordata mixed forest reduced 24% with depth.
(3)The soil pH of nine forest types was between 6.22 and 4.75. And soil pH in the surface layer was higher than the sublayer. The content of soil organic matter in the surface layer was higher than the sublayer. The soil total nutrient under Pinus armandii was the lowest in 9 forest types.
(4)Correlation analysis showed that total nitrogen, the contents of organic matter, alkali-hydro nitrogen and total phosphorus in different types of forests were very significant positive correlated; alkali-hydro nitrogen, the contents of organic matter and total phosphorus were significant positive correlated; soil pH and total nitrogen were very significant negative correlated; the correlation between available phosphorus and total nitrogen was not significant.
(5) Pielou index ranking for: Picea asperata Mast.-Quercus aliena var. > Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr.> Quercus aliena var. acuteserrata.> Betula platyphylla Suk.> Pinus tabulaeformis Carr.> Picea asperata Mast. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Fraxinus mandschurica Rupr. > Pinus armandii Franch. ;
Simpson index (D) of this order: Picea asperata Mast. -Quercus aliena var. acuteserrata> Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr.> Quercus aliena var. acuteserrata.> Betula platyphylla Suk.> Pinus tabulaeformis Carr.> Picea asperata Mast. > Fraxinus mandschurica Rupr. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Pinus armandii Franch.
Shannon - Wiener index (H) of this order: Picea asperata Mast. -Quercus aliena var. acuteserrata.> Betula platyphylla Suk. > Picea asperata Mast. > Quercus aliena var. acuteserrata > Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr. > Fraxinus mandschurica Rupr. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Pinus tabulaeformis Carr. > Pinus armandii Franch..
(6) Most of soil factors studied were significantly correlated with species diversity except pH.
秦岭―黄河、长江两大水系的分水岭,也是中国南北气候的分界线。明显的垂直地带性植被带和土壤类型孕育了秦岭丰富、独特的植被类型。辛家山,位于秦岭西部南坡,嘉陵江上游,北依秦岭主脊,地形地貌波状起伏,具有既复杂又规律的特征。有利于进行植物群落多样性形成、森林结构类型特征以及土壤养分状况变化规律的相关研究。本研究于2008年6月至2010年8月,对秦岭辛家山林区9种不同森林类型(白桦林、水曲柳林、锐齿栎林、油松-水曲柳混交林、云杉-锐齿栎混交林、油松-千金榆混交林、油松林、华山松、云杉林)土壤及地上植被群落进行调查,在此基础上,定量分析了辛家山林区的土壤化学性质及其与不同森林类型之间的关系、不同深度的变化规律、群落物种多样性等。结果表明:
(1)辛家山林区不同森林类型之间土壤养分、pH值都存在显著差异。有机质含量都呈现表层土壤(0~20cm)含量高于中、下层,且随着土层加深有机质含量递减的变化趋势。9种森林类型中,锐齿栎和云杉林下土壤有机质含量较高;云杉林土壤全氮含量(2.97g/kg)、碱解氮含量(257.61mg/kg)均是9种森林中为最高;油松林土壤全磷含量最高,是华山松林的5.64倍,水曲柳林土壤有效磷含量最高;华山松林土壤各项指标均最低。
(2)9种不同森林类型土壤三个土层(0~20cm、20~40cm、40~60cm)在垂直变化上也存在差异。其中,油松-水曲柳混交林表层土壤有机质含量比下层的高出1.39倍,相差最大,相差最小的白桦林土壤表层有机质含量比下层高出了14%;锐齿栎林土壤表层全氮含量比下层高了44%;油松-千金榆混交林土壤有效磷含量递减幅度最大,达到了24%。
(3)该区9种森林类型土壤pH值介于4.75~6.22之间,土壤pH多表现为表层低于下层。
(4)相关分析表明:土壤全氮与有机质、碱解氮、全磷含量达到极显著正相关;碱解氮与有机质、全磷显著正相关;pH与全氮显著负相关;土壤有效磷含量与土壤全磷量相关性不显著。
(5)Pielou指数排序为:云杉+锐齿栎林>油松+水曲柳林>锐齿栎林>白桦林>油松林>云杉林>油松+千金榆>水曲柳林>华山松林;Simpson指数(D)的顺序为:云杉+锐齿栎林>油松+水曲柳林>锐齿栎林>白桦林>油松林>油松-千金榆混交林>云杉林>水曲柳林>华山松林;Shannon-Wiener指数(H)的顺序为:云杉+锐齿栎林>白桦林>云杉林>锐齿栎林>油松+水曲柳林>水曲柳林>油松+千金榆>油松林>华山松林。
(6)本研究所选取的6个土壤因子中,除了pH与植物多样性没有相关性外,其余多成显著相关。
The forest is the important constituent of global ecosystem which has the functions of adjusting the climate, keeping waterhead, purify air, wind-breaking and sand-fixation. It plays an important role in keeping the ecological balance of the earth. It has the vital significance on survival and development of mankind.
Qinling Mountain, the watershed of two major river systems: Yangtze River and Yellow River, is north-south clime boundary of China. The complicated features and varied weather bless this region with advantageous conditions of unique and rich vegetation.
Xinjiashan is located in the south slope of western Qinling and upstream of Jialing river. Next to Qinling major ridge, it has both complicated and regular features, which is helpful to understand forest structure characteristics, biodiversity formation and soil nutrient status. The research investigated 9 different typical forest communities and soil under them in Xinjiashan from June 2008 to September 2010. On this basis, quantitative analysis was done to study the relations between soil fertility and community type, soil depth, diversity of species. The results show that:
(1)The soil nutrient and pH among different forest types were significant different in Xinjiashan. The soil organic matter content had a certain vertical variation, but the maximal value occurs in the surface soil laye(r0~20cm)in most cases. Among 9 different typical forest communities, the organic matter content under Picea asperata and Quercus aliena were higher than others. The soil under Picea asperata had the most total nitrogen and alkali-hydro nitrogen. The total phosphorus content of Pinus tabulaeformis was highest, while the available phosphorus content of Fraxinus mandshurica was the most. Differently, the various indices of soil under Pinus armandii were relatively low.
(2) Soil nutrient was significantly different in three soil layer (0 ~ 20cm, 20 ~ 40cm, 40 ~ 60cm) under different forest types. For instance, the organic matter content of surface soil under Pinus tabulaeformis and Fraxiinus mandshurica mixed forest was 1.39 times higher than the lower. The total nitrogen content of surface soil under Quercus aliena was 44% higher than the lower. The Pinus tabulaeformis and Carpinus cordata mixed forest reduced 24% with depth.
(3)The soil pH of nine forest types was between 6.22 and 4.75. And soil pH in the surface layer was higher than the sublayer. The content of soil organic matter in the surface layer was higher than the sublayer. The soil total nutrient under Pinus armandii was the lowest in 9 forest types.
(4)Correlation analysis showed that total nitrogen, the contents of organic matter, alkali-hydro nitrogen and total phosphorus in different types of forests were very significant positive correlated; alkali-hydro nitrogen, the contents of organic matter and total phosphorus were significant positive correlated; soil pH and total nitrogen were very significant negative correlated; the correlation between available phosphorus and total nitrogen was not significant.
(5) Pielou index ranking for: Picea asperata Mast.-Quercus aliena var. > Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr.> Quercus aliena var. acuteserrata.> Betula platyphylla Suk.> Pinus tabulaeformis Carr.> Picea asperata Mast. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Fraxinus mandschurica Rupr. > Pinus armandii Franch. ;
Simpson index (D) of this order: Picea asperata Mast. -Quercus aliena var. acuteserrata> Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr.> Quercus aliena var. acuteserrata.> Betula platyphylla Suk.> Pinus tabulaeformis Carr.> Picea asperata Mast. > Fraxinus mandschurica Rupr. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Pinus armandii Franch.
Shannon - Wiener index (H) of this order: Picea asperata Mast. -Quercus aliena var. acuteserrata.> Betula platyphylla Suk. > Picea asperata Mast. > Quercus aliena var. acuteserrata > Pinus tabulaeformis Carr. -Fraxinus mandshurica Rupr. > Fraxinus mandschurica Rupr. > Pinus tabulaeformis Carr. - Carpinus cordata Bl. > Pinus tabulaeformis Carr. > Pinus armandii Franch..
(6) Most of soil factors studied were significantly correlated with species diversity except pH.
引文
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杨承栋. 1991.对我国立地分类与评价问题的几点看法.林业科学, 27(1): 60~64
杨承栋.2008.森林土壤学科研究进展与展望.土壤学报,45(5):881~890
杨承栋.论合理保护开发利用中国森林土壤资源.世界林业研究,24(1)19~26
杨小波,张桃林,吴庆书. 2002.海南琼北地区不同植被类型物种多样性与土壤肥力的关系.生态学报, 22(2):190~196
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游秀花,蒋尔可. 2005.不同森林类型土壤化学性质的比较研究.江西农业大学学报, 27(3):357~360.
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张万儒主编. 1997.中国森林立地.北京:科学出版社
张兴义,王树奎,隋跃宇. 2005.东北农田黑土碱解氮现状评价.农业系统科学与综合研究, 21(4):305~309,1581~158 8
周华荣,肖笃宁,周可法.2006.干旱区景观格局空间过程变化的廊道效应——以塔里木河中下游河流廊道区域为例.科学通报,(S1):66~72
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Chapin FS,Zavaleta ES,Eviner VT. 2000.Consequences of changing biodiversity. Nature, 405,234~242 Costanza R D,Arge R,De Groot R. 1997. The value of the world's ecosystemservices and natural capital. Nature, 387:253~260
Du XG,Zhou SR. 2008.Testing the neutral theory of plant communities in subalpine meadow. Journal of Plant Ecology, 32(2):347~354
Donald M. Stoeekel and Mary S. Miller-Goodman,Seasonal Nutrient Dynamics of Forested FloodPlain soil Influenced by MierotoPograPhy and Depth.2001.soil Sci. Soc Am.J,65:922～931.
Elton CS.1958. Ecology of Invasions by Animal and Plant. London:Chapman and Hall.50~72
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Fttier A.H.2005.Darkness visbile: reflections on underground ecology Jurnal of Eeology,93,231～43
Godfray H.C.J and Laeton J.H.2001.Seale and species numbers.Trends in Eeology and Evolution,16:400～404
Hook R B, I C Burke, W K Lauenroth. 1991. Heterogeneity of soil and Plant N and C associated within dividual Plants and openings in North American short grass steppe. Plant and soil, 138:247～256
Gahoonia T S,Nielsen N E. 1992.The effect of too-inducedpH changes on the depletion of inorganic and organicphosphorus in the rhizophere.Plant and soil, 143:185~191
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HarrisonS. 1996.Loealand regional diversity in a partly landscape:native,alien and endemic herbs on serpentine.Eeology,76(2):95~126
HartlletDC,WilsonWT. 1999.Myeorrhizae influence Plant community structure and diversity Intallgr as sprairie.Eeology,80(4):1187～1195
Hou JH,Ma KP. 2002.On mechanisms of species coexistence in plant community.Journal of Plant Ecology, 26:1~8
Kellogg L E,Bridgham S D. 2003.Phosphorus retention andmovement across an ombrotrophic-minerot rophic peat-land gradient.Biogeochemistry, 52:299~315
Lajtha K,Schlesinger W H. 1998.The biochemistry of phosphorus cycling and phosphorus availabilityalong a desert soil chronosequence.Ecology, 69:24~39
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LiebigJ.Die orgnische Chemie in ihrerAnwendung auf Agricultur und Physiologie(Organic Ehemistry in it applications to agriculture and Physiology)
MacArthur RH. 1955. Fluctuations of animal populations and a measure of community stability. Ecology, 36:533~536
Magerran,AE.2001.Ecological diversity and its Measurement.NewJersey:Princeton May RM. 1975. Ecology and Evolution of Communities.Harvard University Press, Cambridge,81~120
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Odum EP.1953.Fundamentals of Ecology. Philadelphia:Saunders,132~157
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熊毅,李庆逵. 1998.中国土壤(第二版).北京:科学出版社:22~24
许广山,张玉华,刘春萍. 1997.长白山阔叶红松林养分循环研究.森林与土壤.北京:中国科学技术出版社, 48~53
薛敬意,唐建维,沙丽清. 2003.西双版纳望天树林土壤养分含量及其季节变化.植物生态学报, 27(3):373~379
薛立,邝立刚,陈红跃. 2003.不同林分土壤养分、微生物与酶活性的研究.土壤学报, 40(2):280~285
杨承栋. 1991.对我国立地分类与评价问题的几点看法.林业科学, 27(1): 60~64
杨承栋.2008.森林土壤学科研究进展与展望.土壤学报,45(5):881~890
杨承栋.论合理保护开发利用中国森林土壤资源.世界林业研究,24(1)19~26
杨小波,张桃林,吴庆书. 2002.海南琼北地区不同植被类型物种多样性与土壤肥力的关系.生态学报, 22(2):190~196
应俊生,李云峰,郭勤峰. 1990.秦岭太白山地区的植物区系和分类.植物分类学报, 28:261~293
应俊生. 1994.秦岭植物区系的特性、特点和起源.植物分类学报,32:389~410
游秀花,蒋尔可. 2005.不同森林类型土壤化学性质的比较研究.江西农业大学学报, 27(3):357~360.
俞元春. 1998.衫木、马尾松,甜储等林分卜土壤养分状况研究[J1.林业科学研究, 11(6):586~591
张秉刚,康慕谊. 1985.鼎湖山自然保护区不同林型下土壤的物理性质.热带亚热带森林生态系统研究, (3):1~10
张秉刚,康慕谊. 1990.亚热带季风常绿阔叶林不同林型下土壤水分状况的研究.热带亚热带森林生态系统研究, (6):75~82
张江英,周华荣,高梅. 2007.白杨河—艾里克湖湿地及周边植物群落与环境因子的关系.干旱区地理,17(6):101~107
张万儒,杨承栋. 1990.卧龙自然保护区森林土壤养分状况.土壤通报, 21(3):97~102
张万儒主编. 1997.中国森林立地.北京:科学出版社
张兴义,王树奎,隋跃宇. 2005.东北农田黑土碱解氮现状评价.农业系统科学与综合研究, 21(4):305~309,1581~158 8
周华荣,肖笃宁,周可法.2006.干旱区景观格局空间过程变化的廊道效应——以塔里木河中下游河流廊道区域为例.科学通报,(S1):66~72
朱祖祥. 1983.土壤学.北京:中国农业出版社: 68~75
Bowen G D, NambiarA G. 1989. Future direction in plantation nutrition research. In: Bowen G D, Nambiar E K S. Nutrition of Plantation Forest. London:Academic Press, 489~505
Braunschweig,Germany.1840. Friedrich Vieweg und Sohn Publ.Co. CarmeanW H. Forest site quality evaluation in theUnite State.Adv. Agron., 1975, (27): 254~255
Chapin FS,Zavaleta ES,Eviner VT. 2000.Consequences of changing biodiversity. Nature, 405,234~242 Costanza R D,Arge R,De Groot R. 1997. The value of the world's ecosystemservices and natural capital. Nature, 387:253~260
Du XG,Zhou SR. 2008.Testing the neutral theory of plant communities in subalpine meadow. Journal of Plant Ecology, 32(2):347~354
Donald M. Stoeekel and Mary S. Miller-Goodman,Seasonal Nutrient Dynamics of Forested FloodPlain soil Influenced by MierotoPograPhy and Depth.2001.soil Sci. Soc Am.J,65:922～931.
Elton CS.1958. Ecology of Invasions by Animal and Plant. London:Chapman and Hall.50~72
Doran J W, Parkin T B. 1994. Defining Soil Quality for Sustainable Environment. Madison Wisconsin: Soil Sci SocAm Spec Publ.35:321～324
Fttier A.H.2005.Darkness visbile: reflections on underground ecology Jurnal of Eeology,93,231～43
Godfray H.C.J and Laeton J.H.2001.Seale and species numbers.Trends in Eeology and Evolution,16:400～404
Hook R B, I C Burke, W K Lauenroth. 1991. Heterogeneity of soil and Plant N and C associated within dividual Plants and openings in North American short grass steppe. Plant and soil, 138:247～256
Gahoonia T S,Nielsen N E. 1992.The effect of too-inducedpH changes on the depletion of inorganic and organicphosphorus in the rhizophere.Plant and soil, 143:185~191
Gose JR. 1989. Biollgical factors influencing nutrient supply in forest soils.In: Bowen G D, Nambiar E K S. Nutrition of Plantation Forests. London:Academic Press, 119~146
HarrisonS. 1996.Loealand regional diversity in a partly landscape:native,alien and endemic herbs on serpentine.Eeology,76(2):95~126
HartlletDC,WilsonWT. 1999.Myeorrhizae influence Plant community structure and diversity Intallgr as sprairie.Eeology,80(4):1187～1195
Hou JH,Ma KP. 2002.On mechanisms of species coexistence in plant community.Journal of Plant Ecology, 26:1~8
Kellogg L E,Bridgham S D. 2003.Phosphorus retention andmovement across an ombrotrophic-minerot rophic peat-land gradient.Biogeochemistry, 52:299~315
Lajtha K,Schlesinger W H. 1998.The biochemistry of phosphorus cycling and phosphorus availabilityalong a desert soil chronosequence.Ecology, 69:24~39
Levins R. 1970.Extinction some mathematical questions in biology. In : Gerstenhaber M ed. Lectures on Mathematics in the Life Sciences, 2:75~107
LiebigJ.Die orgnische Chemie in ihrerAnwendung auf Agricultur und Physiologie(Organic Ehemistry in it applications to agriculture and Physiology)
MacArthur RH. 1955. Fluctuations of animal populations and a measure of community stability. Ecology, 36:533~536
Magerran,AE.2001.Ecological diversity and its Measurement.NewJersey:Princeton May RM. 1975. Ecology and Evolution of Communities.Harvard University Press, Cambridge,81~120
Mead D J. 1989. Diagnosis of nutrient deficiencies in plantation.In:Bowen G D, Nambiar E K S. Nutrition of Plantation Forests.London:Academic Press, 259~291
Odum EP.1953.Fundamentals of Ecology. Philadelphia:Saunders,132~157
Pacala SW,Tilman D. 1993.Limiting similarity in mechanistic and spatial models of plant competition in heterogeneous environments.The American Naturalist, 143:222~257
RboertsonGP,VitowsekPM.1981. Nitrification potentials in primary and Secondary Secondary succession. Ecolgy, 62:376 ~ 386
Reynolds H L. 1997.Soil heterogeneity and plant competition in an annual grassland.Eeology,78(7):2076~ 2090
Stohlgren TJ.. 1998.Comparison of range land vegetation sampling techaiques in theCentralGrasslands.Range.Manage,51(2):164~ 172
TilmanD. 1994.Competition and biodiversity in spatially struetured habitats.Eeology,75(l). TilmanD.PaealaS.1993.TheMaintenanceofsPeeiesriehnessinPlantcommunities.InRichlefs RE. Species diversity in ecological eommunities. Chicago:TheUniversityofChieagoPress,13～25
Vitousek P M, Matson P A , Van ckve K. 1989. Nitrogen availability and nitrification during succession, Primary,secondary and old field seres.Plant and Soil,115:229~239
Zhou SR,Zhang DY. 2006. Neutral theory in community ecology.Journal of Plant Ecology, 30 (5) :868~877