秦岭南坡松栎林群落演替过程中物种组成及多样性动态分析
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摘要
为揭示秦岭南坡植物群落演替与物种多样性之间的复杂关系,采用空间代替时间的方法在秦岭南坡油松林、松栎混交林和锐齿栎林群落典型分布区域设置45个样地进行了群落学调查,统计分析了油松林→松栎混交林→锐齿栎林这一演替序列过程中草、灌、乔3层的物种组成和α物种多样性动态。结果表明:(1)秦岭南坡油松林、松栎混交林和锐齿栎林3种群落共有维管植物312种,隶属于73科183属,其中草本植物136种,灌木(含木质藤本)98种,乔木78种;油松林群落包含52科117属190种,松栎混交林群落含有60科95属129种,锐齿栎林群落则为50科109属178种。(2)沿油松林→松栎混交林→锐齿栎林群落演替序列,草本层物种均匀度和物种多样性显著降低,物种丰富度呈"V"形变化趋势;灌木层物种丰富度、均匀度以及物种多样性均无显著差异;乔木层物种多样性呈明显的单峰型变化,其中松栎混交林群落物种多样性指数最高。研究表明,秦岭南坡松栎林群落物种多样性随演替进展总体上呈现单峰型变化,与中期物种多样性假说一致;物种多样性可能仅仅是群落稳定的一个基础或前提条件。
Species diversity and its change is one of the main indicators of community succession dynamic process.Our objective was to examine the complex relationships between the species diversity and community succession in the successive serials from Pinus tabuliformis forest to P.tabuliformis-Quercus aliena var.acuteserrata mixed forest then to the subsequent Q.alienavar.acuteserrataforest on the south-facing slopes in Qinling Mountains.Changes of species composition and diversity in each vegetation stratum were studied using the method of substituting spatial difference for time change.The results showed that:(1)312 species of vascular plants belonging to 183genera and 73families were recorded in 45quadrats investigated,among which there were 136herbs,98shrubs encompassing woody climbers,and 78trees.The composition of the three communities was that there were 190species belonging to 117genera and 52families,129 species belonging to 95genera and 60families,and 178species belonging to 109genera and 50families inP.tabuliformis forest,P.tabuliformis-Q.aliena var.acuteserrata mixed forest and Q.aliena var. acuteserrataforest,respectively.(2)Species evenness and species diversity in herb layer decreased significantly during community succession(P<0.05),species richness showed a V-shaped trend;shrub layer species richness,evenness and species diversity did not change conspicuously(P>0.05);There was a significant hump-shaped pattern in species diversity in tree layer along the succession process,and the species diversity reached a peak in the community of P.tabuliformis-Q.aliena var.acuteserrata mixed forest.The studies above indicated that:changes of species diversity presented a hump-shaped model as a whole in the process of community succession of pine oak forests on the south-facing slopes in Qinling Mountains,which agrees with the intermediate species diversity hypothesis;The species diversity may simply be one of the basis or prerequisites to the community stability.
Species diversity and its change is one of the main indicators of community succession dynamic process.Our objective was to examine the complex relationships between the species diversity and community succession in the successive serials from Pinus tabuliformis forest to P.tabuliformis-Quercus aliena var.acuteserrata mixed forest then to the subsequent Q.alienavar.acuteserrataforest on the south-facing slopes in Qinling Mountains.Changes of species composition and diversity in each vegetation stratum were studied using the method of substituting spatial difference for time change.The results showed that:(1)312 species of vascular plants belonging to 183genera and 73families were recorded in 45quadrats investigated,among which there were 136herbs,98shrubs encompassing woody climbers,and 78trees.The composition of the three communities was that there were 190species belonging to 117genera and 52families,129 species belonging to 95genera and 60families,and 178species belonging to 109genera and 50families inP.tabuliformis forest,P.tabuliformis-Q.aliena var.acuteserrata mixed forest and Q.aliena var. acuteserrataforest,respectively.(2)Species evenness and species diversity in herb layer decreased significantly during community succession(P<0.05),species richness showed a V-shaped trend;shrub layer species richness,evenness and species diversity did not change conspicuously(P>0.05);There was a significant hump-shaped pattern in species diversity in tree layer along the succession process,and the species diversity reached a peak in the community of P.tabuliformis-Q.aliena var.acuteserrata mixed forest.The studies above indicated that:changes of species diversity presented a hump-shaped model as a whole in the process of community succession of pine oak forests on the south-facing slopes in Qinling Mountains,which agrees with the intermediate species diversity hypothesis;The species diversity may simply be one of the basis or prerequisites to the community stability.
引文
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[12]CHENG R M(程瑞梅),XIAO W F(肖文发),WANG X R(王晓荣),et al.Soil nutrient characteristics in different vegetation succession stages of Three Gorges Reservior area[J].Scientia Silvae Sinicae(林业科学),2010,46(9):1-6(in Chinese).
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[14]LEI N F(雷泞菲),SU ZH X(苏智先),SONG H X(宋会兴),et al.A comparative study on life-form spectra of evergreen broad-leaved forest in different successions in Jinyun Mountain[J].Chinese Journal of Applied Ecology(应用生态学报),2002,13(3):267-270(in Chinese).
[15]YAN X D(延晓东),ZHAO SH D(赵士洞),YU ZH L(于振良).Modeling growth and succession of northeastern China forests and its applications in global change studies[J].Acta Phytoecologica Sinica(植物生态学报),2000,24(1):1-8(in Chinese).
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[18]KANG B(康冰),WANG D X(王得祥),et al.Regeneration characteristics and related affecting factors of Pinus tabulaeformis secondary forests in Qinling Mountains[J].Chinese Journal of Applied Ecology(应用生态学报),2011,22(7):1 659-1 667(in Chinese).
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[20]LIU G Q(刘广全),TU X N(土小宁),ZHAO SH D(赵士洞),et al.Distributional characteristics on biomass and nutrient elements of pine-oak forest belt in MT.Qinling[J].Scientia Silvae Sinicae(林业科学),2001,37(1):28-36(in Chinese).
[21]YU F,WANG D X,SHI X X,et al.Seed dispersal by small rodents favours oak over pine regeneration in the pine-oak forests of the Qinling Mountains,China[J].Scandinavian Journal of Forest Research,2013,28(6):540-549.
[22]YU F,WANG D X,SHI X X,et al.Effects of environmental factors on tree seedling regeneration in a pine-oak mixed forest in the Qinling Mountains,China[J].Journal of Mountain Science,2013,DOI:10.1007/s11629-013-2548-1.
[23]YU F(于飞),WANG D X(王得祥),SHI X X(史晓晓),et al.Species composition and regeneration characteristics of main woody plant seedlings in a pine-oak mixed forest in Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2013,33(3):592-598(in Chinese).
[24]YU F(于飞),WANG D X(王得祥),et al.Regeneration rules of three dominant tree species in a pine-oak mixed forest under different ecological conditions in the Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2013,33(5):1 020-1 026(in Chinese).
[25]ZHAO H(赵桦),YANG P J(杨培君).A floristic analysis of the seed plants in the middle area of south slope of Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2001,21(4):732-739(in Chinese).
[26]张巧明.秦岭南坡中段主要植物群落及物种多样性研究[D].陕西杨陵:西北农林科技大学,2012.
[27]ZHANG M Y(张孟颖),GUO T(郭韬),WANG Y(王源),et al.The ferns study of national field practice base in Xunyang dam of Qinling[J].Journal of Shaanxi Normal University(Nat.Sci.Edi.)(陕西师范大学学报·自然科学版),2009,37(7):42-45(in Chinese).
[28]MA K P(马克平).Measurement of bio-community diversity:Ⅰ.The method ofα-diversity[J].Chinese Biodiversity(生物多样性),1994,2(3):162-168(in Chinese).
[29]MA K P(马克平),LIU Y M(刘玉明).Measurement of bio-community diversity:Ⅰ.The method ofα-diversity[J].Chinese Biodiversity(生物多样性)(second),1994,2(4):231-239(in Chinese).
[30]WANG SH X(王世雄),WANG X A(王孝安),LI G Q(李国庆),et al.Species diversity and environmental interpretation in the process of community succession in the Ziwu Mountain of Shaanxi Province[J].Acta Ecologica Sinica(生态学报),2010,30(6):1 638-1 647(in Chinese).
[31]LI M(李苗),LI K R(李凯荣),YANG X Y(杨晓毅),et al.Study on structure and plant diversity of Pinus tabuliformis stands in Chunhua County[J].Research of Soil and Water Conservation(水土保持研究),2010,17(6):142-147(in Chinese).
[32]李苗.黄土残塬沟壑区人工油松林林分结构及林下植物多样性研究[D].陕西杨陵:西北农林科技大学,2011.
[33]HORN H.The ecology of secondary succession[J].Annual Review of Ecology and Systematics,1974,5:25-27.
[34]ODUM E P.The strategy of ecosystem development[J].Science,1969,164:262-270.
[35]CLEMENTS F E.Nature and structure of the climax[J].The Journal of Ecology,1936,24(1):252-284.
[36]MARGALEF R.Perspectives in ecological theory[M].Chicago:University of Chicago Press,1968.
[37]QIN J(秦娟),WANG K B(王凯博),SHANGGUAN ZH P(上官周平).Change of plant diversity of Populus davidiana,Quercus liaotungensis and their mixed forest in Ziwuling area of Loess Plateau[J].Journal of Northwest A&F University(Nat.Sci.Edi.)(西北农林科技大学学报·自然科学版),2007,35(5):131-140(in Chinese).
[38]ELTON C S.The reasons for conservation[M]//The Ecology of Invasions by Animals and Plants.London:Chapman&Hall,1958:143-153.
[39]LI Y Y(李裕元),SHAO M A(邵明安).The change of plant diversity during natural recovery process of vegetation in Ziwuling area[J].Acta Ecologica Sinica(生态学报),2004,24(2):252-260(in Chinese).
[2]KALACSKA M E R,SANCHEZ-AZOFEIFA G A,CALVO-ALVARADO J C,et al.Effects of season and successional stage on leaf area index and spectral vegetation indices in three Mesoamerican tropical dry forests[J].Biotropica,2005,37:486-496.
[3]ZHANG J Y(张继义),ZHAO H L(赵哈林),ZHANG T H(张铜会),et al.Dynamics of species diversity of communities in restoration processes in Horqin sandy land[J].Acta Phytoecologica Sinica(植物生态学报),2004,28(1):86-92(in Chinese).
[4]SHI SH Y(石胜友),YANG J D(杨季冬),WANG ZH P(王周平),et al.Species diversity in the progression of ecological restoration of artificial mixed forest after wind-damage in Jinyun Mountain,Sichuan[J].Biodiversity Science(生物多样性),2002,10(3):274-279(in Chinese).
[5]SAD S.Floristic and life form diversity in post-pasture successions on a Mediterranean island(Corsica)[J].Plant Ecology,2001,162:67-76.
[6]MADEIRA B G,ESPIRITO-SANTO M M,NETO S D,et al.Changes in tree and liana communities along a successional gradient in a tropical dry forest in south-eastern Brazil[J].Plant Ecology,2009,201:291-304.
[7]LOUCKS O L.Evolution of diversity,efficiency and community stability[J].American Zoologist,1970,10(1):17-25.
[8]HOWARD L F,LEE T D.Temporal patterns of vascular plant diversity in southeastern New Hampshire forests[J].Forest Ecology and Management,2003,185:5-20.
[9]SHAO X Q(邵新庆),WANG K(王堃),WANG Y W(王赟文),et al.Dynamics of plant community during natural restoration and succession of the Inner Mongolia Steppe[J].Acta Ecologica Sinica(生态学报),2008,28(2):856-860(in Chinese).
[10]FIERKE M K,KAUFFMAN J B.Riverscape-level patterns of riparian plant diversity along a successional gradient,Willamette river,Oregon[J].Plant Ecology,2006,185:85-95.
[11]WANG SH S(王树森),YU X X(余新晓),et al.Community structure and species diversity changes of forest vegetation with succession in Rocky Mountain Area of North of China[J].Research of Soil and Water Conservation(水土保持研究),2006,13(6):48-50(in Chinese).
[12]CHENG R M(程瑞梅),XIAO W F(肖文发),WANG X R(王晓荣),et al.Soil nutrient characteristics in different vegetation succession stages of Three Gorges Reservior area[J].Scientia Silvae Sinicae(林业科学),2010,46(9):1-6(in Chinese).
[13]PENG SH L(彭少麟).Studies on edge effect of successional communities and resoration of forest fragmentation in low sub-tropics[J].Acta Ecologica Sinica(生态学报),2000,20(1):1-8(in Chinese).
[14]LEI N F(雷泞菲),SU ZH X(苏智先),SONG H X(宋会兴),et al.A comparative study on life-form spectra of evergreen broad-leaved forest in different successions in Jinyun Mountain[J].Chinese Journal of Applied Ecology(应用生态学报),2002,13(3):267-270(in Chinese).
[15]YAN X D(延晓东),ZHAO SH D(赵士洞),YU ZH L(于振良).Modeling growth and succession of northeastern China forests and its applications in global change studies[J].Acta Phytoecologica Sinica(植物生态学报),2000,24(1):1-8(in Chinese).
[16]徐化成.油松[M].北京:中国林业出版社,1990:113-115.
[17]YU M(岳明),DANG G D(党高弟),YONG L J(雍立军).The basic features of vegetation of Foping Nature Conservation in Shaanxi Province[J].Journal of Wuhan Botanical Research(武汉植物学研究),1999,17(1):22-28(in Chinese).
[18]KANG B(康冰),WANG D X(王得祥),et al.Regeneration characteristics and related affecting factors of Pinus tabulaeformis secondary forests in Qinling Mountains[J].Chinese Journal of Applied Ecology(应用生态学报),2011,22(7):1 659-1 667(in Chinese).
[19]WU H(吴昊),WANG D X(王得祥),HU Y N(胡有宁),et al.Numerical classification and ordination of pine and oak mixed forest communities in the middle part of Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2012,32(8):1 671-1 679(in Chinese).
[20]LIU G Q(刘广全),TU X N(土小宁),ZHAO SH D(赵士洞),et al.Distributional characteristics on biomass and nutrient elements of pine-oak forest belt in MT.Qinling[J].Scientia Silvae Sinicae(林业科学),2001,37(1):28-36(in Chinese).
[21]YU F,WANG D X,SHI X X,et al.Seed dispersal by small rodents favours oak over pine regeneration in the pine-oak forests of the Qinling Mountains,China[J].Scandinavian Journal of Forest Research,2013,28(6):540-549.
[22]YU F,WANG D X,SHI X X,et al.Effects of environmental factors on tree seedling regeneration in a pine-oak mixed forest in the Qinling Mountains,China[J].Journal of Mountain Science,2013,DOI:10.1007/s11629-013-2548-1.
[23]YU F(于飞),WANG D X(王得祥),SHI X X(史晓晓),et al.Species composition and regeneration characteristics of main woody plant seedlings in a pine-oak mixed forest in Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2013,33(3):592-598(in Chinese).
[24]YU F(于飞),WANG D X(王得祥),et al.Regeneration rules of three dominant tree species in a pine-oak mixed forest under different ecological conditions in the Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2013,33(5):1 020-1 026(in Chinese).
[25]ZHAO H(赵桦),YANG P J(杨培君).A floristic analysis of the seed plants in the middle area of south slope of Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2001,21(4):732-739(in Chinese).
[26]张巧明.秦岭南坡中段主要植物群落及物种多样性研究[D].陕西杨陵:西北农林科技大学,2012.
[27]ZHANG M Y(张孟颖),GUO T(郭韬),WANG Y(王源),et al.The ferns study of national field practice base in Xunyang dam of Qinling[J].Journal of Shaanxi Normal University(Nat.Sci.Edi.)(陕西师范大学学报·自然科学版),2009,37(7):42-45(in Chinese).
[28]MA K P(马克平).Measurement of bio-community diversity:Ⅰ.The method ofα-diversity[J].Chinese Biodiversity(生物多样性),1994,2(3):162-168(in Chinese).
[29]MA K P(马克平),LIU Y M(刘玉明).Measurement of bio-community diversity:Ⅰ.The method ofα-diversity[J].Chinese Biodiversity(生物多样性)(second),1994,2(4):231-239(in Chinese).
[30]WANG SH X(王世雄),WANG X A(王孝安),LI G Q(李国庆),et al.Species diversity and environmental interpretation in the process of community succession in the Ziwu Mountain of Shaanxi Province[J].Acta Ecologica Sinica(生态学报),2010,30(6):1 638-1 647(in Chinese).
[31]LI M(李苗),LI K R(李凯荣),YANG X Y(杨晓毅),et al.Study on structure and plant diversity of Pinus tabuliformis stands in Chunhua County[J].Research of Soil and Water Conservation(水土保持研究),2010,17(6):142-147(in Chinese).
[32]李苗.黄土残塬沟壑区人工油松林林分结构及林下植物多样性研究[D].陕西杨陵:西北农林科技大学,2011.
[33]HORN H.The ecology of secondary succession[J].Annual Review of Ecology and Systematics,1974,5:25-27.
[34]ODUM E P.The strategy of ecosystem development[J].Science,1969,164:262-270.
[35]CLEMENTS F E.Nature and structure of the climax[J].The Journal of Ecology,1936,24(1):252-284.
[36]MARGALEF R.Perspectives in ecological theory[M].Chicago:University of Chicago Press,1968.
[37]QIN J(秦娟),WANG K B(王凯博),SHANGGUAN ZH P(上官周平).Change of plant diversity of Populus davidiana,Quercus liaotungensis and their mixed forest in Ziwuling area of Loess Plateau[J].Journal of Northwest A&F University(Nat.Sci.Edi.)(西北农林科技大学学报·自然科学版),2007,35(5):131-140(in Chinese).
[38]ELTON C S.The reasons for conservation[M]//The Ecology of Invasions by Animals and Plants.London:Chapman&Hall,1958:143-153.
[39]LI Y Y(李裕元),SHAO M A(邵明安).The change of plant diversity during natural recovery process of vegetation in Ziwuling area[J].Acta Ecologica Sinica(生态学报),2004,24(2):252-260(in Chinese).