秦岭南坡油松-锐齿槲栎混交林群落不同层次多样性特征及环境解释
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摘要
在秦岭山脉南坡陕西省境内设置11个样地进行实地调查,从植物物种组成、多样性指数差异以及多样性指数沿环境梯度排序等方面对该区域油松-锐齿槲栎天然混交林的群落总体以及乔木层、灌木层、草本层的各自4个α多样性指数(Patrick丰富度指数、Simpson优势度指数、Shannon-Wiener多样性指数、Pielou均匀度指数)特征进行分析,以揭示该群落多样性特征及其与环境因素的关系。结果显示:(1)11个样地共记录维管束植物112种,隶属于45科77属,分布较广的物种有蔷薇科、菊科、百合科、松科、禾本科和忍冬科。(2)研究区群落中建群种锐齿槲栎和油松的优势地位突出,乔木层伴生种稀少,平均仅为4.8种,多样性水平较低;灌木层物种组成丰富,平均11.4种;草本层次之,平均9种。(3)单因素方差分析表明,该群落不同层次的多样性指数之间均存在极显著差异(P<0.01);LSD多重比较及多样性指数变化趋势分析表明,群落多样性水平表现为总体>灌木层>草本层>乔木层。(4)CCA法将群落不同层次多样性指数沿地形因子和土壤养分等12项环境指标梯度的分布划分为界线明显的3个区域,即Ⅰ区(中等浓度土壤磷素和有机质制约区)、Ⅱ区(非环境因子制约区)和Ⅲ区(高浓度土壤磷素制约区);群落总体多样性主要受灌木层影响,同时也受乔木层均匀度影响,但环境因子对乔木层多样性基本无限制性作用。(5)土壤速效磷、全磷和有机质是造成研究区群落不同层次多样性差异的主要因素,海拔、坡向等地形因子对群落多样性影响不大,可能是由于土壤养分的直接影响削弱了地形因子的间接性生态作用。研究表明:保持中等水平的土壤速效磷(3.775±1.529mg·kg-1)和有机质含量[(4.079±1.001)%],并通过合理补植松栎幼树和伐除枯死木等抚育手段提高乔木层均匀度是维持秦岭南坡油松-锐齿槲栎混交林群落生物多样性的有效措施。
Set 11plots in the southern slope of Qinling Mountains in Shaanxi Province,the characteristics of Patrick richness index,Simpson dominance index,Shannon-Wiener diversity index and Pielou evenness index are studied,in terms of the species comprise,diversity index differences and gradient ordination between diversity indices and environmental factors of Pinus tabuliformis and Quercus aliena var.accuteserrata mixed forest,to reveal the relationships between community diversity characteristics and environmental factors.Results indicated that:(1)11plots record 112species vascular plants,belong to 45families 77genera.The wide distribution species include Rosaceae,Asteraceae,Liliaceae,Pinaceae,Poaceae and Caprifoliaceae.(2)The dominant positions of P.tabuliformis and Q.alienavar.accuteserrataare significant.The auxiliary species in tree layer are less with average only 4.8and the biodiversity level is low.The species compositions of shrub layer are rich,average 11.4.The herb layer takes second place,average 9.(3)One-way ANOVA analysis indicates difference of diversity indices from 4different forest layers all reaches extremely significant level(P<0.01),LSD multiple comparison illustrates the values of diversity indices from community overall are the highest,while diversity level of tree layer is the lowest.(4)CCA method divides the distribution of diversity indices from different layers along 12environmental indicators(terrain factors and soil nutrients)gradient into 3sectors,they areⅠ(medium concentration of soil phosphorus and organic matters restricted area),Ⅱ(non-environmental factors restricted area)andⅢ(high concentration of soil phosphorus restricted area);community overall diversity indices are mainly influenced by shrub layer,meanwhile influenced by tree layer evenness indices,but environmental factors nearly have no restrictive roles on tree layer diversity.(5)The main factors causing diversity difference among different layers of community in study area are soil available phosphorus,total phosphorus and organic matter,while elevation,slope and other terrain factors have no significant effects on community diversity.It may be due to soil nutrients direct influence weakens indirect ecological role of terrain factors.Studies show that keeping moderate soil nutrients and improving tree layer evenness by upbringing is an effective measure for maintaining P.tabuliformis and Q.alienavar.accuteserrata mixed forest biodiversity in the southern slope of Qinling Mountains.
Set 11plots in the southern slope of Qinling Mountains in Shaanxi Province,the characteristics of Patrick richness index,Simpson dominance index,Shannon-Wiener diversity index and Pielou evenness index are studied,in terms of the species comprise,diversity index differences and gradient ordination between diversity indices and environmental factors of Pinus tabuliformis and Quercus aliena var.accuteserrata mixed forest,to reveal the relationships between community diversity characteristics and environmental factors.Results indicated that:(1)11plots record 112species vascular plants,belong to 45families 77genera.The wide distribution species include Rosaceae,Asteraceae,Liliaceae,Pinaceae,Poaceae and Caprifoliaceae.(2)The dominant positions of P.tabuliformis and Q.alienavar.accuteserrataare significant.The auxiliary species in tree layer are less with average only 4.8and the biodiversity level is low.The species compositions of shrub layer are rich,average 11.4.The herb layer takes second place,average 9.(3)One-way ANOVA analysis indicates difference of diversity indices from 4different forest layers all reaches extremely significant level(P<0.01),LSD multiple comparison illustrates the values of diversity indices from community overall are the highest,while diversity level of tree layer is the lowest.(4)CCA method divides the distribution of diversity indices from different layers along 12environmental indicators(terrain factors and soil nutrients)gradient into 3sectors,they areⅠ(medium concentration of soil phosphorus and organic matters restricted area),Ⅱ(non-environmental factors restricted area)andⅢ(high concentration of soil phosphorus restricted area);community overall diversity indices are mainly influenced by shrub layer,meanwhile influenced by tree layer evenness indices,but environmental factors nearly have no restrictive roles on tree layer diversity.(5)The main factors causing diversity difference among different layers of community in study area are soil available phosphorus,total phosphorus and organic matter,while elevation,slope and other terrain factors have no significant effects on community diversity.It may be due to soil nutrients direct influence weakens indirect ecological role of terrain factors.Studies show that keeping moderate soil nutrients and improving tree layer evenness by upbringing is an effective measure for maintaining P.tabuliformis and Q.alienavar.accuteserrata mixed forest biodiversity in the southern slope of Qinling Mountains.
引文
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[13]LIU J J(刘建军),WANG D X(王得祥),LEI R D(雷瑞德),et al.Fine roots biomass and spatial dynamic in the natural stands of sharp tooth oak Chinese pine at Huoditang Forest Region[J].Journal of Northwest Forestry College(西北林学院学报),2004,19(1):1-4(in Chinese).
[14]PENG SH L(彭舜磊),WANG D X(王得祥).Comparison of community characteristics of plantation and secondary forest of Pinus armandii in Huoditang Forest Region of Qinling Mountain[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2009,29(11):2 301-2311(in Chinese).
[15]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).
[16]WU H(吴昊),LIU H(刘华),ZHANG Y(张洋),et al.Coupling relationship between species diversity of planted Chinese Pine(Pinus tabulaeformis Carr.)communities and environment factors in mountain area of Shangluo[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2012,32(2):377-383(in Chinese).
[17]DAI L M(代力民),TANG L N(唐立娜),CAO Y M(曹玉明),et al.Quantitative classification and ordination aiming to realize ecological land classification for the mountainous region in eastern Liaoning Province[J].Sci.Silv.Sin.(林业科学),2008,44(3):6-12(in Chinese).
[18]ZHANG F(张峰),ZHANG J T(张金屯),ZHANG F(张峰).Pattern of forest vegetation and its environmental interpretation in Zhuweigou,Lishan Mountain Nature Reserve[J].Acta Ecol.Sin.(生态学报),2003,23(3):422-427(in Chinese).
[19]中国科学院南京土壤研究所.土壤理化性质分析[M].上海:上海科学技术出版社,1978:62-489.
[20]WANG Y S(王育松),SHANGGUAN T L(上官铁梁).Discussion on calculating method of important values[J].Journal of Shanxi University(Nat.Sci.Edi.)(山西大学学报·自然科学版),2010,33(2):312-316(in Chinese).
[21]MA K P(马克平).The measure method of bio-community diversity:the method ofα-diversity[J].Chinese Biodiversity(生物多样性),1994,2(3):162-168(in Chineses).
[22]中国科学院西北植物研究所.秦岭植物志[M].北京:科学出版社,1976:15-16,17-18,79-80.
[23]HU N(胡楠),XU H C(许会才),DING SH Y(丁圣彦).Community ecological characteristics of deciduous-conifer mixed forests in Shiren Mountain Natural Reserve[J].Journal of Henan University(Nat.Sci.Edi.)(河南大学学报·自然科学版),2007,37(5):502-504(in Chinese).
[24]LI X R(李新荣).Analyzing construction of shrub layer in different communities in the spruce broad-leaved mixed geobotanical zone of Moscow State[J].Chinese Bulletin of Botany(植物学通报),1997,14(addition):40-48(in Chinese).
[25]CAO X P(曹旭平),GUO Q Q(郭其强),ZHANG W H(张文辉).Species compositions and preponderant population dynamic of the Pinus tabulaeformis forest and Pinus tabulaeformis-Quercus liaotungensis forest in Huanglong Mountain[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2010,30(5):1 012-1 019(in Chinese).
[26]LIU X D(刘晓东),ZHANG Y L(张彦雷).Natural regeneration of Korean Pine mixed forests in Dong Zhelenghe Forest Station,Xiaoxing’an Mountains[J].Journal of Northeast Forestry University(东北林业大学学报),2009,37(9):8-11(in Chinese).
[27]LI X R(李新荣).Interspecific association and correlation of shrub layer in the coniferous-broad leaved mixed geobotanical zone of Russia Plain[J].Acta Ecol.Sin.(生态学报),1999,19(1):55-60(in Chinese).
[28]SHI J H(史军辉),HUANG ZH L(黄忠良),ZHOU X Y(周小勇),et al.Vertical pattern of plant community and biodiversity on the Dinghu Mountain[J].Chinese Journal of Ecology(生态学杂志),2005,24(10):1 143-1 146(in Chinese).
[29]ZHU ZH L(朱志玲),LI D ZH(李德志),WANG X P(王绪平),et al.Water physiology integration and its ecological effect of clonal plants[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2006,26(12):2 602-2 614(in Chinese).
[30]ZHAO J ZH(赵建中),PENG M(彭敏),LIU W(刘伟),et al.Correlation between growing characters of main species belonging to different functional groups and ground temperature in Kobresia meadow[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2012,32(3):559-565(in Chinese).
[31]WANG CH(汪超),WANG X A(王孝安),GUO H(郭华),et al.Species diversities of major communities in Malan Forest Region of the Loess Plateau[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2006,26(4):791-797(in Chinese).
[32]MO J M(莫江明).Phosphorus availability of soils under degraded pine,mixed and monsoon evergreen broadleaf forests of subtropical China[J].Guihaia(广西植物),2005,25(2):186-192(in Chinese).
[33]XIAO C Y(肖慈英),HUANG Q CH(黄青春),YUAN H H(阮宏华).Characteristics of decomposition of litter from pine,oak and pineoak mixed forests[J].Acta Pedo Sin.(土壤学报),2002,35(9):763-767(in Chinese).
[34]ZHANG L J(张林静),YUE M(岳明),GU F X(顾峰雪),et al.Coupling relationship between plant communities.species diversity and soil factors in ecotone between desert and oasis in Fukang,Xinjiang[J].Chinese Journal of Applied Ecology(应用生态学报),2002,13(6):658-662(in Chinese).
[2]JEAN C B,JOAO A J,ARY T O F.Intermediary disturbance increases tree diversity in riverine forest of Southern Brazil[J].Biodiversity Conservation,2010,19:2 371-2 387.
[3]GUADALUPE W L,FRANCISCO L.Tree species diversity driven by environ mental and anthropogenic factors in tropical dry forest fragments of central Veracruz,Mexico[J].Biodiversity Conservation,2009,18:3 269-3 293.
[4]TANG ZH Y(唐志尧),FANG J Y(方精云).A review on the elevational patterns of plant species diversity[J].Biodiversity Science(生物多样性),2004,12(1):20-28(in Chinese).
[5]XU Y J(徐远杰),CHEN Y N(陈亚宁),LI W H(李卫红),et al.Distribution pattern and environmental interpretation of plant species diversity in the mountainous region of Ili River Valley,Xinjiang,China[J].Chinese Journal of Plant Ecology(植物生态学报),2010,34(10):1 142-1 154(in Chinese).
[6]RANA M S,SAMANT S S,RAWAT Y S.Plant communities and factors responsible for vegetation pattern in an alpine area of the Northwestern Himalaya[J].Journal of Mountain Science,2011,8:817-826.
[7]张金屯.数量生态学(第2版)[M].北京:科学出版社,2004.
[8]WANG D X(王得祥),LIN Y Y(蔺雨阳),LEI R D(雷瑞德),et al.Community composition and classification of natural forest of Chinese pine(Pinus tabulaeformis Carr.)in Qinling Mountains[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2009,29(5):867-873(in Chinese).
[9]牛春山.陕西树木志[M].北京:中国林业出版社,1990:1 189-1 199.
[10]WU H(吴昊),WANG D X(王得祥),HUANG Q P(黄青平),et al.Influence of environmental factors on species diversity of different associations of pine-oak mixed forest in the middle part of south Qinling Mountains[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),2012,40(9):41-50(in Chinese).
[11]ZHANG Y Y(张远迎),ZHANG SH X(张硕新),DANG K L(党坤良),et al.Evaluation on soil nutrient conditions of main types of forests at Huoditang Forest Region[J].Journal of Northwest Forestry College(西北林学院学报),1996,11(4):28-32(in Chinese).
[12]DANG K L(党坤良),LEI R D(雷瑞德),ZHANG SH X(张硕新),et al.Characteristics of sharp tooth oak forest and mixed forest between pine and oak on south facing slope of the Qinling Mountains[J].Journal of Northwest Forestry College(西北林学院学报),1996,11(addition):127-131(in Chinese).
[13]LIU J J(刘建军),WANG D X(王得祥),LEI R D(雷瑞德),et al.Fine roots biomass and spatial dynamic in the natural stands of sharp tooth oak Chinese pine at Huoditang Forest Region[J].Journal of Northwest Forestry College(西北林学院学报),2004,19(1):1-4(in Chinese).
[14]PENG SH L(彭舜磊),WANG D X(王得祥).Comparison of community characteristics of plantation and secondary forest of Pinus armandii in Huoditang Forest Region of Qinling Mountain[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2009,29(11):2 301-2311(in Chinese).
[15]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).
[16]WU H(吴昊),LIU H(刘华),ZHANG Y(张洋),et al.Coupling relationship between species diversity of planted Chinese Pine(Pinus tabulaeformis Carr.)communities and environment factors in mountain area of Shangluo[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2012,32(2):377-383(in Chinese).
[17]DAI L M(代力民),TANG L N(唐立娜),CAO Y M(曹玉明),et al.Quantitative classification and ordination aiming to realize ecological land classification for the mountainous region in eastern Liaoning Province[J].Sci.Silv.Sin.(林业科学),2008,44(3):6-12(in Chinese).
[18]ZHANG F(张峰),ZHANG J T(张金屯),ZHANG F(张峰).Pattern of forest vegetation and its environmental interpretation in Zhuweigou,Lishan Mountain Nature Reserve[J].Acta Ecol.Sin.(生态学报),2003,23(3):422-427(in Chinese).
[19]中国科学院南京土壤研究所.土壤理化性质分析[M].上海:上海科学技术出版社,1978:62-489.
[20]WANG Y S(王育松),SHANGGUAN T L(上官铁梁).Discussion on calculating method of important values[J].Journal of Shanxi University(Nat.Sci.Edi.)(山西大学学报·自然科学版),2010,33(2):312-316(in Chinese).
[21]MA K P(马克平).The measure method of bio-community diversity:the method ofα-diversity[J].Chinese Biodiversity(生物多样性),1994,2(3):162-168(in Chineses).
[22]中国科学院西北植物研究所.秦岭植物志[M].北京:科学出版社,1976:15-16,17-18,79-80.
[23]HU N(胡楠),XU H C(许会才),DING SH Y(丁圣彦).Community ecological characteristics of deciduous-conifer mixed forests in Shiren Mountain Natural Reserve[J].Journal of Henan University(Nat.Sci.Edi.)(河南大学学报·自然科学版),2007,37(5):502-504(in Chinese).
[24]LI X R(李新荣).Analyzing construction of shrub layer in different communities in the spruce broad-leaved mixed geobotanical zone of Moscow State[J].Chinese Bulletin of Botany(植物学通报),1997,14(addition):40-48(in Chinese).
[25]CAO X P(曹旭平),GUO Q Q(郭其强),ZHANG W H(张文辉).Species compositions and preponderant population dynamic of the Pinus tabulaeformis forest and Pinus tabulaeformis-Quercus liaotungensis forest in Huanglong Mountain[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2010,30(5):1 012-1 019(in Chinese).
[26]LIU X D(刘晓东),ZHANG Y L(张彦雷).Natural regeneration of Korean Pine mixed forests in Dong Zhelenghe Forest Station,Xiaoxing’an Mountains[J].Journal of Northeast Forestry University(东北林业大学学报),2009,37(9):8-11(in Chinese).
[27]LI X R(李新荣).Interspecific association and correlation of shrub layer in the coniferous-broad leaved mixed geobotanical zone of Russia Plain[J].Acta Ecol.Sin.(生态学报),1999,19(1):55-60(in Chinese).
[28]SHI J H(史军辉),HUANG ZH L(黄忠良),ZHOU X Y(周小勇),et al.Vertical pattern of plant community and biodiversity on the Dinghu Mountain[J].Chinese Journal of Ecology(生态学杂志),2005,24(10):1 143-1 146(in Chinese).
[29]ZHU ZH L(朱志玲),LI D ZH(李德志),WANG X P(王绪平),et al.Water physiology integration and its ecological effect of clonal plants[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2006,26(12):2 602-2 614(in Chinese).
[30]ZHAO J ZH(赵建中),PENG M(彭敏),LIU W(刘伟),et al.Correlation between growing characters of main species belonging to different functional groups and ground temperature in Kobresia meadow[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2012,32(3):559-565(in Chinese).
[31]WANG CH(汪超),WANG X A(王孝安),GUO H(郭华),et al.Species diversities of major communities in Malan Forest Region of the Loess Plateau[J].Acta Bot.Boreal.-Occident.Sin.(西北植物学报),2006,26(4):791-797(in Chinese).
[32]MO J M(莫江明).Phosphorus availability of soils under degraded pine,mixed and monsoon evergreen broadleaf forests of subtropical China[J].Guihaia(广西植物),2005,25(2):186-192(in Chinese).
[33]XIAO C Y(肖慈英),HUANG Q CH(黄青春),YUAN H H(阮宏华).Characteristics of decomposition of litter from pine,oak and pineoak mixed forests[J].Acta Pedo Sin.(土壤学报),2002,35(9):763-767(in Chinese).
[34]ZHANG L J(张林静),YUE M(岳明),GU F X(顾峰雪),et al.Coupling relationship between plant communities.species diversity and soil factors in ecotone between desert and oasis in Fukang,Xinjiang[J].Chinese Journal of Applied Ecology(应用生态学报),2002,13(6):658-662(in Chinese).