大兴安岭呼中地区3种林分的群落特征、物种多样性差异及其耦合关系
|
摘要
准确理解天然林林分群落特征及其与物种多样性耦合关系是提升天然林管理、达到多样性保护的关键。选择大兴安岭呼中地区典型落叶松林、杂木林、白桦落叶松林为研究对象,分别对乔木层、灌木层和草本层特征(高度、胸径、冠幅、盖度等)进行调查并计算丰富度指数、多样性指数和均匀度指数,旨在探究林分间差异及其耦合关系变化。结果表明:(1)乔木层的树高、枝下高表现出杂木林>落叶松林>白桦落叶松(P<0.05);落叶松林的胸径比白桦落叶松林和杂木林的高出6%和11%;灌木层的高度、盖度、地径、冠幅和草本层多度、盖度、高度在森林类型间均未表现出显著差异。(2)3个森林类型的乔、灌、草丰富度指数R、Shannon-Wiener指数及Simpson指数均表现出杂木林最大,乔木层和草本层呈相同规律,即杂木林>白桦落叶松林>落叶松林,而灌木层表现出杂木林>落叶松林>白桦落叶松林;白桦落叶松林和杂木林的乔木层均匀度Pielou指数和Alatalo指数约为落叶松林的3倍左右,而在灌木层和草本层在森林类型间差异不显著(P>0.05)。(3)典范对应分析(CCA)结果表明林分群落特征和生物多样性关系存在明显森林类型间差异。总体表现为灌木特征(冠幅、地径)、草本层特征(盖度、多度和高度)对多样性均有较大影响;白桦落叶松林和杂木林的胸径对多样性影响明显,而落叶松林的乔木高度(树高、枝下高)对多样性影响较大。杂木林随着灌木盖度、草本高度的增加,草本物种多样性降低、乔木多和灌木物种多样性增加;而落叶松林相同的多样性变化多伴随草本高度增加、多度和盖度变小。灌木层物种多样性增加多与乔木和草本物种多样性降低相伴随,在杂木林中同时伴随着乔木胸径和草本的盖度、多度增大、灌木冠幅变小,而白桦落叶松林则伴随灌木冠幅和草本多度盖度的减小。以上结果表明,林分群落特征与物种多样性存在耦合关系,上述解耦合结果为通过维持良好森林结构、多样性保护具有实践意义。
Exploring the differences and coupling relationships of species diversity and stand characteristics could provide data with which to improve biodiversity and ecological services of natural forests. In this study,three typical communities oflarch forest,shaw( mixed forest with various tree species),and birch-larch forest were selected at Huzhong in the Daxinganling mountains for measuring stand characteristics( height,diameter at breast height [DBH ],crown width,coverage,and others),richness index,diversity indexes,and evenness indexes of the arbor layer,shrub layer,and herb layer. Canonical correspondence analysis( CCA) was used to uncover their differing associations among forest types,together with a multiple analysis of variance to detect the inter-forest differences. The results showed that( 1) the height and clear tree bole height of the arbor layer differed with shaw > larch forest > birch-larch forest( P < 0.05); DBH of the larch forest was 6% and 11% higher than that in the birch-larch forest and the shaw,respectively. There were no significant differences in the shrub layer in the parameters of height,canopy coverage,ground diameter,crown width,as well as in the herb layer for species abundance,herb coverage,and herb height.( 2) The maximum species richness,Shannon-Wiener index,and Simpson index of the arbor layer,shrub layer,and herb layer all occurred in the shaw relative to the larch forest and the birch-larch forest. Moreover,the herb layer showed the same pattern as in the arbor layer,i.e.,shaw > birch-larch forest > larch forest; however,the shrub layer exhibited a different order,with shaw > larch forest > birch-larch forest. The species evenness of the arbor layer in the birch-larch forest and shaw were approximately three times that of the larch forest;however,there was no significant difference in the shrub layer and the herb layer( P > 0.05).( 3) CCA showed that the correspondence associations between community characteristics and species diversity differed among the three forest types. In general,for all three forests the characteristics of the shrub( ground diameter and crown width) and herb layer( coverage,abundance,and height) had a substantial influence on plant species diversity. Plant diversity was significantly affected by tree DBH in the birch-larch forest and the shaw; however,it was highly significantly affected by tree height in the larch forest. With the increase of shrub coverage and herb height,herb diversity decreased; whereas decreasing herb diversity usually corresponded with increasing diversity of the arbor and shrub layer in the shaw. In the larch forest,herb height increase,together with decreases in abundance and coverage usually corresponded with a decrease in herb diversity,and increases in arbor and shrub diversity. An increase in shrub diversity was usually accompanied by a decrease in arbor and herb diversity,but corresponded with increases in tree DBH,herb abundance,and coverage,and decreases in shrub crown width in the shaw; however,it was accompanied by decreases in shrub crown width,as well as herb abundance and coverage in the birch-larch forest. Our findings highlighted that community characteristics were closely coupled with species diversity,and the above-mentioned decoupling results will favor diversity conservation through maintaining good forest structure in the Daxinganling Mountain Region,Northeast China.
Exploring the differences and coupling relationships of species diversity and stand characteristics could provide data with which to improve biodiversity and ecological services of natural forests. In this study,three typical communities oflarch forest,shaw( mixed forest with various tree species),and birch-larch forest were selected at Huzhong in the Daxinganling mountains for measuring stand characteristics( height,diameter at breast height [DBH ],crown width,coverage,and others),richness index,diversity indexes,and evenness indexes of the arbor layer,shrub layer,and herb layer. Canonical correspondence analysis( CCA) was used to uncover their differing associations among forest types,together with a multiple analysis of variance to detect the inter-forest differences. The results showed that( 1) the height and clear tree bole height of the arbor layer differed with shaw > larch forest > birch-larch forest( P < 0.05); DBH of the larch forest was 6% and 11% higher than that in the birch-larch forest and the shaw,respectively. There were no significant differences in the shrub layer in the parameters of height,canopy coverage,ground diameter,crown width,as well as in the herb layer for species abundance,herb coverage,and herb height.( 2) The maximum species richness,Shannon-Wiener index,and Simpson index of the arbor layer,shrub layer,and herb layer all occurred in the shaw relative to the larch forest and the birch-larch forest. Moreover,the herb layer showed the same pattern as in the arbor layer,i.e.,shaw > birch-larch forest > larch forest; however,the shrub layer exhibited a different order,with shaw > larch forest > birch-larch forest. The species evenness of the arbor layer in the birch-larch forest and shaw were approximately three times that of the larch forest;however,there was no significant difference in the shrub layer and the herb layer( P > 0.05).( 3) CCA showed that the correspondence associations between community characteristics and species diversity differed among the three forest types. In general,for all three forests the characteristics of the shrub( ground diameter and crown width) and herb layer( coverage,abundance,and height) had a substantial influence on plant species diversity. Plant diversity was significantly affected by tree DBH in the birch-larch forest and the shaw; however,it was highly significantly affected by tree height in the larch forest. With the increase of shrub coverage and herb height,herb diversity decreased; whereas decreasing herb diversity usually corresponded with increasing diversity of the arbor and shrub layer in the shaw. In the larch forest,herb height increase,together with decreases in abundance and coverage usually corresponded with a decrease in herb diversity,and increases in arbor and shrub diversity. An increase in shrub diversity was usually accompanied by a decrease in arbor and herb diversity,but corresponded with increases in tree DBH,herb abundance,and coverage,and decreases in shrub crown width in the shaw; however,it was accompanied by decreases in shrub crown width,as well as herb abundance and coverage in the birch-larch forest. Our findings highlighted that community characteristics were closely coupled with species diversity,and the above-mentioned decoupling results will favor diversity conservation through maintaining good forest structure in the Daxinganling Mountain Region,Northeast China.
引文
[1]吕忠全.森林资源统计模型理论和方法的研究[D].南京:南京林业大学,2007.
[2]Maltamo M,Kangas A,Uuttera J,Torniainen T,Saramki J.Comparison of percentile based prediction methods and the Weibull distribution in describing the diameter distribution of heterogeneous Scots pine stands.Forest Ecology and Management,2000,133(3):263-274.
[3]汪殿蓓,暨淑仪,陈飞鹏,邢福武,彭少麟.深圳南山区天然森林群落多样性及演替现状.生态学报,2003,23(7):1415-1422.
[4]郑元润.大青沟森林植物群落物种多样性研究.生物多样性,1998,6(3):191-196.
[5]许晴,张放,许中旗,贾彦龙,尤建民.Simpson指数和Shannon-Wiener指数若干特征的分析及“稀释效应”.草业科学,2011,28(4):527-531.
[6]Magurran A E.Ecological Diversity and Its Measurement.Princeton:Princeton University Press,1988.
[7]王泺鑫,韩海荣,程小琴,康峰峰.辽河源自然保护区油松群落结构及物种多样性.生态学杂志,2016.35(2),314-320.
[8]Moeur M.Characterizing spatial patterns of trees using stem-mapped data.Forest Science,1993,39(4):756-775.
[9]郝占庆,张健,李步杭,叶吉,王绪高,姚晓琳.长白山次生杨桦林样地:物种组成与群落结构.植物生态学报,2008,32(2):251-261.
[10]黎芳,潘萍,宁金魁,赖国桢,欧阳勋志,徐辉,郭丽玲,吴自荣,易祖滨.飞播马尾松林林分空间结构对林下植被多样性的影响.东北林业大学学报,2016,44(11):31-35,40-40.
[11]苏日古嘎,张金屯,张斌,程佳佳,田世广,张钦弟,刘素军.松山自然保护区森林群落的数量分类和排序.生态学报,2010,30(10):2621-2629.
[12]郑景明,罗菊春.长白山阔叶红松林结构多样性的初步研究.生物多样性,2003,11(4):295-302.
[13]郭志华,臧润国,蒋有绪.生物多样性的形成、维持机制及其宏观研究方法.林业科学,2002,38(6):116-124.
[14]Lindenmayer D B,Franklin J F,Fischer J.General management principles and a checklist of strategies to guide forest biodiversity conservation.Biological Conservation,2006,131(3):433-445.
[15]Bordelon M A,Mc Allister D C,Holloway R.Sustainable forestry oregon style.Journal of Forestry,2000,98(1):26-34.
[16]贾应舍,于占华,斯日格楞.内蒙古大兴安岭林区生物多样性和自然保护区分析.内蒙古林业调查设计,2008,31(4):105-108.
[17]赵淑清,方精云,朴世龙,宗占江,吴晓莆,古陶.大兴安岭呼中地区白卡鲁山植物群落结构及其多样性研究.生物多样性,2004,12(1):182-189.
[18]陈亚南.落叶松云冷杉林空间结构多样性及择伐模型的研究[D].北京:北京林业大学,2016.
[19]刘燕,支玲,刘佳.国有林场森林资源管理绩效评价——以福建省将乐国有林场为例.林业科学,2015,51(5):117-126.
[20]周道玮,张正祥,靳英华,王平,王学志.东北植被区划及其分布格局.植物生态学报,2010,34(12):1359-1368.
[21]胡海清,魏书精,孙龙.1965—2010年大兴安岭森林火灾碳排放的估算研究.植物生态学报,2012,36(7):629-644.
[22]马克平,黄建辉,于顺利,陈灵芝.北京东灵山地区植物群落多样性的研究——Ⅱ丰富度、均匀度和物种多样性指数.生态学报,1995,15(3):268-277.
[23]Gauch H G J.Multivariate Analysis in Community Ecology.Cambridge:Cambridge University Press,1982.
[24]范文娟.子午岭森林群落草本植物的结构特征研究[D].杨凌:西北农林科技大学,2010.
[25]刘建泉,屈永才,王学福,王零,朱高,郭泽勇.青海云杉群落空间结构特征研究.干旱区资源与环境,2009,23(7):121-125.
[26]石晓东,韩有志.庞泉沟自然保护区寒温性针叶林的种群结构与分布格局.植物科学学报,2008,26(5):489-494.
[27]欧阳勋志,廖为明,彭世揆.天然阔叶林景观质量评价及其垂直结构优化技术.应用生态学报,2007,18(6):1388-1392.
[28]苏文会,朱晓武,范少辉,曾宪礼,刘广路.采伐对森林生态系统的影响综述.林业资源管理,2017,(3):35-40.
[29]张会儒,唐守正.森林生态采伐理论.林业科学,2008,44(10):127-131.
[30]Hao Z Q,Zhang J,Song B,Ye J,Li B H.Vertical structure and spatial associations of dominant tree species in an old-growth temperate forest.Forest Ecology and Management,2007,252(1/3):1-11.
[31]Li L,Huang Z L,Ye W H,Cao H L,Wei S G,Wang Z G,Lian J Y,Sun I F,Ma K P,He F L.Spatial distributions of tree species in a subtropical forest of China.Oikos,2009,118(4):495-502.
[32]Guo Y L,Lu J M,Franklin S B,Wang Q G,Xu Y Z,Zhang K H,Bao D C,Qiao X J,Huang H D,Lu Z J,Jiang M X.Spatial distribution of tree species in a species-rich subtropical mountain forest in central China.Canadian Journal of Forest Research,2013,43(9):826-835.
[33]Wang Q G,Bao D C,Guo Y L,Lu J M,Lu Z J,Xu Y Z,Zhang K H,Liu H B,Meng H J,Jiang M X,Qiao X J,Huang H D.Species associations in a species-rich subtropical forest were not well-explained by stochastic geometry of biodiversity.PLo S One,2014,9(5):e97300.
[34]洪仁辉,薛杨,王小燕,李敦禧,林之盼,宿少锋.文昌8个森林群落结构与物种多样性研究.热带农业科学,2015,35(10):51-56.
[35]李凤英,纪桂琴,石福臣.凉水国家级自然保护区森林群落结构及物种多样性分析.南开大学学报:自然科学版,2009,42(3):38-45.
[36]鱼腾飞,冯起,司建华,席海洋,陈丽娟.黑河下游额济纳绿洲植物群落物种多样性的空间异质性.应用生态学报,2011,22(8):1961-1966.
[37]李俊涛,谭晓京,蔡体久,罗传文,马玉心.凉水国家级自然保护区森林群落林下物种多样性.北京林业大学学报,2007,27(S2):266-271.
[38]Rey B,Jose M.Patterns of diversity in the strata of boreal montane forest in British Columbia.Journal of Vegetation Science,1995,6(1):95-98.
[39]蒋蕾,刘兆刚.大兴安岭天然白桦林群落结构特征及其物种多样性.森林工程,2014,30(4):12-17.
[40]董灵波,刘兆刚,李凤日,蒋蕾.大兴安岭主要森林类型林分空间结构及最优树种组成.林业科学研究,2014,27(6):734-740.
[41]李晓娜,朱莉莉,贺红士,郭瑞超,李旭辉.“天然林保护工程”对大兴安岭呼中森林的中长期影响.北方园艺,2015,(24):195-200.
[42]李月辉,胡远满,常禹,李秀珍,布仁仓,胡长河,王春林.大兴安岭林区道路对植物多样性的影响域.应用生态学报,2010,21(5):1112-1119.
[43]汪媛燕,王立,满多清.民勤绿洲荒漠过渡带群落特征及其物种多样性研究.四川农业大学学报,2014,32(4):355-361.
[44]王磊,高贤明,孙书存.岷江上游人工油松林群落空间结构:物种丰富度和盖度.林业科学,2004,40(6):8-12.
[45]张佳,李生宇,靳正忠,雷加强.防护林下草本植物层片物种多样性与环境因子的关系.干旱区研究,2011,28(1):118-125.
[46]雷相东,陆元昌,张会儒,张则路,陈晓光.抚育间伐对落叶松云冷杉混交林的影响.林业科学,2005,41(4):78-85.
[47]钟昌福,王玉芳,王岩,邱立红.大兴安岭国有林区北药开发的思考.福建林业科技,2006,33(4):243-245.
[2]Maltamo M,Kangas A,Uuttera J,Torniainen T,Saramki J.Comparison of percentile based prediction methods and the Weibull distribution in describing the diameter distribution of heterogeneous Scots pine stands.Forest Ecology and Management,2000,133(3):263-274.
[3]汪殿蓓,暨淑仪,陈飞鹏,邢福武,彭少麟.深圳南山区天然森林群落多样性及演替现状.生态学报,2003,23(7):1415-1422.
[4]郑元润.大青沟森林植物群落物种多样性研究.生物多样性,1998,6(3):191-196.
[5]许晴,张放,许中旗,贾彦龙,尤建民.Simpson指数和Shannon-Wiener指数若干特征的分析及“稀释效应”.草业科学,2011,28(4):527-531.
[6]Magurran A E.Ecological Diversity and Its Measurement.Princeton:Princeton University Press,1988.
[7]王泺鑫,韩海荣,程小琴,康峰峰.辽河源自然保护区油松群落结构及物种多样性.生态学杂志,2016.35(2),314-320.
[8]Moeur M.Characterizing spatial patterns of trees using stem-mapped data.Forest Science,1993,39(4):756-775.
[9]郝占庆,张健,李步杭,叶吉,王绪高,姚晓琳.长白山次生杨桦林样地:物种组成与群落结构.植物生态学报,2008,32(2):251-261.
[10]黎芳,潘萍,宁金魁,赖国桢,欧阳勋志,徐辉,郭丽玲,吴自荣,易祖滨.飞播马尾松林林分空间结构对林下植被多样性的影响.东北林业大学学报,2016,44(11):31-35,40-40.
[11]苏日古嘎,张金屯,张斌,程佳佳,田世广,张钦弟,刘素军.松山自然保护区森林群落的数量分类和排序.生态学报,2010,30(10):2621-2629.
[12]郑景明,罗菊春.长白山阔叶红松林结构多样性的初步研究.生物多样性,2003,11(4):295-302.
[13]郭志华,臧润国,蒋有绪.生物多样性的形成、维持机制及其宏观研究方法.林业科学,2002,38(6):116-124.
[14]Lindenmayer D B,Franklin J F,Fischer J.General management principles and a checklist of strategies to guide forest biodiversity conservation.Biological Conservation,2006,131(3):433-445.
[15]Bordelon M A,Mc Allister D C,Holloway R.Sustainable forestry oregon style.Journal of Forestry,2000,98(1):26-34.
[16]贾应舍,于占华,斯日格楞.内蒙古大兴安岭林区生物多样性和自然保护区分析.内蒙古林业调查设计,2008,31(4):105-108.
[17]赵淑清,方精云,朴世龙,宗占江,吴晓莆,古陶.大兴安岭呼中地区白卡鲁山植物群落结构及其多样性研究.生物多样性,2004,12(1):182-189.
[18]陈亚南.落叶松云冷杉林空间结构多样性及择伐模型的研究[D].北京:北京林业大学,2016.
[19]刘燕,支玲,刘佳.国有林场森林资源管理绩效评价——以福建省将乐国有林场为例.林业科学,2015,51(5):117-126.
[20]周道玮,张正祥,靳英华,王平,王学志.东北植被区划及其分布格局.植物生态学报,2010,34(12):1359-1368.
[21]胡海清,魏书精,孙龙.1965—2010年大兴安岭森林火灾碳排放的估算研究.植物生态学报,2012,36(7):629-644.
[22]马克平,黄建辉,于顺利,陈灵芝.北京东灵山地区植物群落多样性的研究——Ⅱ丰富度、均匀度和物种多样性指数.生态学报,1995,15(3):268-277.
[23]Gauch H G J.Multivariate Analysis in Community Ecology.Cambridge:Cambridge University Press,1982.
[24]范文娟.子午岭森林群落草本植物的结构特征研究[D].杨凌:西北农林科技大学,2010.
[25]刘建泉,屈永才,王学福,王零,朱高,郭泽勇.青海云杉群落空间结构特征研究.干旱区资源与环境,2009,23(7):121-125.
[26]石晓东,韩有志.庞泉沟自然保护区寒温性针叶林的种群结构与分布格局.植物科学学报,2008,26(5):489-494.
[27]欧阳勋志,廖为明,彭世揆.天然阔叶林景观质量评价及其垂直结构优化技术.应用生态学报,2007,18(6):1388-1392.
[28]苏文会,朱晓武,范少辉,曾宪礼,刘广路.采伐对森林生态系统的影响综述.林业资源管理,2017,(3):35-40.
[29]张会儒,唐守正.森林生态采伐理论.林业科学,2008,44(10):127-131.
[30]Hao Z Q,Zhang J,Song B,Ye J,Li B H.Vertical structure and spatial associations of dominant tree species in an old-growth temperate forest.Forest Ecology and Management,2007,252(1/3):1-11.
[31]Li L,Huang Z L,Ye W H,Cao H L,Wei S G,Wang Z G,Lian J Y,Sun I F,Ma K P,He F L.Spatial distributions of tree species in a subtropical forest of China.Oikos,2009,118(4):495-502.
[32]Guo Y L,Lu J M,Franklin S B,Wang Q G,Xu Y Z,Zhang K H,Bao D C,Qiao X J,Huang H D,Lu Z J,Jiang M X.Spatial distribution of tree species in a species-rich subtropical mountain forest in central China.Canadian Journal of Forest Research,2013,43(9):826-835.
[33]Wang Q G,Bao D C,Guo Y L,Lu J M,Lu Z J,Xu Y Z,Zhang K H,Liu H B,Meng H J,Jiang M X,Qiao X J,Huang H D.Species associations in a species-rich subtropical forest were not well-explained by stochastic geometry of biodiversity.PLo S One,2014,9(5):e97300.
[34]洪仁辉,薛杨,王小燕,李敦禧,林之盼,宿少锋.文昌8个森林群落结构与物种多样性研究.热带农业科学,2015,35(10):51-56.
[35]李凤英,纪桂琴,石福臣.凉水国家级自然保护区森林群落结构及物种多样性分析.南开大学学报:自然科学版,2009,42(3):38-45.
[36]鱼腾飞,冯起,司建华,席海洋,陈丽娟.黑河下游额济纳绿洲植物群落物种多样性的空间异质性.应用生态学报,2011,22(8):1961-1966.
[37]李俊涛,谭晓京,蔡体久,罗传文,马玉心.凉水国家级自然保护区森林群落林下物种多样性.北京林业大学学报,2007,27(S2):266-271.
[38]Rey B,Jose M.Patterns of diversity in the strata of boreal montane forest in British Columbia.Journal of Vegetation Science,1995,6(1):95-98.
[39]蒋蕾,刘兆刚.大兴安岭天然白桦林群落结构特征及其物种多样性.森林工程,2014,30(4):12-17.
[40]董灵波,刘兆刚,李凤日,蒋蕾.大兴安岭主要森林类型林分空间结构及最优树种组成.林业科学研究,2014,27(6):734-740.
[41]李晓娜,朱莉莉,贺红士,郭瑞超,李旭辉.“天然林保护工程”对大兴安岭呼中森林的中长期影响.北方园艺,2015,(24):195-200.
[42]李月辉,胡远满,常禹,李秀珍,布仁仓,胡长河,王春林.大兴安岭林区道路对植物多样性的影响域.应用生态学报,2010,21(5):1112-1119.
[43]汪媛燕,王立,满多清.民勤绿洲荒漠过渡带群落特征及其物种多样性研究.四川农业大学学报,2014,32(4):355-361.
[44]王磊,高贤明,孙书存.岷江上游人工油松林群落空间结构:物种丰富度和盖度.林业科学,2004,40(6):8-12.
[45]张佳,李生宇,靳正忠,雷加强.防护林下草本植物层片物种多样性与环境因子的关系.干旱区研究,2011,28(1):118-125.
[46]雷相东,陆元昌,张会儒,张则路,陈晓光.抚育间伐对落叶松云冷杉混交林的影响.林业科学,2005,41(4):78-85.
[47]钟昌福,王玉芳,王岩,邱立红.大兴安岭国有林区北药开发的思考.福建林业科技,2006,33(4):243-245.