古田山常绿阔叶林不同演替群落的萌生特征
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摘要
萌生是植物营养繁殖的重要方式之一,树木的萌生更新对群落结构的维持、植被的动态和演替具有重要意义。本研究基于古田山常绿阔叶林内处于不同演替阶段的25个30 m×30 m样方中胸径≥5 cm的木本植物萌生茎发生情况的调查数据,应用线性回归模型分析了不同演替群落的萌生特征及其地形解释。结果发现:(1)样地内共有56种木本植物存在萌生现象,占总物种数的53.8%。(2)群落(样方)水平的萌生能力与海拔显著负相关,与坡度和坡向的相关性较弱。(3)群落水平的萌生能力与林分年龄显著负相关;在控制地形因子的影响下,林分年龄对萌生指标解释量仍然达到20%–30%。萌生个体数和萌生物种丰富度所占比例随林分年龄显著下降,萌生物种主茎的平均胸径随林分年龄显著增加。研究表明,古田山常绿阔叶林萌生能力随林分年龄的增大而下降,可能由演替序列上物种组成的变化(即萌生能力较强物种的多度下降)和群落结构的变化(即大径级个体增多)导致。另外,萌生能力受到海拔、坡向等地形因素的影响。
Sprouting is an important vegetative reproduction strategy. Sprouting in woody plants is especially of great significance for maintaining forest structure and influencing vegetation dynamics and succession. In this study, we collected sprout data from 25 plots(30 m × 30 m) in an evergreen broad-leaved forest on Gutian Mountain, Zhejiang Province, East China for all trees with diameter at breast height(DBH) equal to or greater than 5 cm. Based on this data, we used general linear regression models to explore the characteristics and the effects of topographic factors and forest age on sprouting ability in forest communities during succession. We found that(1) 53.8%(56/104) of species in this forest showed evidence of sprouting.(2) The plot-level sprouting ability decreased significantly with elevation and marginally decreased significantly with aspect.(3) The plot-level sprouting ability decreased significantly with forest age. Forest age explained 20%–30% of the variation in sprouting ability after being fitted with the models using topographical factors. Both the number of sprouting genets and the proportion of sprouted species richness decreased significantly, while the mean DBH of sprouting genets increased significantly with forest age. This study shows that a decrease in community sprouting ability coupled with an increase in forest age may result from changes in both species composition(i.e. a decreasing number of individuals, but a higher sprouting ability) and structure(i.e. an increasing in proportion of larger individuals) during forest succession. In addition, the sprouting ability of forest communities is also affected by topographical factors, such as elevation and aspect.
Sprouting is an important vegetative reproduction strategy. Sprouting in woody plants is especially of great significance for maintaining forest structure and influencing vegetation dynamics and succession. In this study, we collected sprout data from 25 plots(30 m × 30 m) in an evergreen broad-leaved forest on Gutian Mountain, Zhejiang Province, East China for all trees with diameter at breast height(DBH) equal to or greater than 5 cm. Based on this data, we used general linear regression models to explore the characteristics and the effects of topographic factors and forest age on sprouting ability in forest communities during succession. We found that(1) 53.8%(56/104) of species in this forest showed evidence of sprouting.(2) The plot-level sprouting ability decreased significantly with elevation and marginally decreased significantly with aspect.(3) The plot-level sprouting ability decreased significantly with forest age. Forest age explained 20%–30% of the variation in sprouting ability after being fitted with the models using topographical factors. Both the number of sprouting genets and the proportion of sprouted species richness decreased significantly, while the mean DBH of sprouting genets increased significantly with forest age. This study shows that a decrease in community sprouting ability coupled with an increase in forest age may result from changes in both species composition(i.e. a decreasing number of individuals, but a higher sprouting ability) and structure(i.e. an increasing in proportion of larger individuals) during forest succession. In addition, the sprouting ability of forest communities is also affected by topographical factors, such as elevation and aspect.
引文
Bellingham PJ,Sparrow AD(2000)Resprouting as a life history strategy in woody plant communities.Oikos,89,409-416.
Bellingham PJ,Sparrow AD(2009)Multi-stemmed trees in montane rain forests:Their frequency and demography in relation to elevation,soil nutrients and disturbance.Journal of Ecology,97,472-483.
Bond WJ,Midgley JJ(2001)Ecology of sprouting in woody plants:The persistence niche.Trends in Ecology and Evolution,16,45-51.
Bond WJ,Midgley JJ(2003)The evolutionary ecology of sprouting in woody plants.International Journal of Plant Sciences,164,S103-S114.
Bruelheide H,B?hnke M,Both S,Fang T,Assmann T,Baruffol M,Bauhus J,Buscot F,Chen XY,Ding BY,Durka W,Erfmeier A,Fischer M,Geissler C,Guo DL,Guo LD,Hardtle W,He JS,Hector A,Krober W,Kuhn P,Lang AC,Nadrowski K,Pei KQ,Scherer-Lorenzen M,Shi XZ,Scholten T,Schuldt A,Trogisch S,von Oheimb G,Welk E,Wirth C,Wu YT,Yang XF,Zeng XQ,Zhang SR,Zhou HZ,Ma KP,Schmid B(2011)Community assembly during secondary forest succession in a Chinese subtropical forest.Ecological Monographs,81,25-41.
Chen M,Cao M,Lin LX(2007)Research advances in regeneration of woody plants by sprouting.Chinese Journal of Ecology,26,1114-1118.(in Chinese with English abstract)[陈沐,曹敏,林露湘(2007)木本植物萌生更新研究进展.生态学杂志,26,1114-1118.]
Clarke PJ,Lawes MJ,Midgley JJ(2010)Resprouting as a key functional trait in woody plants-challenges to developing new organizing principles.New Phytologist,188,651-654.
Del Tredici P(2001)Sprouting in temperate trees:A morphological and ecological review.Botanical Review,67,121-140.
Fang JY,Shen ZH,Cui HT(2004)Ecological characteristics of mountains and research issues of mountain ecology.Biodiversity Science,12,10-19.(in Chinese with English abstract)[方精云,沈泽昊,崔海亭(2004)试论山地的生态特征及山地生态学的研究内容.生物多样性,12,10-19.]
Gower ST,Mcmurtrie RE,Murty D(1996)Aboveground net primary production decline with stand age:Potential causes.Trends in Ecology and Evolution,11,378-382.
Guo YL,Wang B,Xiang WS,Ding T,Lu SH,Wen SJ,Huang FZ,Li DX,Li XK(2015)Sprouting characteristics of tree species in 15-hm2 plot of northern tropical karst seasonal rain forest in Nonggang,Guangxi,southern China.Chinese Journal of Ecology,34,955-961.(in Chinese with English abstract)[郭屹立,王斌,向悟生,丁涛,陆树华,文淑均,黄甫昭,李冬兴,李先琨(2015)弄岗北热带喀斯特季节性雨林15 hm2样地木本植物萌生特征.生态学杂志,34,955-961.]
He D,Wei XZ,Li LF,Jiang MX,Yang JY,Yu J(2009)Population structure and dynamics of Cercidiphyllum japonicum in riparian zones of the Shennongjia Mountains region,central China.Chinese Journal of Plant Ecology,33,469-481.(in Chinese with English abstract)[何东,魏新增,李连发,江明喜,杨敬元,喻杰(2009)神农架山地河岸带连香树的种群结构与动态.植物生态学报,33,469-481.]
Hu ZH,Yu MJ(2006)Species diversity in four successional communities of evergreen broad-leaved forest in Gutian Mountain,Zhejiang Province.Chinese Journal of Ecology,25,603-606.(in Chinese with English abstract)[胡正华,于明坚(2006)浙江古田山常绿阔叶林演替序列研究:群落物种多样性.生态学杂志,25,603-606.]
Kammesheidt L(1998)The role of tree sprouts in the restoration of stand structure and species diversity in tropical moist forest after slash-and-burn agriculture in Eastern Paraguay.Plant Ecology,139,155-165.
Lang AC,Hardtle W,Baruffol M,B?hnke M,Bruelheide H,Schmid B,von Wehrden H,von Oheimb G(2012)Mechanisms promoting tree species co-existence:Experimental evidence with saplings of subtropical forest ecosystems of China.Journal of Vegetation Science,23,837-846.
Liu CX,Jin Y,Yu JP,Chen SW,Tian L,Wang YQ,Chen JH(2014)Characteristics of root sprouting trees of Castanopsis eyrei and Schima superba communities in 1 hm2 forest plot at Chawan,Gutianshan National Nature Reserve.Journal of Zhejiang University(Science Edition),41,573-582,592.(in Chinese with English abstract)[刘常幸,金毅,余建平,陈声文,田磊,王云泉,陈建华(2014)古田山茶湾样地甜槠-木荷林根萌特征分析.浙江大学学报(理学版),41,573-582,592.]
Liu HB,Wang QG,Lu JM,Xu YZ,Lu ZJ,Qiao XJ,Bao DC,Guo YL,Meng HJ,Jiang MX(2014)Root-sprouting ability in an evergreen and deciduous broad-leaved mixed forest.Chinese Science Bulletin(Chinese Version),59,3491-3498.(in Chinese with English abstract)[刘海波,王庆刚,路俊盟,徐耀粘,卢志军,乔秀娟,鲍大川,郭屹立,孟红杰,江明喜(2014)八大公山常绿落叶阔叶混交林根萌能力.科学通报,59,3491-3498.]
Nanami S,Kawaguchi H,Tateno R,Li CH,Katagiri S(2004)Sprouting traits and population structure of co-occurring Castanopsis species in an evergreen broad-leaved forest in southern China.Ecological Research,19,341-348.
Paula S,CervellóCP,Pausas JG(2006)Fire as a germination cue:A review for the Mediterranean Basin.Forest Ecology and Management,234,S176.
Paula S,Pausas JG(2006)Leaf traits and resprouting ability in the Mediterranean Basin.Functional Ecology,20,941-947.
Paula S,Arianoutsou M,Kazanis D,Tavsanoglu C,Lloret F,Buhk C,Ojeda F,Luna B,Moreno JM,Rodrigo A,Espelta JM,Palacio S,Fernández-Santos B,Fernandes PM,Pausas JG(2009)Fire-related traits for plant species of the Mediterranean Basin.Ecology,90,1420.
Pausas JG(2001)Resprouting vs.seeding:A Mediterranean perspective.Oikos,94,193-194.
R Core Team(2016)R:A Language and Environment for Statistical Computing.R Foundation for Statistical Computing,Vienna,Austria.https://www.R-project.org/.
Sandel B,Svenning JC(2013)Human impacts drive a global topographic signature in tree cover.Nature Communications,4,1-7.
Shen YX,Yang GR,Huang J(2011)Comparison of tree sprouting in three regeneration stages of an evergreen broadleaved forest in a karst landscape,SW China.Acta Ecologica Sinica,31,126-132.
Su JR,Liu WD,Zhang ZJ,Li SF(2012)Sprouting characteristic in restoration ecosystems of monsoon evergreen broad-leaved forest in south-central of Yunnan Province.Acta Ecologica Sinica,32,805-814.(in Chinese with English abstract)[苏建荣,刘万德,张志钧,李帅峰(2012)云南中南部季风常绿阔叶林恢复生态系统萌生特征.生态学报,32,805-814.]
VerdúM(2000)Ecological and evolutionary differences between Mediterranean seeders and resprouters.Journal of Vegetation Science,11,265-268.
Wang XH,Kent M,Fang XF(2007)Evergreen broad-leaved forest in Eastern China:Its ecology and conservation and the importance of resprouting in forest restoration.Forest Ecology and Management,245,76-87.
Yan ER,Wang XH,Shi JY,Wang XB,Wang LY(2005)Sprouting ecology of woody plants:A research review.Chinese Journal of Applied Ecology,16,2459-2464.(in Chinese with English abstract)[闫恩荣,王希华,施家月,王希波,王良衍(2005)木本植物萌枝生态学研究进展.应用生态学报,16,2459-2464.]
Ye D,Dong RR,Mi XC,Lu W,Zheng ZJ,Yu MJ,Ni J,Chen JH(2017)Characteristics and effects of sprouting on species diversity in a subtropical evergreen broad-leaved forest in Gutianshan,East China.Biodiversity Science,25,393-400.(in Chinese with English abstract)[叶铎,董瑞瑞,米湘成,芦伟,郑振杰,于明坚,倪健,陈建华(2017)古田山常绿阔叶林萌生特征及其与群落物种多样性的关系.生物多样性,25,393-400.]
Ye D,Qian HY,Wang LY,Jin FM,Ni J,Chen SW,Song YF,Chen JH(2018)Sprouting characteristics of woody species in a subtropical evergreen broad-leaved forest in Gutianshan of Qianjiangyuan National Park,East China.Acta Ecologica Sinica,38,3562-3568.(in Chinese with English abstract)[叶铎,钱海源,王璐瑶,金芳梅,倪健,陈声文,宋云峰,陈建华(2018)钱江源国家公园古田山常绿阔叶林木本植物的萌生更新特征.生态学报,38,3562-3568.]
Ye J,Hao ZQ,Wang XG,Bai XJ,Xing DL,Yuan ZQ(2014)Local-scale drivers of multi-stemmed tree formation in Acer,in a temperate forest of Northeast China.Chinese Science Bulletin,59,320-325.
Zhou SX,Jiang MX,Bao DC,Tao M,Huang HD(2011)Population structure and regenerative characteristics of major tree species of rare plant community in Houhe Nature Reserve.Guihaia,31,209-216.(in Chinese with English abstract)[周赛霞,江明喜,鲍大川,陶敏,黄汉东(2011)后河自然保护区珍稀植物群落结构及更新特性.广西植物,31,209-216.]
(1)田磊(2015)古田山5 ha样地木本根萌植物影响因子数值分析.硕士学位论文,浙江师范大学,浙江金华.
Bellingham PJ,Sparrow AD(2009)Multi-stemmed trees in montane rain forests:Their frequency and demography in relation to elevation,soil nutrients and disturbance.Journal of Ecology,97,472-483.
Bond WJ,Midgley JJ(2001)Ecology of sprouting in woody plants:The persistence niche.Trends in Ecology and Evolution,16,45-51.
Bond WJ,Midgley JJ(2003)The evolutionary ecology of sprouting in woody plants.International Journal of Plant Sciences,164,S103-S114.
Bruelheide H,B?hnke M,Both S,Fang T,Assmann T,Baruffol M,Bauhus J,Buscot F,Chen XY,Ding BY,Durka W,Erfmeier A,Fischer M,Geissler C,Guo DL,Guo LD,Hardtle W,He JS,Hector A,Krober W,Kuhn P,Lang AC,Nadrowski K,Pei KQ,Scherer-Lorenzen M,Shi XZ,Scholten T,Schuldt A,Trogisch S,von Oheimb G,Welk E,Wirth C,Wu YT,Yang XF,Zeng XQ,Zhang SR,Zhou HZ,Ma KP,Schmid B(2011)Community assembly during secondary forest succession in a Chinese subtropical forest.Ecological Monographs,81,25-41.
Chen M,Cao M,Lin LX(2007)Research advances in regeneration of woody plants by sprouting.Chinese Journal of Ecology,26,1114-1118.(in Chinese with English abstract)[陈沐,曹敏,林露湘(2007)木本植物萌生更新研究进展.生态学杂志,26,1114-1118.]
Clarke PJ,Lawes MJ,Midgley JJ(2010)Resprouting as a key functional trait in woody plants-challenges to developing new organizing principles.New Phytologist,188,651-654.
Del Tredici P(2001)Sprouting in temperate trees:A morphological and ecological review.Botanical Review,67,121-140.
Fang JY,Shen ZH,Cui HT(2004)Ecological characteristics of mountains and research issues of mountain ecology.Biodiversity Science,12,10-19.(in Chinese with English abstract)[方精云,沈泽昊,崔海亭(2004)试论山地的生态特征及山地生态学的研究内容.生物多样性,12,10-19.]
Gower ST,Mcmurtrie RE,Murty D(1996)Aboveground net primary production decline with stand age:Potential causes.Trends in Ecology and Evolution,11,378-382.
Guo YL,Wang B,Xiang WS,Ding T,Lu SH,Wen SJ,Huang FZ,Li DX,Li XK(2015)Sprouting characteristics of tree species in 15-hm2 plot of northern tropical karst seasonal rain forest in Nonggang,Guangxi,southern China.Chinese Journal of Ecology,34,955-961.(in Chinese with English abstract)[郭屹立,王斌,向悟生,丁涛,陆树华,文淑均,黄甫昭,李冬兴,李先琨(2015)弄岗北热带喀斯特季节性雨林15 hm2样地木本植物萌生特征.生态学杂志,34,955-961.]
He D,Wei XZ,Li LF,Jiang MX,Yang JY,Yu J(2009)Population structure and dynamics of Cercidiphyllum japonicum in riparian zones of the Shennongjia Mountains region,central China.Chinese Journal of Plant Ecology,33,469-481.(in Chinese with English abstract)[何东,魏新增,李连发,江明喜,杨敬元,喻杰(2009)神农架山地河岸带连香树的种群结构与动态.植物生态学报,33,469-481.]
Hu ZH,Yu MJ(2006)Species diversity in four successional communities of evergreen broad-leaved forest in Gutian Mountain,Zhejiang Province.Chinese Journal of Ecology,25,603-606.(in Chinese with English abstract)[胡正华,于明坚(2006)浙江古田山常绿阔叶林演替序列研究:群落物种多样性.生态学杂志,25,603-606.]
Kammesheidt L(1998)The role of tree sprouts in the restoration of stand structure and species diversity in tropical moist forest after slash-and-burn agriculture in Eastern Paraguay.Plant Ecology,139,155-165.
Lang AC,Hardtle W,Baruffol M,B?hnke M,Bruelheide H,Schmid B,von Wehrden H,von Oheimb G(2012)Mechanisms promoting tree species co-existence:Experimental evidence with saplings of subtropical forest ecosystems of China.Journal of Vegetation Science,23,837-846.
Liu CX,Jin Y,Yu JP,Chen SW,Tian L,Wang YQ,Chen JH(2014)Characteristics of root sprouting trees of Castanopsis eyrei and Schima superba communities in 1 hm2 forest plot at Chawan,Gutianshan National Nature Reserve.Journal of Zhejiang University(Science Edition),41,573-582,592.(in Chinese with English abstract)[刘常幸,金毅,余建平,陈声文,田磊,王云泉,陈建华(2014)古田山茶湾样地甜槠-木荷林根萌特征分析.浙江大学学报(理学版),41,573-582,592.]
Liu HB,Wang QG,Lu JM,Xu YZ,Lu ZJ,Qiao XJ,Bao DC,Guo YL,Meng HJ,Jiang MX(2014)Root-sprouting ability in an evergreen and deciduous broad-leaved mixed forest.Chinese Science Bulletin(Chinese Version),59,3491-3498.(in Chinese with English abstract)[刘海波,王庆刚,路俊盟,徐耀粘,卢志军,乔秀娟,鲍大川,郭屹立,孟红杰,江明喜(2014)八大公山常绿落叶阔叶混交林根萌能力.科学通报,59,3491-3498.]
Nanami S,Kawaguchi H,Tateno R,Li CH,Katagiri S(2004)Sprouting traits and population structure of co-occurring Castanopsis species in an evergreen broad-leaved forest in southern China.Ecological Research,19,341-348.
Paula S,CervellóCP,Pausas JG(2006)Fire as a germination cue:A review for the Mediterranean Basin.Forest Ecology and Management,234,S176.
Paula S,Pausas JG(2006)Leaf traits and resprouting ability in the Mediterranean Basin.Functional Ecology,20,941-947.
Paula S,Arianoutsou M,Kazanis D,Tavsanoglu C,Lloret F,Buhk C,Ojeda F,Luna B,Moreno JM,Rodrigo A,Espelta JM,Palacio S,Fernández-Santos B,Fernandes PM,Pausas JG(2009)Fire-related traits for plant species of the Mediterranean Basin.Ecology,90,1420.
Pausas JG(2001)Resprouting vs.seeding:A Mediterranean perspective.Oikos,94,193-194.
R Core Team(2016)R:A Language and Environment for Statistical Computing.R Foundation for Statistical Computing,Vienna,Austria.https://www.R-project.org/.
Sandel B,Svenning JC(2013)Human impacts drive a global topographic signature in tree cover.Nature Communications,4,1-7.
Shen YX,Yang GR,Huang J(2011)Comparison of tree sprouting in three regeneration stages of an evergreen broadleaved forest in a karst landscape,SW China.Acta Ecologica Sinica,31,126-132.
Su JR,Liu WD,Zhang ZJ,Li SF(2012)Sprouting characteristic in restoration ecosystems of monsoon evergreen broad-leaved forest in south-central of Yunnan Province.Acta Ecologica Sinica,32,805-814.(in Chinese with English abstract)[苏建荣,刘万德,张志钧,李帅峰(2012)云南中南部季风常绿阔叶林恢复生态系统萌生特征.生态学报,32,805-814.]
VerdúM(2000)Ecological and evolutionary differences between Mediterranean seeders and resprouters.Journal of Vegetation Science,11,265-268.
Wang XH,Kent M,Fang XF(2007)Evergreen broad-leaved forest in Eastern China:Its ecology and conservation and the importance of resprouting in forest restoration.Forest Ecology and Management,245,76-87.
Yan ER,Wang XH,Shi JY,Wang XB,Wang LY(2005)Sprouting ecology of woody plants:A research review.Chinese Journal of Applied Ecology,16,2459-2464.(in Chinese with English abstract)[闫恩荣,王希华,施家月,王希波,王良衍(2005)木本植物萌枝生态学研究进展.应用生态学报,16,2459-2464.]
Ye D,Dong RR,Mi XC,Lu W,Zheng ZJ,Yu MJ,Ni J,Chen JH(2017)Characteristics and effects of sprouting on species diversity in a subtropical evergreen broad-leaved forest in Gutianshan,East China.Biodiversity Science,25,393-400.(in Chinese with English abstract)[叶铎,董瑞瑞,米湘成,芦伟,郑振杰,于明坚,倪健,陈建华(2017)古田山常绿阔叶林萌生特征及其与群落物种多样性的关系.生物多样性,25,393-400.]
Ye D,Qian HY,Wang LY,Jin FM,Ni J,Chen SW,Song YF,Chen JH(2018)Sprouting characteristics of woody species in a subtropical evergreen broad-leaved forest in Gutianshan of Qianjiangyuan National Park,East China.Acta Ecologica Sinica,38,3562-3568.(in Chinese with English abstract)[叶铎,钱海源,王璐瑶,金芳梅,倪健,陈声文,宋云峰,陈建华(2018)钱江源国家公园古田山常绿阔叶林木本植物的萌生更新特征.生态学报,38,3562-3568.]
Ye J,Hao ZQ,Wang XG,Bai XJ,Xing DL,Yuan ZQ(2014)Local-scale drivers of multi-stemmed tree formation in Acer,in a temperate forest of Northeast China.Chinese Science Bulletin,59,320-325.
Zhou SX,Jiang MX,Bao DC,Tao M,Huang HD(2011)Population structure and regenerative characteristics of major tree species of rare plant community in Houhe Nature Reserve.Guihaia,31,209-216.(in Chinese with English abstract)[周赛霞,江明喜,鲍大川,陶敏,黄汉东(2011)后河自然保护区珍稀植物群落结构及更新特性.广西植物,31,209-216.]
(1)田磊(2015)古田山5 ha样地木本根萌植物影响因子数值分析.硕士学位论文,浙江师范大学,浙江金华.