广西弄岗北热带喀斯特季节性雨林林隙特征与空间分布
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摘要
林隙是森林中一种普遍存在的小尺度森林干扰形式,对促进森林更新和循环演替具有重要作用。对热带喀斯特森林的林隙干扰目前还了解不多。该文采用Worldview-3高空间分辨率遥感影像结合地面调查,对弄岗喀斯特季节性雨林15 hm~2样地林隙干扰特征及其空间分布规律进行了研究。结果表明:样地内共解译到305个林隙,林隙的密度约为每公顷20个,林隙大小的变化范围为10～371 m~2,平均大小为52.3m~2,林隙面积占森林面积的10.6%;林隙在0～50 m尺度整体表现出聚集分布格局;林隙空间分布与大树(DBH≥10 cm)存在显著空间关联,与小树(DBH<10 cm)和枯立木无显著空间关联;不同生态因子对林隙的数量分布和基本结构影响不同,对林隙数量分布影响最显著的包括凹凸度、干旱度指数、丰富度,而坡向、海拔、丰富度对林隙大小有显著影响,海拔、岩石裸露度、地形湿润度指数、干旱度指数、多度等5个生态因子对林隙周长有显著影响。研究结果表明林隙在喀斯特季节性雨林普遍存在,林隙的空间分布与特征存在着地形分异特征,是喀斯特独特的地质地貌和生境条件与喀斯特森林自身生物学特性综合作用的结果。
Forest gap is the dominant form of disturbance in many forest ecosystems.Gaps play an important role in forest helping to maintain forest regeneration and dynamics.Little is known about the gap pattern and characteristics of karst seasonal rain forests.In this study,high-spatial-resolution Worldview-3 satellite imagery was coupled with field data to assess gap disturbance regime and its pattern in a 15 tropical karst seasonal rain forest dynamic plot in Nonggang National Nature Reserve,Guangxi.Total 305 gaps were identified,ranging in size from 10 to 371with mean size 52.3 m~2.The canopy gap fraction was 10.6%,and gap density was about 20 per hectare.The spatial pattern of gaps was clustered at scale 0-50 m.Significant association was found between gaps and big trees(DBH≥10 cm),and null associations between gaps and small trees(DBH<10 cm)and between gaps and standing dead trees.The prevailing ecological factors had different influence on gap distribution and gap characteristics.Convexity,altitude above channel(AAH)and tree species richness had significant effect on the gaps distribution.Gap size was significantly influenced by slope,mean elevation and tree species richness.Gap perimeter was mainly affected by mean elevation,rock-bareness rate(RBR),topographic wetness index(TWI),AAH and tree species abundance.These results indicated that gap disturbance was very frequent in karst seasonal rain forests,and there were topographic differentiation in spatial distribution and characteristics of gaps resulted by the combination effects of unique special karst geological conditions,habitats and the biological characteristics of karst forest.
Forest gap is the dominant form of disturbance in many forest ecosystems.Gaps play an important role in forest helping to maintain forest regeneration and dynamics.Little is known about the gap pattern and characteristics of karst seasonal rain forests.In this study,high-spatial-resolution Worldview-3 satellite imagery was coupled with field data to assess gap disturbance regime and its pattern in a 15 tropical karst seasonal rain forest dynamic plot in Nonggang National Nature Reserve,Guangxi.Total 305 gaps were identified,ranging in size from 10 to 371with mean size 52.3 m~2.The canopy gap fraction was 10.6%,and gap density was about 20 per hectare.The spatial pattern of gaps was clustered at scale 0-50 m.Significant association was found between gaps and big trees(DBH≥10 cm),and null associations between gaps and small trees(DBH<10 cm)and between gaps and standing dead trees.The prevailing ecological factors had different influence on gap distribution and gap characteristics.Convexity,altitude above channel(AAH)and tree species richness had significant effect on the gaps distribution.Gap size was significantly influenced by slope,mean elevation and tree species richness.Gap perimeter was mainly affected by mean elevation,rock-bareness rate(RBR),topographic wetness index(TWI),AAH and tree species abundance.These results indicated that gap disturbance was very frequent in karst seasonal rain forests,and there were topographic differentiation in spatial distribution and characteristics of gaps resulted by the combination effects of unique special karst geological conditions,habitats and the biological characteristics of karst forest.
引文
ALEXANDER HD,MACK MC,2017.Gap regeneration within mature deciduous forests of Interior Alaska:Implications for future forest change[J].For Ecol Manag,396(2017):35-43.
BADDELEY A,DIGGLE PJ,HARDEGEN A,et al.,2014.On tests of spatial pattern based on simulation envelopes[J].Ecol Monogr,84(3):477-489.
BROKAW NVL,1982.The definition of treefall gap and its effect on measures of forest dynamics[J].Biotropica,14(2):158-160.
BURNHAM KP,ANDERSON DR,2002.Model selection and inference:A practical information-theoretic approach[M].New York:Springer:149-199.
GARBARINO M,BORGOGNO MONDINO E,LINGUAE,et al.,2012.Gap disturbances and regeneration patterns in a Bosnian old-growth forest:A multispectral remote sensing and groundbased approach[J].Ann For Sci,69(5):617-625.
GOULAMOUSSNE Y,BEDEAU C,DESCROIX L,et al.,2017.Environmental control of natural gap size distribution in tropical forests[J].Biogeosciences,14(2):1-18.
GUO K,LIU CC,DONG M,2011.Ecological adaptation of plants and control of rocky-desertification on karst region of Southwest China[J].Chin J Plant Ecol,35(10):991-999.[郭柯,刘长成,董鸣,2011.我国西南喀斯特植物生态适应性与石漠化治理[J].植物生态学报,35(10):991-999.]
GUO YL,WANG B,XIANG WS,et al.,2016.Responses of spatial pattern of woody plants’basal area to topographic factors in a tropical karst seasonal rainforest in Nonggang,Guangxi,southern China[J].Biodivers Sci,24(1):30-39.[郭屹立,王斌,向悟生,等,2016.喀斯特季节性雨林木本植物胸高断面积分布格局及其对地形因子的响应[J].生物多样性,24(1):30-39.]
HOBI ML,GINZLER C,COMMARMOT B,et al.,2015.Gap pattern of the largest primeval beech forest of Europe revealed by remote sensing[J].Ecosphere,6(5):t15-t76.
HU LL,LI JS,WU XP,et al.,2010.Rewiews on methods of measuring geometric chracteristics of forest gaps involving gap size,gap shape,and the height of canopy trees surrounding the gap[J].Acta Ecol Sin,30(7):1911-1919.[胡理乐,李俊生,吴晓莆,等,2010.林窗几何特征的测定方法[J].生态学报,30(7):1911-1919.]
HUANG FZ,WANG B,DING T,et al.,2014.Numerical classification of associations in a northern tropical karst seasonal rain forest and the relationships of these associations with environmental factors[J].Biodivers Sci,22(2):157-166.[黄甫昭,王斌,丁涛,等,2014.弄岗北热带喀斯特季节性雨林群丛数量分类及与环境的关系[J].生物多样性,22(2):157-166.]
HUBBELL SP,FOSTER RB,O'BRIEN ST,et al.,1999.Light gap disturbances,recruitment limitation,and tree diversity in a neotropical forest[J].Science,283(5401):554-557.
KATHKE S,BRUELHEID EH,2010.Gap dynamics in a nearnatural spruce forest at Mt.Brocken,Germany[J].For Ecol Manag,259(3):624-632.
KIAN S,TABARI M,ESMAILZADEHO,et al.,2017.Gap characteristics and disturbance regime in an intact hyrcanian oriental beech forest,iran[J].Austr J For Sci,134(4):323-346.
LIU SC,HUA WJ,DUANWB,et al.,2013.Gap characteristics in the mixed broad-leaved Korean pine forest in Xiaoxing'an Mountains[J].Acta Ecol Sin,33(17):5234-5244.[刘少冲,王敬华,段文标,等,2013.小兴安岭阔叶红松混交林林隙特征[J].生态学报,33(17):5234-5244.]
LONG CL,YU SX,WEI LM,et al.,2005.Disturbance regimes and the characteristics of gaps in Maolan Karst Forest,Guizhou Province[J].Sci Silv Sin,41(4):13-19.[龙翠玲,余世孝,魏鲁明,等,2005.茂兰喀斯特森林干扰状况与林隙特征[J].林业科学,41(4):13-19.]
LOOSMORE NB,FORD ED,2006.Statistical inference using the g or k point pattern spatial statistics[J].Ecology,87(8):1925-1931.
MAO XG,DU ZH,LIU JQ,et al.,2018.Object-oriented segmentation and classification of forest gap based on QuickBird remote sensing image[J].Chin J Appl Ecol,29(1):1-10.[毛学刚,杜子涵,刘家倩,等,2018.基于面向对象的QuickBird遥感影像林隙分割与分类[J].应用生态学报,29(1):1-10.]
MAO XG,HOU JY,BAI XF,et al.,2017.Multiscale forest gap segmentation and object-oriented classification based on DOM and LiDAR[J].T Chin Soc Agric Mach,48(9):152-159.[毛学刚,侯吉宇,白雪峰,等,2017.基于DOM及LiDAR的多尺度分割与面向对象林隙分类[J].农业机械学报,48(9):152-159.]
MUSCOLO A,BAGNATO S,SIDARI M,et al.,2014.A review of the roles of forest canopy gaps[J].J For Res,25(4):725-736.
R CORE TEAM D.R:A language and environment for statistical computing[EB/OL].URL https://www.R-project.org/.
RUNKLE JR,1982.Patterns of disturbance in some OldGrowth mesic forests of eastern north america[J].Ecology,63(5):1533-1546.
SCHLIEMANN SA,BOCKHEIM JG,2011.Methods for studying treefall gaps:A review[J].For Ecol Manag,261(7):1143-1151.
SENCAL J,DOYON F,MESSIERC,2018.Tree death not resulting in gap creation:An investigation of canopy dynamics of northern temperate deciduous forests[J].Remote SensBasel,10(1):121.
SHEN ZH,LI DX,WANG GF,2001.Studies on the gap disturbance of the mountain mixed forests of evergreen and deciduous broad-leaved species at mt.Dalaoling in the three gorgesⅠ.The essential characteristics of the gaps[J].Acta Phytoecol Sin,25(3):276-282.[沈泽昊,李道兴,王功芳,2001.三峡大老岭山地常绿落叶阔叶混交林林隙干扰研究I林隙基本特征[J].植物生态学报,25(3):276-282.]
SONG CH,DICKINSON MB,SU LH,et al.,2010.Estimating average tree crown size using spatial information from Ikonos and QuickBird images:Across-sensor and across-site comparisons[J].Remote Sens Environ,114(5):1099-1107.
VEPAKOMMA U,ST-ONGE B,KNEESHAWD,2008.Spatially explicit characterization of boreal forest gap dynamics using multi-temporal lidar data[J].Remote Sens Environ,112(5):2326-2340.
WANG B,HUANG YS,LI XK,et al.,2014.Species composition and spatial distribution of the 15 ha northern tropical karst seasonal rain forest dynamic plot in Nonggang of Guangxi,Southern China[J].Biodivers Sci,22(2):141-156.[王斌,黄俞淞,李先琨,等,2014.弄岗北热带喀斯特季节性雨林15 ha监测样地的树种组成与空间分布[J].生物多样性,22(2):141-156.]
WANG B,XIANG WS,DING T,et al.,2014.Spatial distribution of standing dead trees abundance and its impact factors in the karst seasonal rain forest,Nonggang,southern China[J].Chin Sci Bull,59(35):3479-3490.[王斌,向悟生,丁涛,等,2014.弄岗喀斯特季节性雨林枯立木多度的空间分布及影响因子[J].科学通报,59(35):3479-3490.]
WEN YG,LIN JY,ZHU HG,et al.,2014.Gaps and their natural disturbance regimes in the south subtropical montane evergreen broadleaved forest[J].Guangxi Sci,21(5):447-453.[温远光,林建勇,朱宏光,等,2014.南亚热带山地常绿阔叶林林隙及其自然干扰特征研究[J].广西科学,21(5):447-453.]
WIEGAND T,MOLONEY KA,2004.Rings,circles,and nullmodels for point pattern analysis in ecology[J].Oikos,104(2):209-229.
WU QG,WU FZ,YANG WQ,et al.,2013.Characteristics of gaps and disturbance regimes of the alpine fir forest in Western Sichuan[J].Chin J Appl Environ Biol,19(6):922-928.[吴庆贵,吴福忠,杨万勤,等,2013.川西高山森林林隙特征及干扰状况[J].应用与环境生物学报,19(6):922-928.]
XIANG WS,LI XK,WANG B,2017.A study of mapping forest monitoring plot in rugged topography of South China[J].Guangxi Sci,24(3):219-224.[向悟生,李先琨,王斌,2017.中国南方复杂地形森林样地的测量标定方法[J].广西科学,24(3):219-224.]
YAN SJ,HONG W,WU CZ,et al.,2004.Gaps and their natural disturbance characteristics in mid-subtropical evergreen broad-leaved forest in Wanmulin[J].Chin J Appl Ecol,15(7):1126-1130.[闫淑君,洪伟,吴承祯,等,2004.中亚热带常绿阔叶林林隙及其自然干扰特征的研究[J].应用生态学报,15(7):1126-1130.]
ZANG RG,YANG YC,1999.Gaps and their natural disturbance regimes in the tropical montane rain forest of Hainan Island[J].Sci Silv Sin,35(1):1-8.[臧润国,杨彦承,1999.海南岛热带山地雨林林隙及其自然干扰特征[J].林业科学,35(1):1-8.]
BADDELEY A,DIGGLE PJ,HARDEGEN A,et al.,2014.On tests of spatial pattern based on simulation envelopes[J].Ecol Monogr,84(3):477-489.
BROKAW NVL,1982.The definition of treefall gap and its effect on measures of forest dynamics[J].Biotropica,14(2):158-160.
BURNHAM KP,ANDERSON DR,2002.Model selection and inference:A practical information-theoretic approach[M].New York:Springer:149-199.
GARBARINO M,BORGOGNO MONDINO E,LINGUAE,et al.,2012.Gap disturbances and regeneration patterns in a Bosnian old-growth forest:A multispectral remote sensing and groundbased approach[J].Ann For Sci,69(5):617-625.
GOULAMOUSSNE Y,BEDEAU C,DESCROIX L,et al.,2017.Environmental control of natural gap size distribution in tropical forests[J].Biogeosciences,14(2):1-18.
GUO K,LIU CC,DONG M,2011.Ecological adaptation of plants and control of rocky-desertification on karst region of Southwest China[J].Chin J Plant Ecol,35(10):991-999.[郭柯,刘长成,董鸣,2011.我国西南喀斯特植物生态适应性与石漠化治理[J].植物生态学报,35(10):991-999.]
GUO YL,WANG B,XIANG WS,et al.,2016.Responses of spatial pattern of woody plants’basal area to topographic factors in a tropical karst seasonal rainforest in Nonggang,Guangxi,southern China[J].Biodivers Sci,24(1):30-39.[郭屹立,王斌,向悟生,等,2016.喀斯特季节性雨林木本植物胸高断面积分布格局及其对地形因子的响应[J].生物多样性,24(1):30-39.]
HOBI ML,GINZLER C,COMMARMOT B,et al.,2015.Gap pattern of the largest primeval beech forest of Europe revealed by remote sensing[J].Ecosphere,6(5):t15-t76.
HU LL,LI JS,WU XP,et al.,2010.Rewiews on methods of measuring geometric chracteristics of forest gaps involving gap size,gap shape,and the height of canopy trees surrounding the gap[J].Acta Ecol Sin,30(7):1911-1919.[胡理乐,李俊生,吴晓莆,等,2010.林窗几何特征的测定方法[J].生态学报,30(7):1911-1919.]
HUANG FZ,WANG B,DING T,et al.,2014.Numerical classification of associations in a northern tropical karst seasonal rain forest and the relationships of these associations with environmental factors[J].Biodivers Sci,22(2):157-166.[黄甫昭,王斌,丁涛,等,2014.弄岗北热带喀斯特季节性雨林群丛数量分类及与环境的关系[J].生物多样性,22(2):157-166.]
HUBBELL SP,FOSTER RB,O'BRIEN ST,et al.,1999.Light gap disturbances,recruitment limitation,and tree diversity in a neotropical forest[J].Science,283(5401):554-557.
KATHKE S,BRUELHEID EH,2010.Gap dynamics in a nearnatural spruce forest at Mt.Brocken,Germany[J].For Ecol Manag,259(3):624-632.
KIAN S,TABARI M,ESMAILZADEHO,et al.,2017.Gap characteristics and disturbance regime in an intact hyrcanian oriental beech forest,iran[J].Austr J For Sci,134(4):323-346.
LIU SC,HUA WJ,DUANWB,et al.,2013.Gap characteristics in the mixed broad-leaved Korean pine forest in Xiaoxing'an Mountains[J].Acta Ecol Sin,33(17):5234-5244.[刘少冲,王敬华,段文标,等,2013.小兴安岭阔叶红松混交林林隙特征[J].生态学报,33(17):5234-5244.]
LONG CL,YU SX,WEI LM,et al.,2005.Disturbance regimes and the characteristics of gaps in Maolan Karst Forest,Guizhou Province[J].Sci Silv Sin,41(4):13-19.[龙翠玲,余世孝,魏鲁明,等,2005.茂兰喀斯特森林干扰状况与林隙特征[J].林业科学,41(4):13-19.]
LOOSMORE NB,FORD ED,2006.Statistical inference using the g or k point pattern spatial statistics[J].Ecology,87(8):1925-1931.
MAO XG,DU ZH,LIU JQ,et al.,2018.Object-oriented segmentation and classification of forest gap based on QuickBird remote sensing image[J].Chin J Appl Ecol,29(1):1-10.[毛学刚,杜子涵,刘家倩,等,2018.基于面向对象的QuickBird遥感影像林隙分割与分类[J].应用生态学报,29(1):1-10.]
MAO XG,HOU JY,BAI XF,et al.,2017.Multiscale forest gap segmentation and object-oriented classification based on DOM and LiDAR[J].T Chin Soc Agric Mach,48(9):152-159.[毛学刚,侯吉宇,白雪峰,等,2017.基于DOM及LiDAR的多尺度分割与面向对象林隙分类[J].农业机械学报,48(9):152-159.]
MUSCOLO A,BAGNATO S,SIDARI M,et al.,2014.A review of the roles of forest canopy gaps[J].J For Res,25(4):725-736.
R CORE TEAM D.R:A language and environment for statistical computing[EB/OL].URL https://www.R-project.org/.
RUNKLE JR,1982.Patterns of disturbance in some OldGrowth mesic forests of eastern north america[J].Ecology,63(5):1533-1546.
SCHLIEMANN SA,BOCKHEIM JG,2011.Methods for studying treefall gaps:A review[J].For Ecol Manag,261(7):1143-1151.
SENCAL J,DOYON F,MESSIERC,2018.Tree death not resulting in gap creation:An investigation of canopy dynamics of northern temperate deciduous forests[J].Remote SensBasel,10(1):121.
SHEN ZH,LI DX,WANG GF,2001.Studies on the gap disturbance of the mountain mixed forests of evergreen and deciduous broad-leaved species at mt.Dalaoling in the three gorgesⅠ.The essential characteristics of the gaps[J].Acta Phytoecol Sin,25(3):276-282.[沈泽昊,李道兴,王功芳,2001.三峡大老岭山地常绿落叶阔叶混交林林隙干扰研究I林隙基本特征[J].植物生态学报,25(3):276-282.]
SONG CH,DICKINSON MB,SU LH,et al.,2010.Estimating average tree crown size using spatial information from Ikonos and QuickBird images:Across-sensor and across-site comparisons[J].Remote Sens Environ,114(5):1099-1107.
VEPAKOMMA U,ST-ONGE B,KNEESHAWD,2008.Spatially explicit characterization of boreal forest gap dynamics using multi-temporal lidar data[J].Remote Sens Environ,112(5):2326-2340.
WANG B,HUANG YS,LI XK,et al.,2014.Species composition and spatial distribution of the 15 ha northern tropical karst seasonal rain forest dynamic plot in Nonggang of Guangxi,Southern China[J].Biodivers Sci,22(2):141-156.[王斌,黄俞淞,李先琨,等,2014.弄岗北热带喀斯特季节性雨林15 ha监测样地的树种组成与空间分布[J].生物多样性,22(2):141-156.]
WANG B,XIANG WS,DING T,et al.,2014.Spatial distribution of standing dead trees abundance and its impact factors in the karst seasonal rain forest,Nonggang,southern China[J].Chin Sci Bull,59(35):3479-3490.[王斌,向悟生,丁涛,等,2014.弄岗喀斯特季节性雨林枯立木多度的空间分布及影响因子[J].科学通报,59(35):3479-3490.]
WEN YG,LIN JY,ZHU HG,et al.,2014.Gaps and their natural disturbance regimes in the south subtropical montane evergreen broadleaved forest[J].Guangxi Sci,21(5):447-453.[温远光,林建勇,朱宏光,等,2014.南亚热带山地常绿阔叶林林隙及其自然干扰特征研究[J].广西科学,21(5):447-453.]
WIEGAND T,MOLONEY KA,2004.Rings,circles,and nullmodels for point pattern analysis in ecology[J].Oikos,104(2):209-229.
WU QG,WU FZ,YANG WQ,et al.,2013.Characteristics of gaps and disturbance regimes of the alpine fir forest in Western Sichuan[J].Chin J Appl Environ Biol,19(6):922-928.[吴庆贵,吴福忠,杨万勤,等,2013.川西高山森林林隙特征及干扰状况[J].应用与环境生物学报,19(6):922-928.]
XIANG WS,LI XK,WANG B,2017.A study of mapping forest monitoring plot in rugged topography of South China[J].Guangxi Sci,24(3):219-224.[向悟生,李先琨,王斌,2017.中国南方复杂地形森林样地的测量标定方法[J].广西科学,24(3):219-224.]
YAN SJ,HONG W,WU CZ,et al.,2004.Gaps and their natural disturbance characteristics in mid-subtropical evergreen broad-leaved forest in Wanmulin[J].Chin J Appl Ecol,15(7):1126-1130.[闫淑君,洪伟,吴承祯,等,2004.中亚热带常绿阔叶林林隙及其自然干扰特征的研究[J].应用生态学报,15(7):1126-1130.]
ZANG RG,YANG YC,1999.Gaps and their natural disturbance regimes in the tropical montane rain forest of Hainan Island[J].Sci Silv Sin,35(1):1-8.[臧润国,杨彦承,1999.海南岛热带山地雨林林隙及其自然干扰特征[J].林业科学,35(1):1-8.]