红松存在/不存在数据的多尺度空间分布格局
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摘要
红松属小兴安岭地区地带性植被优势种,该地区也是其分布的北缘。在景观尺度上开展红松的分布格局研究有利于进一步了解红松分布机理、未来迁移过程等问题,对其经营和保护有重要意义。将景观指数法与点格局分析法结合,设定8个空间尺度,利用红松存在/不存在数据,通过计算各空间尺度上红松聚集程度和景观指数,分析小兴安岭地区红松种群在多尺度上的分布格局。研究结果表明,小尺度上红松聚集分布明显,随机分布区多处于其聚集分布区的边缘,均匀分布区则散布在其聚集分布区内。景观指数研究表明,通过景观指数可判断红松聚集分布格局趋势,而不能判断均匀分布、随机分布格局趋势,因为它们在多尺度下景观指数波动大,不能用景观指数来描述分布格局。研究得出如下结论:1)红松主要分布在其分布区的核心区域内,在分布区边缘和过渡带上呈随机分布,2)存在/不存在数据能够用来分析种群的多尺度空间分布格局,3)空间尺度的变化会引起树种分布格局的变化,随机分布随尺度增加,边缘化程度加强,4)单一尺度上,景观格局指数不能完全描述种群分布格局;而在多尺度上,变化趋势稳定的景观指数表明聚集分布存在,而波动剧烈的景观指数常与随机分布和均匀分布联系在一起,5)地形因子中,红松对坡度和海拔两个因子变化敏感。
Spatial patterns reflect the distribution characteristics of populations and the adaptions of populations to environmental resources and ecological processes. Dispersal limitation,inter- and intraspecific competition,and habitat heterogeneity all influence species distribution. In addition,species are affected by local temperature,precipitation,and terrain. Therefore,the relationship among species distribution patterns,landscape indices,and environmental factors are important in population studies affected by changing climates. To reveal these relationships,Korean pine,the dominant species in the Small Khingan Mountains,at the northern boundary of its distribution range,was selected to analyze its distribution patterns at multi-spatial scales. The point pattern analysis and landscape pattern metrics were calculated at eight spatial scales( 90 m × 90 m resolution) using R package( Spatstat 1.42-1) and FRAGSTATS( Version 4.2),based on presence / absence data derived from forest inventory maps and attributed data. The results from the point pattern analysis showed that Korean pine was aggregated in small scales,random distribution areas surround an aggregated pattern,and uniform distribution areas were often mosaicked in aggregated pattern areas. However,the distribution pattern of Korean pine was more aggregated with an increasing spatial scale because the aggregated distribution area increased,especially in the core area of this region. Meanwhile,random distribution areas occurred at the edge of aggregated distribution areas,and uniform distribution areas disappeared from the core area. The results from the landscape pattern analysis indicated that landscape pattern indices could be used to describe an aggregated distribution at multi-spatial scales since the landscape metrics were most stable,or changes were logarithmic; whereas landscape pattern indices dramatically fluctuated with changing spatial scales for the others. The analyses also revealed that Korean pine was sensitive to slope and elevation; most individuals were distributed on flat slopes or elevations between 200 and 800 m. Korean pine distribution was not affected by this aspect. Based on the above results,we concluded that 1) Korean pine was primarily aggregated in the core area of its distribution,and was randomly distributed at the borders or ecotones; 2) presence / absence data of specie was more likely to be implemented in analysis of population distribution type at multi-spatial scales; 3) with changing spatial scales( or extents),species distribution patterns changed,e. g.,the random and uniform distribution areas at small scales often merged into aggregated distribution areas at large scales; 4) landscape pattern indices could not completely describe species distribution patterns at a single scale,but at a multi-spatial scale,stable changes in landscape indices indicated an aggregated distribution,while unstable changes resulted in uniform or random distribution; 5) Korean pine was sensitive to slope and elevation. Elucidation of the spatial patterns of Korean pine aids in understanding its distribution mechanisms,future migration,climate change effects,and promotes strategies for its effective conservation and management.
Spatial patterns reflect the distribution characteristics of populations and the adaptions of populations to environmental resources and ecological processes. Dispersal limitation,inter- and intraspecific competition,and habitat heterogeneity all influence species distribution. In addition,species are affected by local temperature,precipitation,and terrain. Therefore,the relationship among species distribution patterns,landscape indices,and environmental factors are important in population studies affected by changing climates. To reveal these relationships,Korean pine,the dominant species in the Small Khingan Mountains,at the northern boundary of its distribution range,was selected to analyze its distribution patterns at multi-spatial scales. The point pattern analysis and landscape pattern metrics were calculated at eight spatial scales( 90 m × 90 m resolution) using R package( Spatstat 1.42-1) and FRAGSTATS( Version 4.2),based on presence / absence data derived from forest inventory maps and attributed data. The results from the point pattern analysis showed that Korean pine was aggregated in small scales,random distribution areas surround an aggregated pattern,and uniform distribution areas were often mosaicked in aggregated pattern areas. However,the distribution pattern of Korean pine was more aggregated with an increasing spatial scale because the aggregated distribution area increased,especially in the core area of this region. Meanwhile,random distribution areas occurred at the edge of aggregated distribution areas,and uniform distribution areas disappeared from the core area. The results from the landscape pattern analysis indicated that landscape pattern indices could be used to describe an aggregated distribution at multi-spatial scales since the landscape metrics were most stable,or changes were logarithmic; whereas landscape pattern indices dramatically fluctuated with changing spatial scales for the others. The analyses also revealed that Korean pine was sensitive to slope and elevation; most individuals were distributed on flat slopes or elevations between 200 and 800 m. Korean pine distribution was not affected by this aspect. Based on the above results,we concluded that 1) Korean pine was primarily aggregated in the core area of its distribution,and was randomly distributed at the borders or ecotones; 2) presence / absence data of specie was more likely to be implemented in analysis of population distribution type at multi-spatial scales; 3) with changing spatial scales( or extents),species distribution patterns changed,e. g.,the random and uniform distribution areas at small scales often merged into aggregated distribution areas at large scales; 4) landscape pattern indices could not completely describe species distribution patterns at a single scale,but at a multi-spatial scale,stable changes in landscape indices indicated an aggregated distribution,while unstable changes resulted in uniform or random distribution; 5) Korean pine was sensitive to slope and elevation. Elucidation of the spatial patterns of Korean pine aids in understanding its distribution mechanisms,future migration,climate change effects,and promotes strategies for its effective conservation and management.
引文
[1]惠刚盈,李丽,赵中华,党普兴.林木空间分布格局分析方法.生态学报,2007,27(11):4717-4728.
[2]Lin Y C,Chang L W,Yang K C,Wang H H,Sun I F.Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation.Oecologia,2011,165(1):175-184.
[3]Shen G C,Yu M J,Hu X S,Mi X C,Ren H B,Sun I F,Ma K P.Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.Ecology,2009,90(11):3033-3041.
[4]侯向阳,韩进轩.长白山红松林主要树种空间格局的模拟分析.植物生态学报,1997,21(3):242-249.
[5]Martínez I,Wiegand T,González-Taboada F,Obeso J R.Spatial associations among tree species in a temperate forest community in North-western Spain.Forest Ecology and Management,2010,260(4):456-465.
[6]Wiegand T,Gunatilleke S,Gunatilleke N.Species associations in a heterogeneous Sri Lankan dipterocarp forest.The American Naturalist,2007,170(4):E77-E95.
[7]邓莉萍,白雪娇,李露露,牛沙沙,韩美娜,秦胜金,周永斌.辽东山区次生林优势木本植物种间联结与相关分析.生态学杂志,2015,34(6):1473-1479.
[8]康华靖,陈子林,刘鹏,郝朝运,韦福民.大盘山自然保护区香果树种群结构与分布格局.生态学报,2007,27(1):389-396.
[9]张金屯.植物种群空间分布的点格局分析.植物生态学报,1998,22(4):344-349.
[10]刘志华,杨健,贺红士,常禹.黑龙江大兴安岭呼中林区火烧点格局分析及影响因素.生态学报,2011,31(6):1669-1677.
[11]王立海,孟春.小兴安岭带岭林区红松阔叶林景观多样性与稳定性研究.应用生态学报,2005,16(12):2267-2270.
[12]陈文波,肖笃宁,李秀珍.景观空间分析的特征和主要内容.生态学报,2002,22(7):1080-1087.
[13]刘宇,吕一河,傅伯杰.景观格局-土壤侵蚀研究中景观指数的意义解释及局限性.生态学报,2011,31(1):267-275.
[14]布仁仓,王宪礼,肖笃宁.黄河三角洲景观组分判定与景观破碎化分析.应用生态学报,1999,10(3):321-324.
[15]布仁仓,胡远满,常禹,李秀珍,贺红士.景观指数之间的相关分析.生态学报,2005,25(10):2764-2775.
[16]张健,郝占庆,宋波,叶吉,李步杭,姚晓琳.长白山阔叶红松林中红松与紫椴的空间分布格局及其关联性.应用生态学报,2007,18(8):1681-1687.
[17]杨华,李艳丽,沈林,亢新刚.长白山云冷杉针阔混交林主要树种空间分布及其关联性.生态学报,2014,34(16):4698-4706.
[18]徐丽娜,金光泽.小兴安岭凉水典型阔叶红松林动态监测样地:物种组成与群落结构.生物多样性,2012,20(4):470-481.
[19]侯红亚,王立海.小兴安岭阔叶红松林物种组成及主要种群的空间分布格局.应用生态学报,2013,24(11):3043-3049.
[20]Liu Y Y,Li F R,Jin G Z.Spatial patterns and associations of four species in an old-growth temperate forest.Journal of Plant Interactions,2014,9(1):745-753.
[21]中华人民共和国林业部.林业专业调查主要技术规定.北京:中国林业出版社,1990.
[22]Ripley B D.Modelling spatial patterns.Journal of the Royal Statistical Society,Series B,1977,39(2):172-212.
[23]Wiegand T,Moloney K A.Rings,circles,and null-models for point pattern analysis in ecology.Oikos,2004,104(2):209-229.
[24]王计平,杨磊,卫伟,陈利顶,黄志霖.黄土丘陵区景观格局对水土流失过程的影响——景观水平与多尺度比较.生态学报,2011,31(19):5531-5541.
[25]Webb E L,Stanfield B J,Jensen M L.Effects of topography on rainforest tree community structure and diversity in American Samoa,and implications for frugivore and nectarivore populations.Journal of Biogeography,1999,26(4):887-897.
[26]布仁仓,常禹,胡远满,李秀珍,贺红士.小兴安岭针叶树种在不同尺度上对环境因子的敏感性分析.植物生态学报,2008,32(1):80-87.
[27]王晓春,孙龙,周晓峰,王天明,国庆喜,李淑娟.黑龙江省森林景观的格局变化.应用与环境生物学报,2003,9(2):111-116.
[28]陈雄文.1896年和1986年黑龙江省几种森林景观的特征变化.植物学报,2000,42(9):979-984.
[29]白卫国,王祝雄.论我国东北林区森林可持续经营.林业资源管理,2008,(6):1-7.
[30]邓华卫,布仁仓,刘晓梅,贺伟,胡远满,黄乃伟.模拟分类经营对小兴安岭林区森林生物量的影响.生态学报,2012,32(21):6679-6687.
[31]马俊,布仁仓,邓华卫,胡远满,秦秦,韩风林.气候变化对小兴安岭主要阔叶树种地上部分固碳速率影响的模拟.应用生态学报,2014,25(9):2449-2459.
[32]张雷,刘世荣,孙鹏森,王同立.气候变化对马尾松潜在分布影响预估的多模型比较.植物生态学报,2011,35(11):1091-1105.
[33]Falk Wolfgang,Mellert Karl H.Species distribution models as a tool for forest management planning under climate change:risk evaluation of Abies alba in Bavaria.Journal of Vegetation Science,2011,22(4):621-634.
[34]He F L,Duncan R P.Density-dependent effects on tree survival in an old-growth Douglas fir forest.Journal of Ecology,2000,88(4):676-688.
[35]Harms K E,Condit R,Hubbell S P,Foster R B.Habitat associations of trees and shrubs in a 50-ha neotropical forest plot.Journal of Ecology,2001,89(6):947-959.
[36]Zhang Z H,Hu G,Zhu J D,Luo D H,Ni J.Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China.Ecological Research,2010,25(6):1151-1160.
[37]白雪娇,邓莉萍,李露露,牛沙沙,韩美娜,秦胜金,周永斌.辽东山区次生林木本植物空间分布.生态学报,2015,35(1):98-105.
[38]陈利顶,刘洋,吕一河,冯晓明,傅伯杰.景观生态学中的格局分析:现状、困境与未来.生态学报,2008,28(11):5521-5531.
[39]陈利顶,李秀珍,傅伯杰,肖笃宁,赵文武.中国景观生态学发展历程与未来研究重点.生态学报,2014,34(12):3129-3141.
[2]Lin Y C,Chang L W,Yang K C,Wang H H,Sun I F.Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation.Oecologia,2011,165(1):175-184.
[3]Shen G C,Yu M J,Hu X S,Mi X C,Ren H B,Sun I F,Ma K P.Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.Ecology,2009,90(11):3033-3041.
[4]侯向阳,韩进轩.长白山红松林主要树种空间格局的模拟分析.植物生态学报,1997,21(3):242-249.
[5]Martínez I,Wiegand T,González-Taboada F,Obeso J R.Spatial associations among tree species in a temperate forest community in North-western Spain.Forest Ecology and Management,2010,260(4):456-465.
[6]Wiegand T,Gunatilleke S,Gunatilleke N.Species associations in a heterogeneous Sri Lankan dipterocarp forest.The American Naturalist,2007,170(4):E77-E95.
[7]邓莉萍,白雪娇,李露露,牛沙沙,韩美娜,秦胜金,周永斌.辽东山区次生林优势木本植物种间联结与相关分析.生态学杂志,2015,34(6):1473-1479.
[8]康华靖,陈子林,刘鹏,郝朝运,韦福民.大盘山自然保护区香果树种群结构与分布格局.生态学报,2007,27(1):389-396.
[9]张金屯.植物种群空间分布的点格局分析.植物生态学报,1998,22(4):344-349.
[10]刘志华,杨健,贺红士,常禹.黑龙江大兴安岭呼中林区火烧点格局分析及影响因素.生态学报,2011,31(6):1669-1677.
[11]王立海,孟春.小兴安岭带岭林区红松阔叶林景观多样性与稳定性研究.应用生态学报,2005,16(12):2267-2270.
[12]陈文波,肖笃宁,李秀珍.景观空间分析的特征和主要内容.生态学报,2002,22(7):1080-1087.
[13]刘宇,吕一河,傅伯杰.景观格局-土壤侵蚀研究中景观指数的意义解释及局限性.生态学报,2011,31(1):267-275.
[14]布仁仓,王宪礼,肖笃宁.黄河三角洲景观组分判定与景观破碎化分析.应用生态学报,1999,10(3):321-324.
[15]布仁仓,胡远满,常禹,李秀珍,贺红士.景观指数之间的相关分析.生态学报,2005,25(10):2764-2775.
[16]张健,郝占庆,宋波,叶吉,李步杭,姚晓琳.长白山阔叶红松林中红松与紫椴的空间分布格局及其关联性.应用生态学报,2007,18(8):1681-1687.
[17]杨华,李艳丽,沈林,亢新刚.长白山云冷杉针阔混交林主要树种空间分布及其关联性.生态学报,2014,34(16):4698-4706.
[18]徐丽娜,金光泽.小兴安岭凉水典型阔叶红松林动态监测样地:物种组成与群落结构.生物多样性,2012,20(4):470-481.
[19]侯红亚,王立海.小兴安岭阔叶红松林物种组成及主要种群的空间分布格局.应用生态学报,2013,24(11):3043-3049.
[20]Liu Y Y,Li F R,Jin G Z.Spatial patterns and associations of four species in an old-growth temperate forest.Journal of Plant Interactions,2014,9(1):745-753.
[21]中华人民共和国林业部.林业专业调查主要技术规定.北京:中国林业出版社,1990.
[22]Ripley B D.Modelling spatial patterns.Journal of the Royal Statistical Society,Series B,1977,39(2):172-212.
[23]Wiegand T,Moloney K A.Rings,circles,and null-models for point pattern analysis in ecology.Oikos,2004,104(2):209-229.
[24]王计平,杨磊,卫伟,陈利顶,黄志霖.黄土丘陵区景观格局对水土流失过程的影响——景观水平与多尺度比较.生态学报,2011,31(19):5531-5541.
[25]Webb E L,Stanfield B J,Jensen M L.Effects of topography on rainforest tree community structure and diversity in American Samoa,and implications for frugivore and nectarivore populations.Journal of Biogeography,1999,26(4):887-897.
[26]布仁仓,常禹,胡远满,李秀珍,贺红士.小兴安岭针叶树种在不同尺度上对环境因子的敏感性分析.植物生态学报,2008,32(1):80-87.
[27]王晓春,孙龙,周晓峰,王天明,国庆喜,李淑娟.黑龙江省森林景观的格局变化.应用与环境生物学报,2003,9(2):111-116.
[28]陈雄文.1896年和1986年黑龙江省几种森林景观的特征变化.植物学报,2000,42(9):979-984.
[29]白卫国,王祝雄.论我国东北林区森林可持续经营.林业资源管理,2008,(6):1-7.
[30]邓华卫,布仁仓,刘晓梅,贺伟,胡远满,黄乃伟.模拟分类经营对小兴安岭林区森林生物量的影响.生态学报,2012,32(21):6679-6687.
[31]马俊,布仁仓,邓华卫,胡远满,秦秦,韩风林.气候变化对小兴安岭主要阔叶树种地上部分固碳速率影响的模拟.应用生态学报,2014,25(9):2449-2459.
[32]张雷,刘世荣,孙鹏森,王同立.气候变化对马尾松潜在分布影响预估的多模型比较.植物生态学报,2011,35(11):1091-1105.
[33]Falk Wolfgang,Mellert Karl H.Species distribution models as a tool for forest management planning under climate change:risk evaluation of Abies alba in Bavaria.Journal of Vegetation Science,2011,22(4):621-634.
[34]He F L,Duncan R P.Density-dependent effects on tree survival in an old-growth Douglas fir forest.Journal of Ecology,2000,88(4):676-688.
[35]Harms K E,Condit R,Hubbell S P,Foster R B.Habitat associations of trees and shrubs in a 50-ha neotropical forest plot.Journal of Ecology,2001,89(6):947-959.
[36]Zhang Z H,Hu G,Zhu J D,Luo D H,Ni J.Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China.Ecological Research,2010,25(6):1151-1160.
[37]白雪娇,邓莉萍,李露露,牛沙沙,韩美娜,秦胜金,周永斌.辽东山区次生林木本植物空间分布.生态学报,2015,35(1):98-105.
[38]陈利顶,刘洋,吕一河,冯晓明,傅伯杰.景观生态学中的格局分析:现状、困境与未来.生态学报,2008,28(11):5521-5531.
[39]陈利顶,李秀珍,傅伯杰,肖笃宁,赵文武.中国景观生态学发展历程与未来研究重点.生态学报,2014,34(12):3129-3141.