城市化背景下青岛城市森林植被多样性格局成因及其维持
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摘要
快速城市化进程对植被的物种多样性、群落类型及结构、分布格局乃至于种群更新和群落演替都会产生重要影响。自1992年以来,青岛经历了快速而激烈的城市化变迁,其特殊的地形地貌造就了青岛独特的城市发展模式,并保存了相对完好的残存自然植被。本论文在对青岛城市空间扩展历程和土地利用现状进行总结和分析的基础上,定量化地划分了城乡梯度,并据此重点探究了城乡梯度上青岛城市森林植被多样性格局及其主要影响因素,构建了青岛城市森林植被分类体系,对青岛城市森林群落的成因及其动态发展进行了分析；同时以片段化的残存丘陵森林为对象,探讨了城市自然生境岛屿化对物种多样性的影响,以期为青岛“森林城市”和园林绿化建设提供理论支撑和科学依据。主要结论如下：
(1)青岛城市空间扩展模式大致分为两个阶段：1992年以前的轴向带状扩展模式和1992年至今的多中心、飞地式扩展模式。对2009年遥感影像的土地利用类型分析表明：下垫面不透水率大于90%的区域与百年前最早城市化的区域一致；不透水率介于70%至90%的区域为老城区外延和新城区中心地带；不透水率介于40%至70%的区域为新城区外延；不透水率低于40%的区域为丘陵林地和包含农田的村镇。据此,青岛城乡梯度可被划分为高密度城区、中密度城区、低密度城区和非城市区域。
(2)以丘陵、公园和道路3种绿地为对象,分别在城乡梯度上进行样线调查,共设置336个样方,记录木本植物52科111属189种。参考样方聚类分析结果,在56%水平上划分出51个群落类型；依据群落的生活型组成、建群种亲缘性和优势种组成,建立植被型组-群系组-群系3级分类等级,并结合群落的起源性质(残存自然林和人工林),构建了青岛城市森林植被分类体系：其中残存自然林中包括1个植被型组、2个群系组、3个群系,人工林包括4个植被型组、15个群系组、50个群系。
(3)沿城乡梯度,丘陵、公园和道路绿地在物种丰富度和个体密度上呈现不同变化格局。公园绿地在城乡梯度上无显著的丰富度和个体密度变化。在丘陵和道路绿地中,乡土物种的丰富度和个体密度由高密度城区至非城市区逐渐增加；外来物种的丰富度则逐渐降低,但个体密度无显著变化。由于丘陵绿地的乡土物种增加趋势强于外来物种减少趋势,总体上其物种丰富度和个体密度呈增加趋势；而道路绿地的乡土物种增加趋势和外来物种减少趋势接近,总体上无显著变化。
(4)选取自然环境、群落结构和人为活动3大类16个因子对群落的物种丰富度和物种组成进行了分析,乡土物种的丰富度在丘陵和公园绿地中均受3类因素影响,其中人为干扰影响最大；在道路绿地中主要由群落结构决定,且不受自然环境影响。外来物种的丰富度在丘陵绿地中仅由人为活动因素决定,在其他两类绿地中则由自然环境和群落结构决定。此外,随人为管控强度增加,在丘陵-公园-道路中,乡土物种受3类因素的影响逐渐降低,而外来物种逐渐增加。3类因子对群落物种组成的总体解释率很低,其中人为干扰的影响最大。
(5)以逆-J字型、L型、间歇(多峰)型、单峰型和单柱型为五种基本更新格局,分别对51个群落的主要物种进行种群结构分析,依据各类更新格局出现频次的分布,判定每一乔木树种的更新潜力,进而对各群落的短期发展趋势进行预判,并最终将各群落归纳为自我维持型群落、干扰依赖维持型群落、自优势种替代型群落和侵入优势种替代型群落四种类型。
(6)群落的林下更新层幼苗的丰富度和密度主要取决于群落结构和人为活动两种因素；前者中上木层个体密度的影响最大,表明了种源对幼苗更新的重要贡献；后者中人为干扰强度影响较大,表明了森林管理和利用的方式、频度及强度对幼苗更新的抑制作用。此外,残存自然林中乡土种的幼苗丰富度受3类因子的影响较人工林中的大；外来物种则相反。这与残存自然林乡土种种源丰富、人工林外来种种源丰富有关。
(7)以片段化残存森林斑块为对象,采用斑块面积、距离种源(崂山)远近、斑块隔离度和人为干扰强度4个影响因子,通过CCA分析量化影响因子与物种组成的关系,赋予各影响因子权重,形成岛屿化综合指数,进而探讨其与物种丰富度的关系。结果表明,城市中片段化森林斑块的物种丰富度与岛屿化综合指数的关系随着种类性质的变化而不同。乡土物种丰富度与岛屿化综合指数有显著关系,但外来物种无此关系；鸟类传播和风力传播物种的丰富度与岛屿化综合指数关系有显著关系,而重力传播物种无此关系；除常绿物种、灌木物种和针叶物种外,其他生活型的物种丰富度与岛屿化综合指数均以对数函数拟合效果最佳。这表明,城市化带来的片段化效应对位于暖温带落叶阔叶林区的青岛森林植被所产生的负面影响,集中体现在散播能力强的落叶阔叶乔木乡土物种上。
The rapid urbanization process has a significant impact on species diversity and distribution, community types and structure, even population regeneration and community succession. Since1992, Qingdao has experienced rapid and intense urbanization, and formed a special urban space expansion mode while the remaining natural vegetation is preserved well on hills in urban area. This study summarized the urban development process and analyzed the status of land use of Qingdao, according to which the urban-rural gradients were divided, built a vegetation classification system for Qingdao urban forests, and analyzed the community assemblage rules and dynamics of Qingdao urban forests. Moreover, fragmentation effects on species diversity were studied on remaining natural forests in hills scattered in urban region. We hope this study can provide the theoretical support and scientific basis for "Qingdao Forest City" and landscaping construction. The main conclusions are as follows:
(1) Generally, Qingdao urban space expansion mode was divided into two phases: axial belt expansion mode before1992and multi-center expansion mode after1992. The land use type analysis based on2009remote sensing image showed that, the regions where impervious land percentage greater than90%are consistent with the earliest urbanized area before a century; the regions where impervious land percentage between70%and90%are the extension of old urban area and new urban centers; the regions where impervious land percentage between40%and70%are the extension of new urban area; the regions where impervious land percentage lower than40%are hill forests and farm land in villages and towns. According to these, urban-rural gradients were divided into high-density urban areas, medium-density urban areas, low-density urban areas and non-urban areas, in Qingdao.
(2) Three transects were established along urban-rural gradients for urban forests in hills (HF), parks (PF) and aside roads(RF), respectively. The surveys showed that 111genera,52families,189species of woody plants were recorded in336plots, totally. In56%level,51community types were divided based on clustering analysis. We established a four-level vegetation classification system for Qingdao urban forest. In the first level, urban forests were divided into two groups:remain natural forests and plantations. In the second level, both of two groups were classified into vegetation type groups by life form composition. In the third level, all vegetation type groups were classified into formation groups by constructive species' affinity. Finally, all formation groups were classified into formations by dominant species. In summary, there were one vegetation type group, two formation groups and three formations for the remaining natural forests and four vegetation type groups, fifteen formation groups and fifty formations for plantations.
(3) Species richness and individual density along urban-rural gradients were different among HF, PF and RF. In PF, there were no significant variations for both richness and individual density. In HF and RF, from high-density urban areas to non-urban areas, it showed an increasing tendency in richness and individual density for native species, but decreased tendency in richness for alien species with consistent individual density. Because the increasing rate of native species was greater than the decreasing rate of alien species in HF, it showed an increasing trend of species richness and individual density, totally. In contrast, there was no significant variation in RF due to the similar relative variation rates between the increasing trend of native species and decreasing trend of alien species.
(4) Multiple regression and canonical correspondence analysis (CCA) were used to determine the effects on species richness and species composition, by using sixteen influencing factors that divided into three types:natural environment (NEFs), community structure (CSFs) and human activities (HAFs). Native species richness was influenced by all three types factors in which HAFs was the most important for both of HF and PF, and was only determined by CSFs for RF. Alien species richness was determined by HAFs in HF and determined by NEFs&CSFs in PF and RF. In addition, the explanations of variation of native species richness decreased along HF-PF-RF with increased human management intensity. Inversely, the explanations of variation of alien species richness increased. All factors had little contribution on explaining the community species composition according to CCA analysis, in which human disturbance intensity was the most effective factor.
(5) Population size-structure was analyzed for main species in51communities and classified into five basic regeneration patterns:reverse-J type, L-type, sporadic type, unimodal type and unibar type. According to frequency in five basic regeneration patterns, regeneration potential of each species was determined, and then the short-term dynamics of communities were predicted, based on which communities were summarized into four groups:self-sustaining community, disturbance-dependent sustaining community, coexist species dominance replacing community and post-colonized species dominant community.
(6) For seedlings, richness and individual density mainly depends on CSFs and HAFs. Density of individual taller than1.5m was the greatest influencing factor in the former, which suggested a significant contribution of seed production on seedling recruitment. Human disturbance intensity was the greatest influencing factor in the latter, which suggested forest management and utilization had negative effects on seedling recruitment. In addition, the explanations of variation of seedling richness decreased for native species but increased for alien species, along remain natural forests-plantations with increased human management intensity, which could be explained by higher native species richness in HF and higher alien species richness in RF.
(7) To explore the effects of habitat fragment on remnant natural forests, CCA was used to analysis the relationship between species composition (present-absence data) and four factors:area, distance to species pool, human disturbance intensity and the isolation degree. The length of vector in CCA biplot was used as weights for each influencing factor, to calculate the sum, named isolated island index (Ⅲ). Then, the relationships between III and species richness were simulated by linear model, logarithmic curve, power curve and S curve. The results indicated that relationship between III and richness was significant negative for native species, but not significant for alien species richness, and relationship between III and richness was significant negative for bird dispersal species and wind dispersal species, but not significant for gravity dispersal species. Moreover, logarithmic curve was the best simulating model for any life form group, except for evergreen broadleaved species, shrub species and coniferous species which cannot be simulated by all four models. All those results indicated that the negative fragmentation impact on Qingdao urban forest which belongs to warm temperate deciduous broad-leaved forest region was mainly embodied in tree native deciduous broad-leaved species with great dispersal ability.
(1)青岛城市空间扩展模式大致分为两个阶段：1992年以前的轴向带状扩展模式和1992年至今的多中心、飞地式扩展模式。对2009年遥感影像的土地利用类型分析表明：下垫面不透水率大于90%的区域与百年前最早城市化的区域一致；不透水率介于70%至90%的区域为老城区外延和新城区中心地带；不透水率介于40%至70%的区域为新城区外延；不透水率低于40%的区域为丘陵林地和包含农田的村镇。据此,青岛城乡梯度可被划分为高密度城区、中密度城区、低密度城区和非城市区域。
(2)以丘陵、公园和道路3种绿地为对象,分别在城乡梯度上进行样线调查,共设置336个样方,记录木本植物52科111属189种。参考样方聚类分析结果,在56%水平上划分出51个群落类型；依据群落的生活型组成、建群种亲缘性和优势种组成,建立植被型组-群系组-群系3级分类等级,并结合群落的起源性质(残存自然林和人工林),构建了青岛城市森林植被分类体系：其中残存自然林中包括1个植被型组、2个群系组、3个群系,人工林包括4个植被型组、15个群系组、50个群系。
(3)沿城乡梯度,丘陵、公园和道路绿地在物种丰富度和个体密度上呈现不同变化格局。公园绿地在城乡梯度上无显著的丰富度和个体密度变化。在丘陵和道路绿地中,乡土物种的丰富度和个体密度由高密度城区至非城市区逐渐增加；外来物种的丰富度则逐渐降低,但个体密度无显著变化。由于丘陵绿地的乡土物种增加趋势强于外来物种减少趋势,总体上其物种丰富度和个体密度呈增加趋势；而道路绿地的乡土物种增加趋势和外来物种减少趋势接近,总体上无显著变化。
(4)选取自然环境、群落结构和人为活动3大类16个因子对群落的物种丰富度和物种组成进行了分析,乡土物种的丰富度在丘陵和公园绿地中均受3类因素影响,其中人为干扰影响最大；在道路绿地中主要由群落结构决定,且不受自然环境影响。外来物种的丰富度在丘陵绿地中仅由人为活动因素决定,在其他两类绿地中则由自然环境和群落结构决定。此外,随人为管控强度增加,在丘陵-公园-道路中,乡土物种受3类因素的影响逐渐降低,而外来物种逐渐增加。3类因子对群落物种组成的总体解释率很低,其中人为干扰的影响最大。
(5)以逆-J字型、L型、间歇(多峰)型、单峰型和单柱型为五种基本更新格局,分别对51个群落的主要物种进行种群结构分析,依据各类更新格局出现频次的分布,判定每一乔木树种的更新潜力,进而对各群落的短期发展趋势进行预判,并最终将各群落归纳为自我维持型群落、干扰依赖维持型群落、自优势种替代型群落和侵入优势种替代型群落四种类型。
(6)群落的林下更新层幼苗的丰富度和密度主要取决于群落结构和人为活动两种因素；前者中上木层个体密度的影响最大,表明了种源对幼苗更新的重要贡献；后者中人为干扰强度影响较大,表明了森林管理和利用的方式、频度及强度对幼苗更新的抑制作用。此外,残存自然林中乡土种的幼苗丰富度受3类因子的影响较人工林中的大；外来物种则相反。这与残存自然林乡土种种源丰富、人工林外来种种源丰富有关。
(7)以片段化残存森林斑块为对象,采用斑块面积、距离种源(崂山)远近、斑块隔离度和人为干扰强度4个影响因子,通过CCA分析量化影响因子与物种组成的关系,赋予各影响因子权重,形成岛屿化综合指数,进而探讨其与物种丰富度的关系。结果表明,城市中片段化森林斑块的物种丰富度与岛屿化综合指数的关系随着种类性质的变化而不同。乡土物种丰富度与岛屿化综合指数有显著关系,但外来物种无此关系；鸟类传播和风力传播物种的丰富度与岛屿化综合指数关系有显著关系,而重力传播物种无此关系；除常绿物种、灌木物种和针叶物种外,其他生活型的物种丰富度与岛屿化综合指数均以对数函数拟合效果最佳。这表明,城市化带来的片段化效应对位于暖温带落叶阔叶林区的青岛森林植被所产生的负面影响,集中体现在散播能力强的落叶阔叶乔木乡土物种上。
The rapid urbanization process has a significant impact on species diversity and distribution, community types and structure, even population regeneration and community succession. Since1992, Qingdao has experienced rapid and intense urbanization, and formed a special urban space expansion mode while the remaining natural vegetation is preserved well on hills in urban area. This study summarized the urban development process and analyzed the status of land use of Qingdao, according to which the urban-rural gradients were divided, built a vegetation classification system for Qingdao urban forests, and analyzed the community assemblage rules and dynamics of Qingdao urban forests. Moreover, fragmentation effects on species diversity were studied on remaining natural forests in hills scattered in urban region. We hope this study can provide the theoretical support and scientific basis for "Qingdao Forest City" and landscaping construction. The main conclusions are as follows:
(1) Generally, Qingdao urban space expansion mode was divided into two phases: axial belt expansion mode before1992and multi-center expansion mode after1992. The land use type analysis based on2009remote sensing image showed that, the regions where impervious land percentage greater than90%are consistent with the earliest urbanized area before a century; the regions where impervious land percentage between70%and90%are the extension of old urban area and new urban centers; the regions where impervious land percentage between40%and70%are the extension of new urban area; the regions where impervious land percentage lower than40%are hill forests and farm land in villages and towns. According to these, urban-rural gradients were divided into high-density urban areas, medium-density urban areas, low-density urban areas and non-urban areas, in Qingdao.
(2) Three transects were established along urban-rural gradients for urban forests in hills (HF), parks (PF) and aside roads(RF), respectively. The surveys showed that 111genera,52families,189species of woody plants were recorded in336plots, totally. In56%level,51community types were divided based on clustering analysis. We established a four-level vegetation classification system for Qingdao urban forest. In the first level, urban forests were divided into two groups:remain natural forests and plantations. In the second level, both of two groups were classified into vegetation type groups by life form composition. In the third level, all vegetation type groups were classified into formation groups by constructive species' affinity. Finally, all formation groups were classified into formations by dominant species. In summary, there were one vegetation type group, two formation groups and three formations for the remaining natural forests and four vegetation type groups, fifteen formation groups and fifty formations for plantations.
(3) Species richness and individual density along urban-rural gradients were different among HF, PF and RF. In PF, there were no significant variations for both richness and individual density. In HF and RF, from high-density urban areas to non-urban areas, it showed an increasing tendency in richness and individual density for native species, but decreased tendency in richness for alien species with consistent individual density. Because the increasing rate of native species was greater than the decreasing rate of alien species in HF, it showed an increasing trend of species richness and individual density, totally. In contrast, there was no significant variation in RF due to the similar relative variation rates between the increasing trend of native species and decreasing trend of alien species.
(4) Multiple regression and canonical correspondence analysis (CCA) were used to determine the effects on species richness and species composition, by using sixteen influencing factors that divided into three types:natural environment (NEFs), community structure (CSFs) and human activities (HAFs). Native species richness was influenced by all three types factors in which HAFs was the most important for both of HF and PF, and was only determined by CSFs for RF. Alien species richness was determined by HAFs in HF and determined by NEFs&CSFs in PF and RF. In addition, the explanations of variation of native species richness decreased along HF-PF-RF with increased human management intensity. Inversely, the explanations of variation of alien species richness increased. All factors had little contribution on explaining the community species composition according to CCA analysis, in which human disturbance intensity was the most effective factor.
(5) Population size-structure was analyzed for main species in51communities and classified into five basic regeneration patterns:reverse-J type, L-type, sporadic type, unimodal type and unibar type. According to frequency in five basic regeneration patterns, regeneration potential of each species was determined, and then the short-term dynamics of communities were predicted, based on which communities were summarized into four groups:self-sustaining community, disturbance-dependent sustaining community, coexist species dominance replacing community and post-colonized species dominant community.
(6) For seedlings, richness and individual density mainly depends on CSFs and HAFs. Density of individual taller than1.5m was the greatest influencing factor in the former, which suggested a significant contribution of seed production on seedling recruitment. Human disturbance intensity was the greatest influencing factor in the latter, which suggested forest management and utilization had negative effects on seedling recruitment. In addition, the explanations of variation of seedling richness decreased for native species but increased for alien species, along remain natural forests-plantations with increased human management intensity, which could be explained by higher native species richness in HF and higher alien species richness in RF.
(7) To explore the effects of habitat fragment on remnant natural forests, CCA was used to analysis the relationship between species composition (present-absence data) and four factors:area, distance to species pool, human disturbance intensity and the isolation degree. The length of vector in CCA biplot was used as weights for each influencing factor, to calculate the sum, named isolated island index (Ⅲ). Then, the relationships between III and species richness were simulated by linear model, logarithmic curve, power curve and S curve. The results indicated that relationship between III and richness was significant negative for native species, but not significant for alien species richness, and relationship between III and richness was significant negative for bird dispersal species and wind dispersal species, but not significant for gravity dispersal species. Moreover, logarithmic curve was the best simulating model for any life form group, except for evergreen broadleaved species, shrub species and coniferous species which cannot be simulated by all four models. All those results indicated that the negative fragmentation impact on Qingdao urban forest which belongs to warm temperate deciduous broad-leaved forest region was mainly embodied in tree native deciduous broad-leaved species with great dispersal ability.
引文
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