哈尔滨城区杂草群落分布格局及其对生境异质化的响应
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摘要
杂草是城市植被的重要组成部分,可维持与保育城市生物多样性。由于具有敏感性较高、可塑性与抗逆性较强的特点,能够在短时间内迅速响应城市生境异质化,杂草因此已成为研究城市化过程与城市植被互馈机制的绝好对象。本论文在国家自然科学基金面上项目(编号40971041)“城市化背景下杂草对城市生境异质化的响应及其适应对策”的资助下,以杂草为研究对象,选择哈尔滨近30年来城市化最为剧烈的外环线内区域(即哈尔滨城区),通过对城市化梯度上不同生境类型中杂草群落进行调查,阐明城市化背景下杂草多样性分布格局及其成因：并通过分析广布型杂草的表型可塑性,探讨杂草对城市异质性生境的响应机理。本研究填补了我国中温带大城市杂草群落的研究空白,研究成果为深入开展城市化与城市生态系统的互馈机制研究提供理论基础,可指导、优化城市生物多样性管理与保育,具重要的科学价值与实践意义。论文主要结论如下：
(1)快速城市化进程对杂草多样性造成了剧烈影响
调查的1763个样方中,共记录杂草38科128属175种,高比例的单属种及丰富的地理成分,一方面反映了哈尔滨城区生境的高度异质化,另一方面也体现出杂草进化更为完全。城市生境的不稳定与异质化使得杂草生存空间缩小,森林和灌草丛空隙、废弃地以及缝隙是杂草的主要生存空间。可塑性高和生活史周期短的广布性杂草一般对高频度与高强度的人为干扰具有较强的适应性。根据生境情况和适应特征,175种杂草可划分为广适型、阴湿型、中生型和耐践踏-刈割型杂草4个种组。
同20世纪50年代记录的611种相比,杂草总数减少了436种,表征了城市化降低了杂草物种丰富度。相比半世纪前记录,本次调查新调查到杂草63种,未被调查到的杂草有499种,留存物种112种。留存物种近半数为夏季一年生广布型杂草,虽然半个多世纪来城市生境发生了剧烈变化,但是由于其耐受性较强,可塑性较高,因此能够留存；而土地利用类型的剧烈变化,一方面造成生境的极不稳定,可能是大量多年生杂草消失的主要原因,另一方面杂草生境发生变化,大量湿生杂草、农田杂草以及草原性杂草的原有生境遭到破坏,又是物种消失的重要因素；同时城市的再自然化在一定程度上使得部分森林性杂草及园林性杂草有机会出现。
采用植物与生境相结合的方法,针对杂草敏感性高的特点,在以往研究的基础上,增加了生态指示值的划分,建立了杂草群落四级分类系统,将1763个杂草群落划分为2种生活型类型,5种休眠型类型,22种群落组,119种优势种群落。城市生境异质化,以及相同生境中的杂草共生与季相尺度上的生态位分化导致了杂草群落类型的多样化。低群落物种丰富度、高比例的单优势种表征了杂草群落多样性低,而高比例的夏季一年生生活型是杂草群落对高强度人为干扰下、不稳定的城市生境的响应。
(2)生境类型及直接的人为干扰影响杂草群落分布格局
根据群落在不同城市化梯度上的分布情况,结合其优势种及主要伴生种对生境的适应特征,哈尔滨城区杂草群落可被划分为3个类群：1)广泛分布于三个区域的广适型杂草群落,具有较短的生活史,较强的繁殖力,较高的形态可塑性,不论干扰强度如何,均能通过调节自身可塑性,正常的生长、繁殖；2)主要分布在Z2和Z3区域内的嫌城市杂草群落,生长在城市再自然化过程中的人工森林、人工湿地等相对阴湿的生境中,以夏季一年生为主,抗干扰能力相对较差,包括低矮的森林性杂草群落以及逸生杂草群落；3)主要分布在Z3区域内的极嫌城市杂草群落,以多年生生活型为主,抗干扰能力差,植株相对高大,多以直立生长为主。
Z1区城市化程度最高,硬质化地面比例较高,杂草的生长空间有限,加之热岛效应,可能在一定程度上导致了该区域杂草群落类型较少,大多为适应性较强、可塑性较高的广布型杂草群落,极适城市杂草群落还未出现。Z2区域在土地利用类型与Z1区域相似,城市化程度基本一致,杂草群落多样性指数亦无显著性差异,森林空隙比例增大,出现嫌城市杂草群落。Z3区域的城市化程度最低,土地利用类型与杂草生境类型多样,一定程度上使得该区域杂草群落类型多样,同时适宜的生长环境与低强度的人为管护,亦可能是该区杂草群落多样性显著高于Z1与Z2区域的原因之一。城市化梯度这一个综合环境因子对杂草群落分布的影响并不显著,生境类型及直接的人为干扰方式与强度是主要因素。
在城市土地利用的不断变化过程中,城市人为干扰较多,造成其偶然性与随机性多,笔者认为生境不稳定状态时,杂草分布可能与自然选择无关,其共存和分布由随机因素决定,符合生态漂变,所有物种进入新生境的能力理论上相同,即更容易被中性理论所解释。在生境相对稳定后,即杂草占据相应生境后,其分布由生境和植物共同决定,在营养条件充足且干扰较小的生境条件下,杂草群落类型丰富且群落物种丰富度较高；营养条件有所威胁的生境中竞争性强的CR型杂草可占据生态位形成群落；条件恶劣的生境中耐受性强的SR型杂草可适应该生态位形成群落,即生境相对稳定后杂草分布更容易被生态位理论所解释。如果生境稳定持续,杂草群落也必然从一年生演替到多年生,最终出现木本群落。然而城市中往往在木本群落出现之前生境就遭到了干扰与破坏,重新开始了上述循环。
(3)表型可塑性是杂草对异质化生境的适应对策之一
6种广布型杂草叶性状在不同生境类型中的变化格局基本一致,废弃地与农田往往具有较大的叶面积,较小的比叶面积；灌草丛与森林空隙则具有较大的比叶面积,叶干物质含量小；草坪具有较小的叶面积,湿地则具有较小的比叶面积与大的叶干物质含量。杂草叶面积在农田与废弃地中最大,而在草坪中最小,可能主要是受光照与土壤中水分因子的影响；杂草比叶面积在森林与灌草丛空隙大,在废弃地与湿地中小,主要是受土壤水分与养分决定,但较多的影响因素彼此相互作用；叶干物质含量灌草丛空隙中最低,在湿地中最高,主要与土壤水分呈负相关；叶片氮磷含量在不同生境类型中差异不大,与土壤总氮、总磷含量无明显相关性。
杂草各器官生物量在不同生境类型中变化大、可塑性强。主要受生境中光照影响,各器官生物量与总生物量均在废弃地、湿地2种生境类型中最大,并显著高于其余4种生境类型。然而由于多种多样的生境因子影响趋势不同,造成不同生境类型中杂草生物量分配差异并不显著。与个体水平上研究结果一致,杂草群落生物量在不同生境类型中变化明显,均以废弃地中最大,湿地生境中次之,并显著高于其余4种生境类型,此现象亦主要是受光照条件的影响。杂草群落地下/地上生物量比值(R/S)差异不大,更倾向于满足等速生长假说。
Urban ruderal is an important part of urban vegetation, which is essential for urban biodiversity conservation. Because of their high sensitivity, plasticity and resistance, ruderal species could respond to the urban habitat heterogeneity and rapidly adapt to changing habitats, ruderal species are ideal to facilitate studies on the impact of urbanization on urban ecosystems and subsequent ecosystems response. Supported by the National Natural Science Foundation of China "Response of ruderal to urban habitat heterogeneity and the adaptation strategies under urbanization", the present study took urban ruderal as the research object, and selected the study area in the Outer-Ring Road which is under the most dramatic urbanization during the nearly30years (i.e. the urban area of Harbin). To clarify the distribution pattern of ruderal communities and its causes in the context of urbanization, ruderal communities were investigated through the different habitat types along the urbanization gradient. To explore the adaptive strategy of ruderal species in response to high heterogeneous habitats, phenotypic plasticity of eurytopic ruderal species was analyzed. The present study filled the gap of ruderal research in middle latitude temperate zone cities in China, and also provided the theoretical basis for the study of influencing mechanism between urbanization and urban ecosystems, and the useful information for the local government to help them improve urban ecosystem conservation and management. The research results are as follows:
(1) Rapid urbanization has resulted in obvious changes in ruderal species diversity.
There were38families,128genera and175ruderal species in1763plots, with large proportion of monotypic-genus and rich geographical areal-types reflecting a substantially heterogeneous habitat condition, and the more completely of ruderal species evolution. Ruderal species in the urban area resided mainly in gaps constructed by each of forests, shrub-grasslands, abandoned fields and roadsides, indicating a progressively declined living space with increased heterogeneity and un-stability in species habitats. The widespread ruderal species with strong plasticity and short life cycle exhibited a relatively great tolerance with respect to intensive and frequent human disturbance. Based on its habitat properties and adaptive ability,175ruderal species could be classified into four ecological groups:eurytopic species, shady-and wet-tolerant species, mesophytic species, and trample-and mowing-tolerant species.
There were436of ruderal species that recorded in1955had reduced. The pronounced changes indicated that the rapid urbanization had reduced the ruderal species richness.63species were new record,112species were remained,499species were failed to investigate over the past half century. More than a half of the remaining species were summer annual widespread ruderal species, although the urban habitat chaged obviously, they could remain because of the high resistance and plasticity.
The unstable urban habitats under the obvious land use and land cover change maybe one of the reason of the disappeared of large number of perennial species. Moreover, large of hygrophytic species, farmland species and grassland species, which were suitable their relevant habitat, had a drought trend in urban habitats that consistent with the change of land use characterized. The urban re-naturalization was considered to relate with the emergence of plantations and ornamental species.
Considered the high sensitivity of ruderal ruderal, ecological indicator value was added to the classification system of ruderal community on the basis of previous studies. A four-level classification system was established. Within this system, the identified1763ruderal communities could be categorized into two types of life form at the first level, five types of dormancy form at the second level,22community groups at the third level,119dominance communities at the lowest level. Urban habitat heterogenization, the commensalism of ruderal in the same habitat and species' seasonal aspect niche differentiation resulted in the high diversity of ruderal community type growing in the urban area. The low species richness and high rates of uni-dominant species represented the rudral speciesdiversity was low; The high proportions of dominant mono-species and dominant annual species were the adaptive strategies in which ruderal community responses to unstable urban habitats under human disturbances with high intensity and frequency.
(2) Habitat type and the direct human disturbance have affected the distribution pattern of ruderal communities.
The number of ruderal community types showed a obvious trend along the urbanization gradient, followed by Z1(55) The impervious surface with poor infiltration had the higher proportion in Z1with the highest urbanized degree, maybe made a declined living space of ruderal species. It also maybe the important reason of the low of the number of community types together with the heat island effect. Most of the ruderal communities in Z1were widely distributed ruderal communities with strong adaptation and high plasticity. The communities of the most highly adapt to urban habitat had not yet appeared. The land use types in Z2were similar to Z1, and consistent with the degree of urbanization in Z1. The ruderal community diversity index also had no significant difference in Z1and Z2. Dislike urban ruderal communities were appeared because of the forest gap.The urbanization degree of Z3was relatively lower, the various land use types maybe resulted in the high diversity of ruderal community type in a certain extent. Moreover, the suitable environment for ruderal growth and the lower strength of the human management and protection maybe the important reason of significantly higher ruderal community diversity in Z1and Z2. Non-significant in the distribution pattern of ruderal communities along the urbanization gradient (i.e. a comprehensive environmental factor), the mainly cause was the habitattype and the direct human disturbance type and intensity.
There was more human disturbance in the process of urban land use changed, causing the more of chance and randomness. Therefore, the authors thought that when the habitat was unstable, the distribution of ruderal species was likely had no relation with natural selection, instead its distribution and coexistence was determined by the random factors, which means all the ruderal species had the same capacity of entering into new habitats theoretically, that is more easily explained by the neutral theory. When the habitat was relatively stable, the ruderal species distribution was determined by the habitat and itself, that is more easily explained by the niche theory. The ruderal community had high diversity of ruderal community type and richness under the high nutrition and less interference habitat conditions; The CR-strategy ruderal type was grown to be ruderal communities when the nutrition conditions had threatened; The SR-strategy ruderal type was grown to be ruderal communities in poor habitat conditions. If the ruderal habitat sustained stable, rudral communities will be succession from annual to perennial, culminating in woody communities. However, new disturbance always appeared before that in the urban habitat.
(3) Phenotypic plasticity is one of the adaptive strategies of ruderal to heterogeneous habitat.
Change patterns of the leaf traits of the six widely distributed ruderal species in different habitat types were the same. There was higher leaf area and lower specific leaf area in abandoned land and farmland; there was higher specific leaf area and lower leaf dry matter content in shrub and forest gap; and there was lower leaf area in lawn and there were lower specific leaf area and higher leaf dry matter content in wetland. Ruderal had the highest leaf area in farmland and abandoned land and lowest leaf area in lawn, mainly affected by light and soil moisture factors possibely; Ruderal had the higher specific leaf area in shrub and forest gap and lower leaf area in abandoned land and wetland, mainly affected by soil moisture and nutrient factors, however, interaction was occurred in most factors; Ruderal had the lowest leaf dry matter content in shrub gap and highest leaf dry matter content in wetland, mainly negatively correlated with soil moisture; no difference of Nmass and Pmass in different habitats type, and it was no significant correlation with soil total nitrogen and phosphorus content.
The ruderal biomass in different organs in different habitat types had obvious change and strong plasticity. The biomass in each organ and total biomass were largest both in abandoned land and wetland and significant higher than the other four habitat types, mainly affected by light. However, interaction was occurred in most factors, which had different affected trend, led to no significant variation of ruderal biomass distribution in different habitats. Consistent with the individual level, ruderal community biomass in different habitat types had obvious change, with the largest in in abandoned land, and then in wetland, and significant higher than other four habitat types, mainly affected by light conditions. R/S of ruderal community had no obvious change? which it was more inclined to met the allometric relationship.
(1)快速城市化进程对杂草多样性造成了剧烈影响
调查的1763个样方中,共记录杂草38科128属175种,高比例的单属种及丰富的地理成分,一方面反映了哈尔滨城区生境的高度异质化,另一方面也体现出杂草进化更为完全。城市生境的不稳定与异质化使得杂草生存空间缩小,森林和灌草丛空隙、废弃地以及缝隙是杂草的主要生存空间。可塑性高和生活史周期短的广布性杂草一般对高频度与高强度的人为干扰具有较强的适应性。根据生境情况和适应特征,175种杂草可划分为广适型、阴湿型、中生型和耐践踏-刈割型杂草4个种组。
同20世纪50年代记录的611种相比,杂草总数减少了436种,表征了城市化降低了杂草物种丰富度。相比半世纪前记录,本次调查新调查到杂草63种,未被调查到的杂草有499种,留存物种112种。留存物种近半数为夏季一年生广布型杂草,虽然半个多世纪来城市生境发生了剧烈变化,但是由于其耐受性较强,可塑性较高,因此能够留存；而土地利用类型的剧烈变化,一方面造成生境的极不稳定,可能是大量多年生杂草消失的主要原因,另一方面杂草生境发生变化,大量湿生杂草、农田杂草以及草原性杂草的原有生境遭到破坏,又是物种消失的重要因素；同时城市的再自然化在一定程度上使得部分森林性杂草及园林性杂草有机会出现。
采用植物与生境相结合的方法,针对杂草敏感性高的特点,在以往研究的基础上,增加了生态指示值的划分,建立了杂草群落四级分类系统,将1763个杂草群落划分为2种生活型类型,5种休眠型类型,22种群落组,119种优势种群落。城市生境异质化,以及相同生境中的杂草共生与季相尺度上的生态位分化导致了杂草群落类型的多样化。低群落物种丰富度、高比例的单优势种表征了杂草群落多样性低,而高比例的夏季一年生生活型是杂草群落对高强度人为干扰下、不稳定的城市生境的响应。
(2)生境类型及直接的人为干扰影响杂草群落分布格局
根据群落在不同城市化梯度上的分布情况,结合其优势种及主要伴生种对生境的适应特征,哈尔滨城区杂草群落可被划分为3个类群：1)广泛分布于三个区域的广适型杂草群落,具有较短的生活史,较强的繁殖力,较高的形态可塑性,不论干扰强度如何,均能通过调节自身可塑性,正常的生长、繁殖；2)主要分布在Z2和Z3区域内的嫌城市杂草群落,生长在城市再自然化过程中的人工森林、人工湿地等相对阴湿的生境中,以夏季一年生为主,抗干扰能力相对较差,包括低矮的森林性杂草群落以及逸生杂草群落；3)主要分布在Z3区域内的极嫌城市杂草群落,以多年生生活型为主,抗干扰能力差,植株相对高大,多以直立生长为主。
Z1区城市化程度最高,硬质化地面比例较高,杂草的生长空间有限,加之热岛效应,可能在一定程度上导致了该区域杂草群落类型较少,大多为适应性较强、可塑性较高的广布型杂草群落,极适城市杂草群落还未出现。Z2区域在土地利用类型与Z1区域相似,城市化程度基本一致,杂草群落多样性指数亦无显著性差异,森林空隙比例增大,出现嫌城市杂草群落。Z3区域的城市化程度最低,土地利用类型与杂草生境类型多样,一定程度上使得该区域杂草群落类型多样,同时适宜的生长环境与低强度的人为管护,亦可能是该区杂草群落多样性显著高于Z1与Z2区域的原因之一。城市化梯度这一个综合环境因子对杂草群落分布的影响并不显著,生境类型及直接的人为干扰方式与强度是主要因素。
在城市土地利用的不断变化过程中,城市人为干扰较多,造成其偶然性与随机性多,笔者认为生境不稳定状态时,杂草分布可能与自然选择无关,其共存和分布由随机因素决定,符合生态漂变,所有物种进入新生境的能力理论上相同,即更容易被中性理论所解释。在生境相对稳定后,即杂草占据相应生境后,其分布由生境和植物共同决定,在营养条件充足且干扰较小的生境条件下,杂草群落类型丰富且群落物种丰富度较高；营养条件有所威胁的生境中竞争性强的CR型杂草可占据生态位形成群落；条件恶劣的生境中耐受性强的SR型杂草可适应该生态位形成群落,即生境相对稳定后杂草分布更容易被生态位理论所解释。如果生境稳定持续,杂草群落也必然从一年生演替到多年生,最终出现木本群落。然而城市中往往在木本群落出现之前生境就遭到了干扰与破坏,重新开始了上述循环。
(3)表型可塑性是杂草对异质化生境的适应对策之一
6种广布型杂草叶性状在不同生境类型中的变化格局基本一致,废弃地与农田往往具有较大的叶面积,较小的比叶面积；灌草丛与森林空隙则具有较大的比叶面积,叶干物质含量小；草坪具有较小的叶面积,湿地则具有较小的比叶面积与大的叶干物质含量。杂草叶面积在农田与废弃地中最大,而在草坪中最小,可能主要是受光照与土壤中水分因子的影响；杂草比叶面积在森林与灌草丛空隙大,在废弃地与湿地中小,主要是受土壤水分与养分决定,但较多的影响因素彼此相互作用；叶干物质含量灌草丛空隙中最低,在湿地中最高,主要与土壤水分呈负相关；叶片氮磷含量在不同生境类型中差异不大,与土壤总氮、总磷含量无明显相关性。
杂草各器官生物量在不同生境类型中变化大、可塑性强。主要受生境中光照影响,各器官生物量与总生物量均在废弃地、湿地2种生境类型中最大,并显著高于其余4种生境类型。然而由于多种多样的生境因子影响趋势不同,造成不同生境类型中杂草生物量分配差异并不显著。与个体水平上研究结果一致,杂草群落生物量在不同生境类型中变化明显,均以废弃地中最大,湿地生境中次之,并显著高于其余4种生境类型,此现象亦主要是受光照条件的影响。杂草群落地下/地上生物量比值(R/S)差异不大,更倾向于满足等速生长假说。
Urban ruderal is an important part of urban vegetation, which is essential for urban biodiversity conservation. Because of their high sensitivity, plasticity and resistance, ruderal species could respond to the urban habitat heterogeneity and rapidly adapt to changing habitats, ruderal species are ideal to facilitate studies on the impact of urbanization on urban ecosystems and subsequent ecosystems response. Supported by the National Natural Science Foundation of China "Response of ruderal to urban habitat heterogeneity and the adaptation strategies under urbanization", the present study took urban ruderal as the research object, and selected the study area in the Outer-Ring Road which is under the most dramatic urbanization during the nearly30years (i.e. the urban area of Harbin). To clarify the distribution pattern of ruderal communities and its causes in the context of urbanization, ruderal communities were investigated through the different habitat types along the urbanization gradient. To explore the adaptive strategy of ruderal species in response to high heterogeneous habitats, phenotypic plasticity of eurytopic ruderal species was analyzed. The present study filled the gap of ruderal research in middle latitude temperate zone cities in China, and also provided the theoretical basis for the study of influencing mechanism between urbanization and urban ecosystems, and the useful information for the local government to help them improve urban ecosystem conservation and management. The research results are as follows:
(1) Rapid urbanization has resulted in obvious changes in ruderal species diversity.
There were38families,128genera and175ruderal species in1763plots, with large proportion of monotypic-genus and rich geographical areal-types reflecting a substantially heterogeneous habitat condition, and the more completely of ruderal species evolution. Ruderal species in the urban area resided mainly in gaps constructed by each of forests, shrub-grasslands, abandoned fields and roadsides, indicating a progressively declined living space with increased heterogeneity and un-stability in species habitats. The widespread ruderal species with strong plasticity and short life cycle exhibited a relatively great tolerance with respect to intensive and frequent human disturbance. Based on its habitat properties and adaptive ability,175ruderal species could be classified into four ecological groups:eurytopic species, shady-and wet-tolerant species, mesophytic species, and trample-and mowing-tolerant species.
There were436of ruderal species that recorded in1955had reduced. The pronounced changes indicated that the rapid urbanization had reduced the ruderal species richness.63species were new record,112species were remained,499species were failed to investigate over the past half century. More than a half of the remaining species were summer annual widespread ruderal species, although the urban habitat chaged obviously, they could remain because of the high resistance and plasticity.
The unstable urban habitats under the obvious land use and land cover change maybe one of the reason of the disappeared of large number of perennial species. Moreover, large of hygrophytic species, farmland species and grassland species, which were suitable their relevant habitat, had a drought trend in urban habitats that consistent with the change of land use characterized. The urban re-naturalization was considered to relate with the emergence of plantations and ornamental species.
Considered the high sensitivity of ruderal ruderal, ecological indicator value was added to the classification system of ruderal community on the basis of previous studies. A four-level classification system was established. Within this system, the identified1763ruderal communities could be categorized into two types of life form at the first level, five types of dormancy form at the second level,22community groups at the third level,119dominance communities at the lowest level. Urban habitat heterogenization, the commensalism of ruderal in the same habitat and species' seasonal aspect niche differentiation resulted in the high diversity of ruderal community type growing in the urban area. The low species richness and high rates of uni-dominant species represented the rudral speciesdiversity was low; The high proportions of dominant mono-species and dominant annual species were the adaptive strategies in which ruderal community responses to unstable urban habitats under human disturbances with high intensity and frequency.
(2) Habitat type and the direct human disturbance have affected the distribution pattern of ruderal communities.
The number of ruderal community types showed a obvious trend along the urbanization gradient, followed by Z1(55)
There was more human disturbance in the process of urban land use changed, causing the more of chance and randomness. Therefore, the authors thought that when the habitat was unstable, the distribution of ruderal species was likely had no relation with natural selection, instead its distribution and coexistence was determined by the random factors, which means all the ruderal species had the same capacity of entering into new habitats theoretically, that is more easily explained by the neutral theory. When the habitat was relatively stable, the ruderal species distribution was determined by the habitat and itself, that is more easily explained by the niche theory. The ruderal community had high diversity of ruderal community type and richness under the high nutrition and less interference habitat conditions; The CR-strategy ruderal type was grown to be ruderal communities when the nutrition conditions had threatened; The SR-strategy ruderal type was grown to be ruderal communities in poor habitat conditions. If the ruderal habitat sustained stable, rudral communities will be succession from annual to perennial, culminating in woody communities. However, new disturbance always appeared before that in the urban habitat.
(3) Phenotypic plasticity is one of the adaptive strategies of ruderal to heterogeneous habitat.
Change patterns of the leaf traits of the six widely distributed ruderal species in different habitat types were the same. There was higher leaf area and lower specific leaf area in abandoned land and farmland; there was higher specific leaf area and lower leaf dry matter content in shrub and forest gap; and there was lower leaf area in lawn and there were lower specific leaf area and higher leaf dry matter content in wetland. Ruderal had the highest leaf area in farmland and abandoned land and lowest leaf area in lawn, mainly affected by light and soil moisture factors possibely; Ruderal had the higher specific leaf area in shrub and forest gap and lower leaf area in abandoned land and wetland, mainly affected by soil moisture and nutrient factors, however, interaction was occurred in most factors; Ruderal had the lowest leaf dry matter content in shrub gap and highest leaf dry matter content in wetland, mainly negatively correlated with soil moisture; no difference of Nmass and Pmass in different habitats type, and it was no significant correlation with soil total nitrogen and phosphorus content.
The ruderal biomass in different organs in different habitat types had obvious change and strong plasticity. The biomass in each organ and total biomass were largest both in abandoned land and wetland and significant higher than the other four habitat types, mainly affected by light. However, interaction was occurred in most factors, which had different affected trend, led to no significant variation of ruderal biomass distribution in different habitats. Consistent with the individual level, ruderal community biomass in different habitat types had obvious change, with the largest in in abandoned land, and then in wetland, and significant higher than other four habitat types, mainly affected by light conditions. R/S of ruderal community had no obvious change? which it was more inclined to met the allometric relationship.
引文
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