生境破碎化对生物多样性的影响

生境破碎化对生物多样性和生态系统功能的影响是当前国内外生态学家研究的热点问题之一。生境破碎化导致原生境的总面积减小,产生隔离的异质种群,从而影响个体行为特性、种群间基因交换、物种间相互作用及生态过程。生境破碎化的过程引起栖息地内部食物、繁殖场所、局部小气候、边缘效应等生物和非生物条件的变化,从而影响植物种群的大小和灭绝速率、扩散和迁入、遗传和变异以及存活力等,影响动物种群的异质种群动态、适宜生境比例、灭绝阈值、种间关系等。随着景观生态学与农业科学的融合,探索利用景观布局控制害虫发生将是人类利用生境破碎化为人类服务的一条新途径。     

生境破碎化对生物多样性的影响研究综述

生境破碎化与生物多样性的关系是理论生态学的研究热点。本文在综述国内外相关研究内容的基础上,阐述了生境破碎化的概念内涵与度量,介绍了生境破碎化研究的主要理论基础、研究内容与研究进展;总结了目前生境破碎化研究存在度量破碎景观格局的方法尚未统一、研究方法有待精确以及生境破碎化与生物多样性的阈值效应尚未发现等问题。今后,半干旱地区生境破碎化对生物多样性影响的研究需要加强,空间分析理论和方法将会在生境破碎化对生物多样性影响的研究中得到更多应用。     

扩散对破碎化景观上宿主-寄生种群动态的影响

景观破碎化和扩散是空间种群模型的重要因素,对生物入侵存在着深远的影响。本章将基于偶对近似模型,探讨由局部和全局宿主-寄生相互作用共同决定的扩散模式对破坏性景观上疾病入侵与传播的影响。其中,生境破坏由生境丧失量与生境破碎化程度来描述。模拟结果显示,宿主和病毒的全局扩散对疾病的入侵与种群密度产生不对称效应：病毒的全局扩散对系统产生的影响较宿主的全局扩散更为显著。不同扩散模式下,生境丧失越高或破碎化程度越低,均将越有害于寄生病毒的入侵;同时,生境的破坏程度也显著地影响了入侵阈值对扩散模式的响应机制。本文研究结果暗示,景观破碎化的空间分布格局以及病毒扩散的限制均可作为物种保护与管理中有效的疾病控制策略。该研究结果在一定意义上丰富和发展了寄生感染理论,为物种保护提供了生态学理论依据。     

近40年东北地区陆栖脊椎动物种群数量及其生境变化评估

生物多样性是生态平衡维持和生态过程与功能实现的基础,东北地区是我国乃至全球生物多样性最为丰富的地区之一。为研究和探讨东北地区陆栖脊椎动物种群数量与生境变化之间的关系,利用物种调查数据和生境遥感观测数据,以地球生命力指数、生态系统面积和破碎度等指示性指标,综合评估了近40年东北地区陆栖脊椎动物种群数量及其生境变化。结果表明:1970—2010年,东北地区陆栖脊椎动物种群数量下降了近70.1%,森林脊椎动物种群数量减少了近80.9%,草原和荒漠生态系统脊椎动物种群数量增加了近180.9%。1980—2010年,湿地物种种群数量减少了近75.7%。1980—2015年期间,农业和城镇建设用地增幅分别达到25.2%和32.3%,不断挤占和蚕食着自然生态空间,致使自然生境面积不断减少,减幅约为8.0%。自然生境景观破碎化程度总体呈现加重趋势,尤其是森林生境,破碎化指数增加约42.7%。但是,2005年之后,自然生境景观破碎化程度加重趋势趋缓,与2005年之后脊椎动物种群数量减少幅度减缓趋势一致。森林砍伐、人口增长、城镇化、交通建设等造成的自然生态系统破碎度增加和栖息地质量下降对大型兽类影响比较显著。     

异质种群动态模型:破碎化景观动态模拟的新途径

景观破碎化导致物种以异质种群方式存活,使得基于异质种群动态模拟破碎化景观动态成为可能。异质种群动态模型的发展为景观动态模拟奠定了良好基础。根据空间处理方式的不同,异质种群模型可分为三大类,可不同程度地用于描述破碎化景观动态。(1)空间不确定异质种群模型,假定所有局域种群间均等互联,模型中不包含空间信息,仅能用于景观斑块动态描述;(2)空间确定异质种群模型,假设局域种群在二维空间上以规则格子形式排列,是一种准现实的空间处理方式,可用于景观动态的简单描述;(3)空间现实异质种群模型,包含了破碎化景观中局域种群的几何特征,可直接用于真实景观动态的模拟研究。空间现实的和基于个体的异质种群模型不但是未来异质种群模型发展的主流,也将成为未来破碎化景观动态研究的重要工具。为了更加准确完整地描述破碎化景观动态,不但应该综合运用已有的各种异质种群模型方法,更要引进新模型来刎画多物种、多变量、高维度、复杂连接的破碎化景观格局与过程。     

中国野生动物生境适宜性评价和生境破碎化研究

生境是生物出现的环境空间, 开展野生动物的生境适宜性评价和生境破碎化研究, 有助于濒危动物的保育。随着生态学科的发展, 多元统计分析、景观生态学和3S 技术被用于生境适宜性评价中, 使其研究结果广泛应用于生境质量评估、生境承载力分析、物种潜在分布预测和物种濒危机制评价等方面。然而研究对象基础资料的缺乏和研究时间较短常局限生境适宜性评价研究继续深入。生境破碎化研究常集中在破碎化现状及其对生物的影响。时空尺度的扩展和研究方向的分化应是今后生境破碎化研究的发展趋势。     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

京杭运河邵伯高邮段栖息地破碎化对狗獾种群数量分布的影响

栖息地破碎是生物多样性下降的主要原因之一。栖息地破碎引起的面积效应、隔离效应和边缘效应能影响动物种群的绝灭阈值、分布、多度、种间关系以及生态系统过程, 最终影响动物种群的数量分布。2006年10-11月和2007年10-11月, 利用全球定位系统(GPS)、地理信息系统(GIS)和样方法定量分析京杭运河邵伯至高邮段狗獾栖息地破碎化程度, 确定不同斑块的面积、斑块距离、斑块隔离度以及栖息地质量对斑块中狗獾数量分布的影响。结果表明, 各个斑块内狗獾的洞口数、粪堆数与该斑块的面积显著的正相关(r=0.961, P=0.039; r=0.999, P=0.023), 但与斑块距离、斑块隔离度无显著的相关性(P>0.05)。栖息地的质量也会影响狗獾的数量分布, 多元线性逐步回归分析表明, 人类干扰和与栖息地的郁闭性显著的影响狗獾的数量分布。以上结果说明, 京杭运河邵伯高邮段栖息地的破碎化程度对狗獾的数量分布还没有造成显著的直接影响, 但会间接降低栖息地的质量, 进而影响狗獾的生存。     

农业景观生物多样性与害虫生态控制

现代农业的一个重要特征就是人类对农田生态系统的干扰强度及频率不断增加,严重影响农业景观的结构及其生物多样性.农业景观结构的变化及其生物多样性的丧失,必然引起生态系统服务功能的弱化,不利于实施以保护自然天敌为主的害虫生态控制.农业的集约化经营导致自然生境破碎化,减少了农业景观的复杂性,使得作物和非作物变成一种相对离散化的生境类型和镶嵌的景观格局;破碎化的生境不仅会减少某些物种的丰度,还会影响物种之间的相互关系及生物群落的多样性和稳定性.非作物生境类型如林地、灌木篱墙、田块边缘区、休耕地和草地等,是一种比较稳定的异质化环境.非作物生境较少受到干扰,可以为寄生性和捕食性节肢动物提供适宜的越冬或避难场所以及替代猎物、花粉和花蜜等资源,因此,非作物生境有利于自然天敌的栖息和繁衍,也有利于它们迁入邻近的作物生境中对害虫起到调节和控制作用.景观的格局-过程-尺度影响农田生物群落物种丰富度、多度、多样性以及害虫与天敌之间的相互作用.从区域农业景观系统的角度出发,运用景观生态学的理论和方法来研究作物、害虫、天敌等组分在不同斑块之间的转移过程和变化规律,揭示害虫在较大尺度和具有异质性的空间范围内的灾变机理,可为利用农业景观生物多样性来保护农田自然天敌,实施害虫的区域性生态控制提供新的研究思路和手段.     

棉花生境面积及其破碎化对烟粉虱种群的影响

【目的】阐明棉花生境面积变化及其破碎化对烟粉虱Bemisiatabaci种群的作用规律,为合理利用作物布局进行害虫生态调控提供理论支撑。【方法】采用国际流行的微景观试验模型系统(Experimental model landscape system,EMLS)进行试验设计,田间条件下连续两年研究了棉花生境面积变化(20%、40%、60%、80%和100%;其他为玉米生境面积)及2种极端破碎化(完全连通C clumped:H=1.0;完全破碎F fragmented:H=0.0)下烟粉虱种群数量变化,采用广义线性模型(GLM)分析各因素对烟粉虱种群数量的影响。【结果】棉花生境面积及其破碎化单独作用时均对烟粉虱种群数量无显著影响,而取样时间则有显著影响。烟粉虱种群数量也没有受到取样时间与棉花生境面积、取样时间与破碎化以及三者交互作用的显著影响。但是,棉花生境面积与破碎化的互作效应则存在年度变化,2014年无显著作用,2015年显著影响烟粉虱种群。当棉花生境面积较小(20%)或较大(80%)时,破碎化程度高,烟粉虱种群数量少;棉花生境面积中等(40%和60%)时,破碎化程度低,烟粉虱种群数量少。【结论】烟粉虱种群对棉花生境面积变化有较强的适应性,而生境破碎化只能在一定程度上产生影响。     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Population Dynamics of the Andean Lizard Anolis heterodermus: Fast‐slow Demographic Strategies in Fragmented Scrubland Landscapes

Habitat fragmentation and loss affect population stability and demographic processes, increasing the extinction risk of species. We studied Anolis heterodermus populations inhabiting large and small Andean scrubland patches in three fragmented landscapes in the Sabana de Bogotá (Colombia) to determine the effect of habitat fragmentation and loss on population dynamics. We used the capture‐mark‐recapture method and multistate models to estimate vital rates for each population. We estimated growth population rate and the most important processes that affect λ by elasticity analysis of vital rates. We tested the effects of habitat fragmentation and loss on vital rates of lizard populations. All six isolated populations showed a positive or an equilibrium growth rate (λ = 1), and the most important demographic process affecting λ was the growth to first reproduction. Populations from landscapes with less scrubland natural cover showed higher stasis of young adults. Populations in highly fragmented landscapes showed highest juvenile survival and growth population rates. Independent of the landscape's habitat configuration and connectivity, populations from larger scrubland patches showed low adult survivorship, but high transition rates. Populations varied from a slow strategy with low growth and delayed maturation in smaller patches to a fast strategy with high growth and early maturation in large patches. This variation was congruent with the fast‐slow continuum hypothesis and has serious implications for Andean lizard conservation and management strategies. We suggest that more stable lizard populations will be maintained if different management strategies are adopted according to patch area and habitat structure.     

Habitat fragmentation and extinction thresholds on fractal landscapes

Habitat fragmentation is a potentially critical factor in determining population persistence. In this paper, we explore the effect of fragmentation when the fragmentation follows a fractal pattern. The habitat is divided into patches, each of which is suitable or unsuitable. Suitable patches are either occupied or unoccupied, and change state depending on rates of colonization and local extinction. We compare the behaviour of two models: a spatially implicit patch-occupancy (PO) model and a spatially explicit cellular automaton (CA) model. The PO model has two fixed points: extinction, and a stable equilibrium with a fixed proportion of occupied patches. Global extinction results when habitat destruction reduces the proportion of suitable patches below a critical threshold. The PO model successfully recreates the extinction patterns found in other models. We translated the PO model into a stochastic cellular automaton. Fractal arrangements of suitable and unsuitable patches were used to simulate habitat fragmentation. We found that: (i) a population on a fractal landscape can tolerate more habitat destruction than predicted by the patch-occupancy model, and (ii) the extinction threshold decreases as the fractal dimension of the landscape decreases. These effects cannot be seen in spatially implicit models. Landscape struc-ture plays a vital role in mediating the effects of habitat fragmentation on persistence.     

Metapopulation dynamics: Does it help to have more of the same?

With the interest in conservation biology shifting towards processes from patterns, and to populations from communities, the theory of metapopulation dynamics is replacing the equilibrium theory of island biogeography as the population ecology paradigm in conservation biology. The simplest models of metapopulation dynamics make predictions about the effects of habitat fragmentation - size and isolation of habitat patches - on metapopulation persistence. The simple models may be enriched by considerations of the effects of demographic and environmental stochasticity on the size and extinction probability of local populations. Environmental stochasticity affects populations at two levels: it makes local extinctions more probable, and it also decreases metapopulation persistence time by increasing the correlation of extinction events across populations. Some controversy has arisen over the significance of correlated extinctions, and how they may affect the optimal subdivision of metapopulations to maximize their persistence time.     

Spatially structured metapopulation models: global and local assessment of metapopulation capacity

We model metapopulation dynamics in finite networks of discrete habitat patches with given areas and spatial locations. We define and analyze two simple and ecologically intuitive measures of the capacity of the habitat patch network to support a viable metapopulation. Metapopulation persistence capacity lambda(M) defines the threshold condition for long-term metapopulation persistence as lambda(M)>delta, where delta is defined by the extinction and colonization rate parameters of the focal species. Metapopulation invasion capacity lambda(I) sets the condition for successful invasion of an empty network from one small local population as lambda(I)>delta. The metapopulation capacities lambda(M) and lambda(I) are defined as the leading eigenvalue or a comparable quantity of an appropriate "landscape" matrix. Based on these definitions, we present a classification of a very general class of deterministic, continuous-time and discrete-time metapopulation models. Two specific models are analyzed in greater detail: a spatially realistic version of the continuous-time Levins model and the discrete-time incidence function model with propagule size-dependent colonization rate and a rescue effect. In both models we assume that the extinction rate increases with decreasing patch area and that the colonization rate increases with patch connectivity. In the spatially realistic Levins model, the two types of metapopulation capacities coincide, whereas the incidence function model possesses a strong Allee effect characterized by lambda(I)=0. For these two models, we show that the metapopulation capacities can be considered as simple sums of contributions from individual habitat patches, given by the elements of the leading eigenvector or comparable quantities. We may therefore assess the significance of particular habitat patches, including new patches that might be added to the network, for the metapopulation capacities of the network as a whole. We derive useful approximations for both the threshold conditions and the equilibrium states in the two models. The metapopulation capacities and the measures of the dynamic significance of particular patches can be calculated for real patch networks for applications in metapopulation ecology, landscape ecology, and conservation biology.     

景观生态学:两栖动物生态学研究的新途径

全球两栖动物正以远超过自然灭绝的高速率灭绝,这与生境丧失和景观破碎化有着直接关系。生境丧失导致两栖动物的生存空间减少,使局部种群消失,而景观破碎化则导致两栖动物种群之间的隔离度增加,不利于动物的繁殖和扩散。但两者往往是同时出现,相互作用。复合种群、景观连接度、景观遗传学及景观模型模拟等理论和方法的发展,为在生境丧失与破碎化景观下两栖动物的种群结构、组成和动态变化研究提供了理论基础和技术方法。同时景观生态学中特别重视研究的尺度,生境破碎化是发生在景观尺度下的生境变化过程,因此对生境破碎化的影响应该从现有的主要集中在斑块尺度和斑块-景观尺度转变到景观尺度上来。     

Anthropogenic landscape change promotes asymmetric dispersal and limits regional patch occupancy in a spatially structured bird population

1. Local extinctions in habitat patches and asymmetric dispersal between patches are key processes structuring animal populations in heterogeneous environments. Effective landscape conservation requires an understanding of how habitat loss and fragmentation influence demographic processes within populations and movement between populations. 2. We used patch occupancy surveys and molecular data for a rainforest bird, the logrunner (Orthonyx temminckii), to determine (i) the effects of landscape change and patch structure on local extinction; (ii) the asymmetry of emigration and immigration rates; (iii) the relative influence of local and between-population landscapes on asymmetric emigration and immigration; and (iv) the relative contributions of habitat loss and habitat fragmentation to asymmetric emigration and immigration. 3. Whether or not a patch was occupied by logrunners was primarily determined by the isolation of that patch. After controlling for patch isolation, patch occupancy declined in landscapes experiencing high levels of rainforest loss over the last 100 years. Habitat loss and fragmentation over the last century was more important than the current pattern of patch isolation alone, which suggested that immigration from neighbouring patches was unable to prevent local extinction in highly modified landscapes. 4. We discovered that dispersal between logrunner populations is highly asymmetric. Emigration rates were 39% lower when local landscapes were fragmented, but emigration was not limited by the structure of the between-population landscapes. In contrast, immigration was 37% greater when local landscapes were fragmented and was lower when the between-population landscapes were fragmented. Rainforest fragmentation influenced asymmetric dispersal to a greater extent than did rainforest loss, and a 60% reduction in mean patch area was capable of switching a population from being a net exporter to a net importer of dispersing logrunners. 5. The synergistic effects of landscape change on species occurrence and asymmetric dispersal have important implications for conservation. Conservation measures that maintain large patch sizes in the landscape may promote asymmetric dispersal from intact to fragmented landscapes and allow rainforest bird populations to persist in fragmented and degraded landscapes. These sink populations could form the kernel of source populations given sufficient habitat restoration. However, the success of this rescue effect will depend on the quality of the between-population landscapes.     

生境丧失对具有似Allee效应集合种群的影响及对策——以江苏盐城为例

似Allee效应对物种续存是潜在的扰动因素,稀有物种更易受其影响,可能增加生存于破碎化栖息地中的珍稀物种的死亡风险;但似Allee效应对多物种集合种群续存的影响及其在珍稀物种保护中的应用未能引起足够重视。将似Allee效应引入集合种群动力模式,建立了生境丧失下具有似Allee效应的n-珍稀物种的集合种群模式,并以江苏盐城滩涂湿地中的29种珍稀物种为研究实例。研究结果表明:(1)似Allee效应导致n-物种集合种群多度作长期变周期振荡,原本竞争共存物种可能无法继续共存,甚至灭绝。(2)似Allee效应增强对次强种及劣势种的生存极为不利,导致次强物种由强至弱灭绝,劣势物种由弱至强依次灭绝。(3)盐城天然湿地丧失29%后,11种劣势物种的集合种群由弱到强将最终依次灭绝,灭绝迟豫时间为304～890a,这些物种即Hanski所指的"活死者"。(4)适度增加栖息地面积是保护珍稀物种多样性的有效方法之一,在盐城现存3200km2的天然湿地基础上适度增加1801～2064km2左右栖息地面积,可以有效保护29种濒危物种的多样性,同时应注意结合针对具体物种的保护措施来提高濒危物种多度。研究结果对物种多样性保护及自然保护区建设具有重要的理论指导意义。     

集合种群动态对生境毁坏空间异质性的响应

首次将分形几何(Fractal geometry)与元胞自动机(Cellular automata)相结合,研究了破碎化生境中集合种群的空间分布格局动态,以及集合种群动态对生境毁坏空间异质性的响应。研究发现:(1)各个物种种群在生境中的分布具有很好的分形特征,物种的计盒维数(Box dimension)不仅可以很好地反映种群的空间分布结构,也能很好地反映种群动态。(2)如果将空间因素考虑进来的话,生境毁坏的灭绝债务(Time debt)将大于空间隐含模式所模拟的结果。(3)物种灭绝同时存在强物种灭绝和弱物种灭绝。并且只有在生境随机毁坏下,才与空间隐含的模拟结果比较接近,即强物种中将是最强物种率先灭绝。而在边缘毁坏这种比较集中成块的开发方式下,将是较强的物种灭绝。(4)边缘毁坏相对随机毁坏有利于物种,尤其是弱物种的长期续存。