生境破坏的模式对集合种群动态和续存的影响

构建了空间关联的集合种群模型,该模型不但包含了种群的空间结构信息,而且引入了破坏生境的全局密度和局部密度两个指标,它们描述了破坏生境的模式.模型揭示了破坏生境的空间分布格局复杂地影响了集合种群的动态和续存,破坏和未破坏生境斑块的均匀混合不利于集合种群的增长和续存,而生境类型聚集分布可以促进集合种群的快速增长和长期续存;对于两种斑块类型相对均匀混合的生境来说,均匀场假设可能会高估集合种群的续存,对于相对斑块类型高度聚集的生境,均匀场假设可能会低估集合种群的续存;物种的迁移范围也会影响集合种群的续存,迁移范围越大的物种越容易抵御生境的破坏而免遭灭绝.这意味着在生物保护中不能仅仅考虑生境的恢复和斑块质量的改善,生境结构的构建也是很重要的,加强生境斑块之间的连通性也有利于物种的长期续存.     

集合种群的理论框架与应用研究进展

集合种群的研究是当今国际生态学的重要前沿与热点。随着全球范围的生境破坏和破碎化,集合种群的研究方法已成为数学生态学、理论生态学和保护生物学的重要手段。由于其迅速的发展,集合种群的概念与理论得到迅速扩展与丰富。为了能总观集合种群进展的全局并开展进一步的工作．首先对集合种群的已有概念、理论和模型做了全面的分析和总结;其次对集合种群的发展和概念进行了探讨,以集合种群模型的中心框架：Levins的斑块占据模型为基础,展开对其它原理、效应和机制的探讨;主要包括了Levins原理．即当生境遭到破坏时,空斑块比例在集合种群灭绝前保持不变,然后还分析了Allee效应(集合种群的Allee效应主要是由于建群困难和扩散损失造成的);第三,分析了援救效应：迁入个体可以降低斑块中现有局域种群的灭绝风险。援救效应会增强集合种群的生存力,使空斑块比例下降。第四,探讨了两竞争集合种群的共存机制,即竞争,侵占妥协,其共存机制为空间生境中物种共存提供了有力的理论解释。最后,对集合种群群落中的灭绝债务进行了讨论。并给出了2种最为主要的集合种群空间模拟方法。     

集合种群的似Allee效应

从局域种群出发,建立了一个既包括局域种群动态,又包含集合种群侵占率的整合模型,并在这两个层次上进行了计算机模拟,结果表明：（1）同局域种群的Allee效应相类似,集合种群的斑块（适宜生境）侵占比例也存在一个临界值,即使有足够的适宜生境,当斑块的侵占比例低于这个临界值时,集合种群优将趋于灭绝。（2）这个临界值与局域种各的Allee效应密切相关,这将给自然保护,尤其稀有生物的保护以很大的启示。     

集合群落(metacommunity)研究动态

由于人类活动等原因,自然生境在日益丧失和破碎化。因此,集合种群结构作为空间生态学的一个重要的研究途径越来越受到人们的重视,而斑块化环境中的生物群落即集合群落也成为生态学的一个研究热点。文中综述了集合群落动态,群落物种丰富度和物种分布,生境的丧失和破坏对集合群落的影响等的最新研究成果。     

相互作用的集合种群研究动态

在集合种群水平上,两个或更多物种可以生活在同一个斑块网络中而没有相互作用.但在很多情况下,种间的相互作用会影响种群的迁移率、灭绝率和侵占率,从而调节相应物种的集合种群动态.这方面的研究主要有集合种群水平上物种之间的竞争、捕食以及在没有任何环境异质性的条件下物种在空间上聚集分布的产生和维持等.综述了近年来关于集合种群水平上的竞争,捕食者和猎物系统以及捕食与复杂空间动态的最新研究成果.     

Allee效应对具有生境恢复的集合种群的影响

Allee效应与种群的灭绝密切相关,其研究对生态保护和管理至关重要。Allee效应对物种续存是潜在的干扰因素,濒危物种更容易受其影响,可能会增加生存于生境破碎化斑块的濒危物种的死亡风险,因此研究Allee效应对种群的动态和续存的影响是必要的。从包含由生物有机体对环境的修复产生的Allee效应的集合种群模型出发,引入由其他机制形成的Allee效应,建立了常微分动力系统模型和基于网格模型的元胞自动机模型。通过理论分析和计算机模拟表明:(1)强Allee效应不利于具有生境恢复的集合种群的续存;(2)生境恢复有利于种群续存;(3)局部扩散影响了集合种群的空间结构、动态行为和稳定性,生境斑块之间的局部作用将会减缓或消除集合种群的Allee效应,有利于集合种群的续存。     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

集合种群理论在生态恢复中的应用

环境问题的改善是生态与环境学家所面临的挑战之一 ,其核心是生态系统的稳定性和平衡性遭到极大的破坏 ,最明显的表现是生境破碎和栖息地丧失。本文探讨了集合种群理论的形成与成因 ,概述了该理论发展的最新成果及应用前景。在集合种群理论应用于生态恢复实践的论述中 ,阐述了一些原则性问题 ,包括集合种群平衡观、最小可存活集合种群、最适斑块密度以及高质量生境斑块等 ,以求在人们设计物种保护和恢复对策时 ,有一定的指导价值。     

生境变化对集合种群系统生态效的影响

通过大量的数值模拟发现：生境恢复或扩展将导致集合种群的强弱序由自然数的顺序规律演变为奇数种群强－偶数种群弱,同时集合种群里的最优秀种群将迅速扩张,发展为更为强大的最优势种,而当生境遭受到破坏（毁坏）,集合种群里的最优秀种群将迅速地伦为最弱者,如果栖息地的毁坏率大于集合种群优势种对栖息地的占有率,不仅集合种群里的优势种群将不可避免地灭绝,伴随最优秀种群走向灭绝的种群依次还有第二、第三、第四强等的种群。同时将导致集合种群的强弱序由自然数的顺序规律演变为偶数种群强－奇数种群弱。     

集合种群空间混沌的模拟研究以及Allee效应、拥挤效应与捕食效应对空间模式的影响

集合种群的空间模式研究是当今生态学的核心问题之一。本研究利用常微分动力系统以及基于网格模型的元胞自动机模型对Allee效应、拥挤效应以及捕食作用集合种群的空间分布模式做了全面的模拟研究。Allee效应描述当种群水平低于某一阈值时会发生由生殖成功几率下降造成的种群负增长率,而拥挤效应是指当种群密度过高时引起的个体性为异常从而达到调节种群增长率的作用。文章组建了3个空间确定性模型：局部作用模型(CIM)、距离敏感模型(DSM)和集合种群捕食模型(MMP)。局部作用模型显示在一维生境中空斑块形成金字塔状,二维模型显示出明显的动态拟周期性以及由空间混沌所形成的异质性。距离敏感模型可导致由迁移个体中密度制约强度决定的集合种群大小复杂动态与种群密度的双峰分布。这些结果说明动态行为的复杂性,不仅可用于表征研究物种的特性,而且可以表明该物种的续存能力与灭绝风险。集合种群捕食模型是概率转换空间模型,利用该模型得出了依赖于模型参数和生境尺度的白组织种群概率空间分布模式。模拟的结果表明,系统的内在机制和这种白组织模式导致捕食者形成集团型不明显的“捕食小组”或“杀手小组”,并具有较高扩散力．但却包括侵占率低、灭绝率高的特点。而使猎物种群形成高集团性、高侵占率、低灭绝率、低扩散力的种群集团。这种特点又使捕食者种群在生境中处于中心地带,而使猎物种群形成在捕食者和生境边缘间的环状分布。这些结果还说明了尺度对于生态学的研究是至关重要的,不同的尺度将产生不同的系统模式。     

Metapopulation models for extinction threshold in spatially correlated landscapes

Simple analytical models assuming homogeneous space have been used to examine the effects of habitat loss and fragmentation on metapopulation size. The models predict an extinction threshold, a critical amount of suitable habitat below which the metapopulation goes deterministically extinct. The consequences of non-random loss of habitat for species with localized dispersal have been studied mainly numerically. In this paper, we present two analytical approaches to the study of habitat loss and its metapopulation dynamic consequences incorporating spatial correlation in both metapopulation dynamics as well as in the pattern of habitat destruction. One approach is based on a measure called metapopulation capacity, given by the dominant eigenvalue of a "landscape" matrix, which encapsulates the effects of landscape structure on population extinctions and colonizations. The other approach is based on pair approximation. These models allow us to examine analytically the effects of spatial structure in habitat loss on the equilibrium metapopulation size and the threshold condition for persistence. In contrast to the pair approximation based approaches, the metapopulation capacity based approach allows us to consider species with long as well as short dispersal range and landscapes with spatial correlation at different scales. The two methods make dissimilar assumptions, but the broad conclusions concerning the consequences of spatial correlation in the landscape structure are the same. Our results show that increasing correlation in the spatial arrangement of the remaining habitat increases patch occupancy, that this increase is more evident for species with short-range than long-range dispersal, and that to be most beneficial for metapopulation size, the range of spatial correlation in landscape structure should be at least a few times greater than the dispersal range of the species.     

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。     

Spatial Patterns of Prisoner’s Dilemma Game in Metapopulations

Because to defect is the evolutionary stable strategy in the prisoner’s dilemma game (PDG), understanding the mechanism generating and maintaining cooperation in PDG, i.e. the paradox of cooperation, has intrinsic significance for understanding social altruism behaviors. Spatial structure serves as the key to this dilemma. Here, we build the model of spatial PDG under a metapopulation framework: the sub-populations of cooperators and defectors obey the rules in spatial PDG as well as the colonization–extinction process of metapopulations. Using the mean-field approximation and the pair approximation, we obtain the differential equations for the dynamics of occupancy and spatial correlation. Cellular automaton is also built to simulate the spatiotemporal dynamics of the spatial PDG in metapopulations. Join-count statistics are used to measure the spatial correlation as well as the spatial association of the metapopulation. Simulation results show that the distribution is self-organized and that it converges to a static boundary due to the boycotting of cooperators to defectors. Metapopulations can survive even when the colonization rate is lower than the extinction rate due to the compensation of cooperation rewards for extinction debt. With a change of parameters in the model, a metapopulation can consist of pure cooperators, pure defectors, or cooperator–defector coexistence. The necessary condition of cooperation evolution is the local colonization of a metapopulation. The spatial correlation between the cooperators tends to be weaker with the increase in the temptation to defect and the habitat connectivity; yet the spatial correlation between defectors becomes stronger. The relationship between spatial structure and the colonization rate is complicated, especially for cooperators. The metapopulation may undergo a temporary period of prosperity just before the extinction, even while the colonization rate is declining. An erratum to this article can be found at     

物种灭绝对不同时间尺度栖息地毁坏响应的空间模拟

人类活动所引起的栖息地毁坏已成为当前物种多样性丧失的最主要的原因之一。空间显含模型相对于空间隐含模型来说,更加接近于现实,因此,通过元胞自动机,模拟了物种多样性对万年、千年、百年时间尺度人类活动所引起的栖息地毁坏的响应。研究结果表明：万年时间尺度上,物种是由强到弱的灭绝;而在千年时间尺度上,物种灭绝的序受集合种群结构的影响较大;在百年时间尺度上。物种由于栖息地毁坏过于剧烈和迅速,来不及作出响应。在栖息地完全毁坏时集体灭绝。因此,物种灭绝序不只是受竞争-侵占均衡机制的影响,还受不同时间尺度（不同速率）栖息地毁坏的影响。以及集合种群结构的影响。     

Are spatially correlated or uncorrelated disturbance regimes better for the survival of species?

The survival of species in dynamic landscapes (characterised by patch destruction and subsequent regeneration) depends on both the species' attributes and the disturbance pattern. Using a spatially explicit model we explored how the mean time to extinction of a metapopulation depends on the spatial correlation of patch destruction in relation to the population growth and dispersal abilities of species. Two contrasting answers are possible. On the one hand, increasing spatial correlation of patch destruction increases the spatial correlation of population growth and this is known to decrease metapopulation persistence. On the other hand, spatially correlated patch destruction and regeneration can lead to clustered habitat patches and this is known to increase metapopulation persistence. Therefore, we hypothesised that some species are better off under spatially correlated and alternatively uncorrelated disturbance regimes. However, contrary to this hypothesis, in all kinds of cases spatial correlation reduced metapopulation persistence. We found this to be due to the fact that the spatial correlation of patch destruction causes increasing temporal fluctuations in the regional carrying capacity of the metapopulation and is hence generally disadvantageous for long-term persistence. The main consequence for conservation biology is that reducing spatial correlation in disturbances is likely to be a reliable strategy in a dynamic landscape that will benefit practically all species with a low risk of adverse side effects .     

集合种群动态对栖息地毁坏时空异质性的响应

栖息地毁坏既有时间异质性,也有空间异质性,而以往的研究往往只关注其中的一种。将两种不同的异质性共同引入到元胞自动机中,模拟了集合种群动态对栖息地毁坏时空异质性的响应。发现,在随机离散的栖息地毁坏下,由于物种的迁移繁殖力受栖息地毁坏的影响很大,迁移繁殖力弱而竞争力强的物种先灭绝。在连续的栖息地毁坏下,物种的迁移繁殖力受栖息地毁坏的影响较小,物种的灭绝由竞争力和迁移繁殖力共同决定:在有绝对优势种的群落里,种间竞争显著,弱物种先灭绝,而在没有绝对优势种的群落里,种间竞争较小,则以强物种先灭绝。因此,随机毁坏不利于强物种续存,而连续毁坏则不利于具有绝对优势种群的群落里的弱物种续存。在实际开发某一栖息地时,根据集合种群结构和被保护的对象采取相应的开发模式。     

Merging spatial and temporal structure within a metapopulation model

Current research recognizes that both the spatial and temporal structure of the landscape influence species persistence. Patch models that incorporate the spatial structure of the landscape have been used to investigate static habitat destruction by comparing persistence results within nested landscapes. Other researchers have incorporated temporal structure into their models by making habitat suitability a dynamic feature of the landscape. In this article, we present a spatially realistic patch model that allows patches to be in one of three states: uninhabitable, habitable, or occupied. The model is analytically tractable and allows us to explore the interactions between the spatial and temporal structure of the landscape as perceived by the target species. Extinction thresholds are derived that depend on habitat suitability, mean lifetime of a patch, and metapopulation capacity. We find that a species is able to tolerate more ephemeral destruction, provided that the rate of the destruction does not exceed the scale of its own metapopulation dynamics, which is dictated by natural history characteristics and the spatial structure of the landscape. This model allows for an expansion of the classic definition of a patch and should prove useful when considering species inhabiting complex dynamic landscapes, for example, agricultural landscapes.     

Responses of Metapopulation Dynamics to Two Different Kinds of Habitat Destruction Caused by Human Activities

Habitat destruction can be classified into instantaneous destruction and continuous destruction by the different ways of human destroying habitat. Previous studies, however, always focused on instantaneous destruction. In this study, we develop a universal model, Multi-time scale N-species model, to study and compare the responses of metapopulation dynamics to both kinds of habitat destruction. The model explores that: (1) under instantaneous habitat destruction, species extinction is determined by the proportion of habitat destruction (D) and the structure of metapopulation (q). When D>q, species will go extinct ranked from the best competitor to the worst. When D≤ q, no species will go extinct, but the equilibrium abundances of odd-ranked competitors will decrease, and the equilibrium abundances of even-ranked competitors will increase; (2) under continuous destruction, species extinction is dependent on the speed of habitat destruction and the metapopulation structure. The higher the speed of habitat destruction and the bigger q are, the earlier species go extinct. Usually, there are two possible mechanisms of species extinction: one is that all species go extinct collectively following complete destruction, and the other is that species go extinct in ranked competitive order from best to worst, and the survivals, if they exist, will go extinct collectively following complete destruction. The oscillation amplitudes of inferior competitors are so large as to increase the probability of stochastic extinction under instantaneous destruction. Therefore, it is relatively propitious for the persistence of rare species under slow and continuous destruction, especially when continuous destruction stops.     

Habitat Deterioration, Habitat Destruction, and Metapopulation Persistence in a Heterogenous Landscape

Levins's unstructured metapopulation model predicts that the equilibrium fraction of empty habitat patches is a constant function of the fractionhof suitable patches in the landscape and that this constant equals the threshold value for metapopulation persistence. Levins's model thus suggests that the minimum amount of suitable habitat necessary for metapopulation persistence can be estimated from the fraction of empty patches at steady state. In this paper we construct several more realistic structured metapopulation models that include variation in patch quality and the rescue effect. These models predict both positive and negative correlations between the fractions of suitable patches and empty patches. The type of correlation depends in an intricate manner on the strength of the rescue effect and on the quality distribution of the patches to be destroyed. Empty patches can be considered as the resource limiting metapopulation growth. Our results demonstrate that the correlation between the fractions of suitable patches and empty patches is positive if and only if the average value of the resource decreases as the number of patches increases.