集合种群与生物多样性保护

集合种群的概念受到空前的重视,其精髓是强调物种受局域和区域两个空间尺度上生态学过程的共同作用。主要介绍了集合种群概念的由来、集合种群动态理论以及集合种群理论在生物多样性保护及生物防治中一些可能的应用。     

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。     

物种保护的理论基础——从岛屿生物地理学理论到集合种群理论

全球面临着生境破碎化的危机,物种保护已成为人类面临的重大课题,并不是所有的人对岛屿生物地理学理论的产生及其关注的海洋岛屿都很熟悉,但是越来越多生物赖以生存的自然栖息地的丧失和破碎化都是有目共睹的,岛屿生物地理学和集合种群理论是目前物种保护的两个基本理论,物种迁入率和绝灭率的动态变化决策岛屿上的物种丰富度是岛屿生物地理学理论的核心内容,而集合种群理论关注的是局部种群之间个体迁移的动态以及物种的续存条件,在概述两个理论形成、发展及其核心内容的基础上,着重比较它们的异同点以及在生态学理论和实践中的应用,并论述物种保护理论范式从岛屿生物地理学向集合种群理论转变的基本背景和原因。     

局域种群的Allee效应和集合种群的同步性

从包含Allee效应的局域种群出发,建立了耦合映像格子模型,即集合种群模型.通过分析和计算机模拟表明:(1)当局域种群受到Allee效应强度较大时,集合种群同步灭绝;(2)而当Allee效应强度相对较弱时,通过稳定局域种群动态(减少混沌)使得集合种群发生同步波动,而这种同步波动能够增加集合种群的灭绝风险;(3)斑块间的连接程度对集合种群同步波动的发生有很大的影响,适当的破碎化有利于集合种群的续存.全局迁移和Allee效应结合起来增加了集合种群同步波动的可能,从而增加集合种群的灭绝风险.这些结果对理解同步性的机理、利用同步机理来制定物种保护策略和害虫防治都有重要的意义.     

岛屿生物地理学与集合种群理论的本质与渊源

岛屿生物地理学和集合种群理论是目前生物多样性保育所依赖的主要生态学理论。人们通常强调这两种理论的区别,对它们之间的关联却很少注意到。事实上,这两种理论是同根同源的。以经典集合种群理论的创始者R．Levim对他与岛屿生物地理学的创始者R．H．MaeArthur的合作过程以及岛屿生物地理学对他提出集合种群理论的影响的回顾为基础,分析比较了岛屿生物地理学、经典集合种群理论、以Hanski为代表的现代集合种群理论的基本假设、研究范式和核心思想的异同,简要介绍了多物种集合种群与集合群落研究的差异,最后分析了岛屿生物地理学和集合种群理论在生物多样性保育实践中的应用和存在问题。     

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。     

通过网蛱蝶的例证研究试论集合种群的理论和方法

空间生态学 (Spatial Ecology) 近来引起了生态学界的广泛注目。集合种群 (又译作异质种群)生态学的理论和方法是它的重要组成内容。有关的文章在近二三年急剧增加。本文试图通过Hanski等人对网蛱蝶的研究,了解其所采用的研究途径,得到了什么样的数据,如何进行分析,最后得到了什么结果,并结合我们对网蛱蝶进行研究的一些体会,试图分析集合种群的理论和方法的实际内涵、特色及应用前景。     

集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。     

鸟类栖息地片段化研究的理论基础

栖息地片段化是导致许多森林鸟类种群下降的主要原因之一,而对栖息地片段化的形成及其影响的研究已是成为鸟类生态学的研究热点之一。介绍了鸟类栖息地片段化研究的理论基础,即岛屿生物地理学理论、景观生态学理论以及集合种群理论等,并阐述了鸟类栖息地片段化研究范式转变的原因。     

中国湿地物种多样性与生境面积关系及其生态学机理的模拟研究

生物群落中的物种多样性是群落生态学研究的一个基本问题。大量的实验研究表明物种多样性和生境面积的常数次幂成比例,这说明它们的对数数量级呈线性正相关。我们研究了中国境内15块湿地的高等植物和32块湿地鸟类多样性,发现它们分别和湿地面积在对数尺度上呈线性正相关．进一步支持了种数与面积的幂指数关系。我们还借助计算机模拟系统地讨论了产生这种简单规律的生态学机理,包括中性理论、集合种群动态和物种分布的自相似性。中性理论假设了群落中物种的个体之间只有竞争关系,忽略了其它的种间关系。集合种群动态理论考虑的是由多个亚群落构成的集合群落,在研究种数和面积关系时也忽略了种间关系,所以也是中性的。尽管物种分布的自相似性可导致这种面积幂指数关系,但在自然界中自相似性也可能不成立。     

Unexpected coherence and conservation.

The effects of migration in a network of patch populations, or metapopulation, are extremely important for predicting the possibility of extinctions both at a local and a global scale. Migration between patches synchronizes local populations and bestows upon them identical dynamics (coherent or synchronous oscillations), a feature that is understood to enhance the risk of global extinctions. This is one of the central theoretical arguments in the literature associated with conservation ecology. Here, rather than restricting ourselves to the study of coherent oscillations, we examine other types of synchronization phenomena that we consider to be equally important. Intermittent and out-of-phase synchronization are but two examples that force us to reinterpret some classical results of the metapopulation theory. In addition, we discuss how asynchronous processes (for example, random timing of dispersal) can paradoxically generate metapopulation synchronization, another non-intuitive result that cannot easily be explained by the standard theory.     

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.     

The dynamics of two diffusively coupled predator-prey populations

I analyze the dynamics of predator and prey populations living in two patches. Within a patch the prey grow logistically and the predators have a Holling type II functional response. The two patches are coupled through predator migration. The system can be interpreted as a simple predator-prey metapopulation or as a spatially explicit predator-prey system. Asynchronous local dynamics are presumed by metapopulation theory. The main question I address is when synchronous and when asynchronous dynamics arise. Contrary to biological intuition, for very small migration rates the oscillations always synchronize. For intermediate migration rates the synchronous oscillations are unstable and I found periodic, quasi-periodic, and intermittently chaotic attractors with asynchronous dynamics. For large predator migration rates, attractors in the form of equilibria or limit cycles exist in which one of the patches contains no prey. The dynamical behavior of the system is described using bifurcation diagrams. The model shows that spatial predator-prey populations can be regulated through the interplay of local dynamics and migration.     

Area-dependent migration by ringlet butterflies generates a mixture of patchy population and metapopulation attributes

Interpretation of spatially structured population systems is critically dependent on levels of migration between habitat patches. If there is considerable movement, with each individual visiting several patches, there is one ”patchy population”; if there is intermediate movement, with most individuals staying within their natal patch, there is a metapopulation; and if (virtually) no movement occurs, then the populations are separate (Harrison 1991, 1994). These population types actually represent points along a continuum of much to no mobility in relation to patch structure. Therefore, interpretation of the effects of spatial structure on the dynamics of a population system must be accompanied by information on mobility. We use empirical data on movements by ringlet butterflies, Aphantopus hyperantus, to investigate two key issues that need to be resolved in spatially-structured population systems. First, do local habitat patches contain largely independent local populations (the unit of a metapopulation), or merely aggregations of adult butterflies (as in patchy populations)? Second, what are the effects of patch area on migration in and out of the patches, since patch area varies considerably within most real population systems, and because human landscape modification usually results in changes in habitat patch sizes? Mark-release-recapture (MRR) data from two spatially structured study systems showed that 63% and 79% of recaptures remained in the same patch, and thus it seems reasonable to call both systems metapopulations, with some capacity for separate local dynamics to take place in different local patches. Per capita immigration and emigration rates declined with increasing patch area, while the resident fraction increased. Actual numbers of emigrants either stayed the same or increased with area. The effect of patch area on movement of individuals in the system are exactly what we would have expected if A. hyperantus were responding to habitat geometry. Large patches acted as local populations (metapopulation units) and small patches simply as locations with aggregations (units of patchy populations), all within 0.5 km². Perhaps not unusually, our study system appears to contain a mixture of metapopulation and patchy-population attributes.     

Population history and life history influence the migration rate of female Glanville fritillary butterflies

This study examines the causes of emigration from small fragments of suitable habitat in a species that has a distinct metapopulation structure, frequent turnover of local populations, and substantial migration among local populations and currently unoccupied habitat fragments. We conducted a field experiment in which 727 individuals of the Glanville fritillary butterfly ( Melitaea cinxia ) originating from four regions were marked and released simultaneously in a natural environment. In three of the four source regions, larvae for the experiment were collected from dozens of small local populations, some of which had been established in the previous summer (new populations), whereas the remaining populations were older. In two of the source regions, female butterflies prefer a host plant ( Veronica spicata ) that is not present in the release area, where there is only Plantago lanceolata , the preferred host plant of females from the other two source regions. We found that migration rate of males was unrelated to any of the factors studied in this experiment. In contrast, two factors influenced the migration rate of females. First, Veronica -preferring females had higher emigration rate than Plantago- preferring females from the Plantago -containing release patches, demonstrating that the individual perception of habitat quality significantly influences the migration rate of females. Second, females from newly-established populations were more dispersive than females from older populations, supporting the notion that metapopulation processes (recurrent colonizations) select for increased migration. The observed migration rate was not correlated with any body size measurements, and thus the observed differences in migration rate were apparently caused by differences in the behaviour of female butterflies rather than in their flight capacity.     

Allee-like effects in metapopulation dynamics

The existences of the Allee effect at the local population level and of the Allee-like effect at the metapopulation level are important for both ecology and conservation. Although there have been a great many papers on the Allee effect, they have mainly referred to only local populations and have not dealt with the relationship between the two. In this paper, we begin with local population dynamics and then construct a model including both local population and metapopulation dynamics. Then we simulate with computer at these two levels. The results indicate that the Allee-like effect in a metapopulation may emerge from the imposed Allee effect at the local population level. This threshold fraction of occupied patches below which the metapopulation goes extinct is seriously affected by the per capita migration rate, the survival rate during migration and the initial population size on the occupied patches. We also find that severe demographic stochasticity may compound the metapopulation extinction risk posed by the Allee effect. These conclusions are helpful for nature conservation, especially for the preservation of rare species.     

The applicability of metapopulation theory to large mammals

Metapopulation theory has become a common framework in conservation biology and it is sometimes suggested that a metapopulation approach should be used for management of large mammals. However, it has also been suggested that metapopulation theory would not be applicable to species with long generations compared to those with short ones. In this paper, we review how and on what empirical ground metapopulation terminology has been applied to insects, small mammals and large mammals. The review showed that the metapopulation term sometimes was used for population networks which only fulfilled the broadest possible definition of a metapopulation, i.e. they were subpopulations connected by migrating individuals. We argue that the metapopulation concept should be reserved for networks that also show some kind of metapopulation dynamics. Otherwise it applies to almost all populations and loses its substance. We found much empirical support for metapopulation dynamics in both insects and small mammals, but not in large mammals. A possible reason is the methods used to confirm the existence of metapopulation dynamics. For insects and small mammals, the common approach is to study population turnover through patch occupancy data. Such data is difficult to obtain for large mammals, since longer temporal scales need to be covered to record extinctions and colonizations. Still, many populations of large mammals are exposed to habitat fragmentation and the resulting subpopulations sometimes have high risks of extinction. If there is migration between the subpopulations, the metapopulation framework could provide valuable information on their population dynamics. We suggest that a metapopulation approach can be interesting for populations of large mammals, when there are discrete breeding subpopulations and when these subpopulations have different growth rates and demographic fates. Thus, a comparison of the subpopulations’ demographic fates, rather than subpopulation turnover, can be a feasible alternative for studies of metapopulation dynamics in large mammals.     

Linking extinction-colonization dynamics to genetic structure in a salamander metapopulation

Theory predicts that founder effects have a primary role in determining metapopulation genetic structure. However, ecological factors that affect extinction-colonization dynamics may also create spatial variation in the strength of genetic drift and migration. We tested the hypothesis that ecological factors underlying extinction-colonization dynamics influenced the genetic structure of a tiger salamander (Ambystoma tigrinum) metapopulation. We used empirical data on metapopulation dynamics to make a priori predictions about the effects of population age and ecological factors on genetic diversity and divergence among 41 populations. Metapopulation dynamics of A. tigrinum depended on wetland area, connectivity and presence of predatory fish. We found that newly colonized populations were more genetically differentiated than established populations, suggesting that founder effects influenced genetic structure. However, ecological drivers of metapopulation dynamics were more important than age in predicting genetic structure. Consistent with demographic predictions from metapopulation theory, genetic diversity and divergence depended on wetland area and connectivity. Divergence was greatest in small, isolated wetlands where genetic diversity was low. Our results show that ecological factors underlying metapopulation dynamics can be key determinants of spatial genetic structure, and that habitat area and isolation may mediate the contributions of drift and migration to divergence and evolution in local populations.     

The genetic effective size of a metapopulation

The structure of a population over time, space and categories of social and sexual role governs its ability to retain genetic variation in the face of drift. A metapopulation is an extreme form of spatial structure in which loosely coupled local populations 'turnover', that is, suffer extinction followed by recolonization from elsewhere within the metapopulation. These local populations turn over with a characteristic half-life. Based on a simulation model that incorporates both realistic features of population ecology and population genetics, the ability of such a metapopulation to retain genetic variation, which may be defined as proportional to its so-called effective population size, denoted N_e(^meta), can be one to two orders of magnitude lower than the maximum total number of individuals in the system. N_e(meta) depends on the persistence time associated with longevity of local populations (the turnover half-life), the average number of local populations extant in the metapopulation and the gene flow between local populations. Habitat fragmentation, which can create a metapopulation from a formerly continuously distributed species, may have unappreciated large genetic consequences for species impacted by human development.