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。     

兰科植物种群动态研究进展

兰科植物种群动态研究中,种群统计学分析能够很好地揭示植物个体在时空上的变化,是研究种群动态的核心.在自然生境中,许多附生兰科植物更倾向于离散或斑块状分布,可以通过集合种群研究分析斑块之间个体的基因流动,判断物种种群保护的规模.长期的种群动态研究能够获得兰科植物生活史和种群动态方面的可靠信息,以及一定环境条件下其时空波动及与种群功能之间的关系;短期的研究能够更好地理解具有结构性的独立植株与其所处的群落间的关系.本文根据种群生态学原理以及兰科植物的生态特点,从种群的密度及分布、种群统计学、种群的调节、集合种群和种群生存力分析（PVA）模型等方面阐述了国内外兰科植物种群动态研究进展.     

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。     

荒漠破碎化生境中长爪沙鼠集合种群野外验证研究

近年来,人类活动和自然干扰,导致内蒙古阿拉善荒漠区生境的破碎化,出现了长爪沙鼠在不同斑块间的不连续分布,每一斑块内可能存在一个局域种群,而集合种群建立的前提条件,是局域种群斑块状分布在离散的栖息地环境中。2002~2012年每年的4~10月,在阿拉善荒漠区禁牧、轮牧、过牧和开垦4种人为不同利用方式形成的生境斑块中,采用标志重捕法对长爪沙鼠（Meriones unguiculatus）种群进行定点监测。通过分析长爪沙鼠种群动态,计算各局域种群的灭绝风险,利用Spearman秩相关系数检验种群动态的空间同步性,同时以种群周转率对长爪沙鼠扩散能力进行评估,以检验阿拉善荒漠区长爪沙鼠种群空间结构是否具有经典集合种群的功能。结果表明：（1） 不同生境斑块可被长爪沙鼠局域种群占据,11年间捕获长爪沙鼠2~7次不等;（2） 长爪沙鼠所有局域种群均具有灭绝风险,在轮牧区和禁牧区灭绝率高达1.000 0,开垦区灭绝率最低,也达到0.333 4,而本研究期间最大局域种群（2008年过牧区,26只/hm²）,在2010年发生了局域灭绝;（3） 不同生境斑块间没有明显的空间隔离而阻碍局域种群的重新建立,长爪沙鼠扩散能力较强,绝大部分月份的种群周转率在50.0%以上,特别是周转率达到100.0%的月份较多;（4） 不同生境斑块间仅轮牧区和禁牧区中长爪沙鼠种群密度显著正相关（P<0.05）,而其他生境斑块间相关性均不显著（P >0.05）,长爪沙鼠局域种群整体显示出明显的非同步空间动态。阿拉善荒漠区长爪沙鼠种群满足作为经典集合种群物种区域续存的4个条件,具有作为研究小哺乳动物集合种群的潜在价值。     

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。     

集合种群模拟模型及其在竞争共存机制与物种多样性研究中的应用

以荒漠区人工植被的恢复与重建为背景,从宏观尺度研究了很集合种群的空间分布新模式,建立了基于Levins集合种群模型的数值模拟方法。对两物种的模拟结果表明：在适当选择参数下,模拟植被区的集合种群可以形成“海藻式”稳定的时空分布结构,在理论上表明相同生态特征的物种在空间生境中可以达成共存。为了达到物种丰富度和生产力最佳,实现持续发展,对多物种集合种群进行了模拟。模拟结果显示当物种的种数为5时,空间上随机播种的模拟种群覆盖率达到最大,因而可发挥最大的治沙作用。另外,模拟还显示在播种时应采取集聚式的空间播种模式,以使种群具有较高的防沙能力。该结果可为生物防沙治沙领域提供理论依据。     

生境变化对集合种群系统生态效应的影响

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。     

用空间直观模型是否足以从斑块占据性资料中推断集合种群的动态过程?

在集合种群的研究中,经常要根据空间占据性数据应用斑块模型来推断种群的动态过程,在保护生物学应用中,斑块占据性模型的参数估测对于阐释集合种群动态和预测种群对生境破坏的反应极为重要。我们探讨了一种广泛应用的空间直观模型——率函数模型(Incidence function model)中参数估测的不确定性问题,通过构建由50个斑块组成的网络和两个假想的已知参数的集合种群,应用模拟模型产生集合种群随时间变化的斑块占据性数据系列：即快照(snapshot)。然后,根据这些快照,应用率函数模型和最大似然法估测种群动态参数。此外,我们还给出了传统的率函数模型的一个变形,这个变形包含了目标区效应(Target area effect)：即一个斑块的占据概率不但取决于空间隔离度,也取决于斑块本身面积的大小。结果表明：根据同一个集合种群不同的快照所估测的参数可以有很大差异,一个快照得出的参数提示的是占据性强但存活率低的集合种群,而另一个快照可能反映的是一个占据性弱但存活率高的集合种群。应用传统的率函数模型于一个包含了目标区效应的集合种群,导致斑块大小相关的灭绝率参数估测的正偏差。因此,仅根据一个快照的空间占据性数据来推测集合种群的过程有很大的不确定性[动物学报49(6)：787～794,2003]。     

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.     

Optimal harvesting strategies for a metapopulation

We consider optimal strategies for harvesting a population that is composed of two local populations. The local populations are connected by the dispersal of juveniles, e.g. larvae, and together form a metapopulation. We model the metapopulation dynamics using coupled difference equations. Dynamic programming is used to determine policies for exploitation that are economically optimal. The metapopulation harvesting theory is applied to a hypothetical fishery and optimal strategies are compared to harvesting strategies that assume the metapopulation is composed either of single unconnected populations or of one well-mixed population. Local populations that have high per capita larval production should be more conservatively harvested than would be predicted using conventional theory. Recognizing the metapopulation structure of a stock and using the appropriate theory can significantly improve economic gains.     

Costly dispersal can destabilize the homogeneous equilibrium of a metapopulation

I investigate the stability of the homogeneous equilibrium of a discrete-time metapopulation assuming costly dispersal with arbitrary (but fixed) spatial pattern of connectivity between the local populations. First, I link the stability of the metapopulation to the stability of a single isolated population by proving that the homogeneous metapopulation equilibrium, provided that it exists, is stable if and only if a single population, which is subject to extra mortality matching the average dispersal-induced mortality of the metapopulation, has a stable fixed point. Second, I demonstrate that extra mortality may destabilize the fixed point of a single population. Taken together, the two results imply that costly dispersal can destabilize the homogeneous equilibrium of a metapopulation. I illustrate this by simulations and discuss why earlier work, arriving at the opposite conclusion, was flawed.     

Metapopulation dynamics: brief history and conceptual domain

We review the early development of metapopulation ideas, which culminated in the well-known model by Levins in 1969. We present a survey of metapopulation terminology and outline the kinds of studies that have been conducted on single-species and multispecies metapopulations. Metapopulation studies have important conceptual links with the equilibrium theory of island biogeography and with studies on the dynamics of species living in patchy environments. Metapopulation ideas play an increasingly important role in landscape ecology and conservation biology.     

The contribution of patch topology and demographic parameters to population viability analysis predictions: the case of the European tree frog

Population viability analyses (PVA) are increasingly used in metapopulation conservation plans. Two major types of models are commonly used to assess vulnerability and to rank management options: population-based stochastic simulation models (PSM such as RAMAS or VORTEX) and stochastic patch occupancy models (SPOM). While the first set of models relies on explicit intrapatch dynamics and interpatch dispersal to predict population levels in space and time, the latter is based on spatially explicit metapopulation theory where the probability of patch occupation is predicted given the patch area and isolation (patch topology). We applied both approaches to a European tree frog (Hyla arborea) metapopulation in western Switzerland in order to evaluate the concordances of both models and their applications to conservation. Although some quantitative discrepancies appeared in terms of network occupancy and equilibrium population size, the two approaches were largely concordant regarding the ranking of patch values and sensitivities to parameters, which is encouraging given the differences in the underlying paradigms and input data.     

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.     

Metapopulation models for seasonally migratory animals

Metapopulation models are widely used to study species that occupy patchily distributed habitat, but are rarely applied to migratory species, because of the difficulty of identifying demographically independent subpopulations. Here, we extend metapopulation theory to describe the directed seasonal movement of migratory populations between two sets of habitat patches, breeding and non-breeding, with potentially different colonization and extinction rates between patch types. By extending the classic metapopulation model, we show that migratory metapopulations will persist if the product of the two colonization rates exceeds the product of extinction rates. Further, we develop a spatially realistic migratory metapopulation model and derive a landscape metric-the migratory metapopulation capacity-that determines persistence. This new extension to metapopulation theory introduces an important tool for the management and conservation of migratory species and may also be applicable to model the dynamics of two host-parasite systems.     

Habitat fragmentation and population extinction of birds

It has not been established that a major cause of extinction in birds or any other taxa is failure of metapopulation dynamics: the collapse of a network of ephemeral but discrete populations as movement between them becomes increasingly infrequent. The few data on who goes where and who mates with whom suggest that most species are structured as either a single large population or a small set of source populations and a larger set of sinks. The extinction of the latter is irrelevant to the persistence of the species. However, regional decline of a species in the face of habitat destruction and fragmentation can mimic a failure of metapopulation dynamics, because distinct aggregations of individuals will disappear much as they would if populations in an interacting network were eliminated one by one. Any species with highly restricted range is at great risk of extinction from spatially localized forces, such as cyclones or deforestation. Restricted range rather than inherent weakness is the main reason that so many island species have gone extinct or are endangered. Species with small populations in contact with much larger heterospecific ones with which they are interfertile are threatened with extinction by hybridization. Finally, the disappearance of a species from a site may be due to subtle habitat change, even if this observation seems superficially consistent with some general population theory, such as the dynamic equilibrium theory of island biogeography. Current theory is an inadequate substitute for intensive field studies as a means to address the conservation problems of individual species.     

How much does an individual habitat fragment contribute to metapopulation dynamics and persistence?

We derive measures for assessing the value of an individual habitat fragment for the dynamics and persistence of a metapopulation living in a network of many fragments. We demonstrate that the most appropriate measure of fragment value depends on the question asked. Specifically, we analyse four alternative measures: the contribution of a fragment to the metapopulation capacity of the network, to the equilibrium metapopulation size, to the expected time to metapopulation extinction and the long-term contribution of a fragment to colonization events in the network. The latter measure is comparable to density-dependent measures in general matrix population theory, though some differences are introduced by the fact that "density dependence" is spatially localized in the metapopulation context. We show that the value of a fragment depends not only on the properties of the landscape but also on the properties of the species. Most importantly, variation in fragment values between the habitat fragments is greatest in the case of rare species that occur close to the extinction threshold, as these species are likely to be restricted to the most favorable parts of the landscape. We expect that the measures of habitat fragment value described and analysed here have applications in landscape ecology and in conservation biology.