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。     

自然保护区学说与麦克阿瑟-威尔逊理论

全球正两临着一场物种灭绝的大危机,自然保护己成为人类面临的重大课题,自然保护区的建立是减缓物种灭绝速率、保护生物多样性的重要途径。岛屿生物地理学为自然保护区理论的产生起了重要的作用。然而,许多研究表明,该学科尚未成熟,尤其在自然保护区方面的应用颇为有限,无疑,现代自然保护区理论应是多学科研究的综合产物。本文概述了岛屿生物地理学的基本内容及其在自然保护方面的影响,并对种群脆性分析和景观生态学途径在自然保护区理论中的作用进行讨论。     

自然保护区学说与麦克阿瑟-威尔逊理论

全球正两临着一场物种灭绝的大危机,自然保护己成为人类面临的重大课题,自然保护区的建立是减缓物种灭绝速率、保护生物多样性的重要途径。岛屿生物地理学为自然保护区理论的产生起了重要的作用。然而,许多研究表明,该学科尚未成熟,尤其在自然保护区方面的应用颇为有限,无疑,现代自然保护区理论应是多学科研究的综合产物。本文概述了岛屿生物地理学的基本内容及其在自然保护方面的影响,并对种群脆性分析和景观生态学途径在自然保护区理论中的作用进行讨论。     

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。     

岛屿生物地理学理论在物种保育方面的应用误区

岛屿生物地理学理论疏忽岛屿与大陆环境存在的实质性差异:岛屿是由性质均一的海水环绕,导致岛屿周边气流温度、湿度等因子较一致,使得岛屿环境多样性降低.岛屿环境单一化和气候变化是物种较少和易灭绝主要原因.岛屿生物地理学理论在物种保育方面的应用误区,主要是片面强调自然保护区面积效应和距离效应,忽视气候变化等产生的环境单一化对物种濒危作用.说明岛屿生物地理学理论必须修正,只有将岛屿环境单一化以及物种脆弱性作用等方面纳入理论框架内,才是正确理论和可应用于物种保育.对自然保护区设置和物种保育提出新思路和应用方法.     

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。     

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。     

集合种群动态:理论与应用

集合种群是指一相对独立地理区域内各个局域种群的集合,这些局域种群通过一定程度的个体迁移而连结在一起。集合种群理论是生态学最新分支领域——空间生态学的主要研究途径之一;它关注的是局域种群之间个体迁移的动力学后果,以及具有不稳定局域种群物种的区域续存的条件。本文较为全面地介绍了集合种群理论的基本内容,并展望其应用前景。     

种群生存力分析（PVA）的方法与应用

随着人们对资源的加速利用,生境丧失和破碎化导致物种濒危问题日益严重.以岛屿生物地理学为理论起源的种群生存力分析（PVA）,通过分析和模拟种群动态过程并建立灭绝概率与种群数量之间的关系,为濒危物种保护提供了重要的理论依据和研究途径.在过去的几十年中,种群生存力分析已成为保护生物学中一项重要的研究内容.目前种群生存力分析发展稳定,但对其实际预测能力和准确性尚存质疑,应用方面也有待进一步发展.种群生存力分析的进一步完善还需要在理论和方法上的创新,特别是籍于景观生态学和可持续性科学的理念,将空间分析手段、经济社会因素纳入到物种和种群的预测和管理上,从而使其 具有更完整的理论基础和更高的实用价值.为此,本文对种群生存力分析的历史、基本概念、研究方法、模型应用和准确性进行了综述,并提出了有关的研究展望.     

生物多样性保护的一个理论框架

稳定而复杂多样的自然生境有利于多样性的形成和保存,剧变并趋于简单化的干扰生境常使多样性丧失。现实中的森林破碎化使区域物种丧失。多样性保护要求具备促使物种能长久生存的生境。生物最小面积概念集中讨论物种长久生存与群落（景观）面积的关系,是多样性保护的最基本的理论基础。根据自然保护实践,提出最小景观,扩充了生物最小面积概念。讨论了生物最小面积概念在建立自然保护区的理论框架、了解被保护生物及其生境的自然特点以及建立更全面的自然保护网络等方面的应用     

The importance of matrix quality in fragmented landscapes: Understanding ecosystem collapse through a combination of deterministic and stochastic forces

Increasing fragmentation of natural landscapes due to anthropogenic forces are changing the ecological structure of many systems. The corpus of literature has addressed the fact that there are many complex mechanisms that may lead to population and ecosystem collapse. Here, we propose an additional mechanism that combines the deterministic framework of complex oscillatory phenomena with the stochastic framework based on the spatial phenomena of island biogeography and metapopulation theory. Combining these two frameworks, which we believe exist in disturbed natural systems, we construct a situation in which collections of competitors are controlled by specialist natural enemies. Species are isolated in fragmented habitats separated by a matrix of decreasing ecological quality, which will determine immigration potential. In this system, species are exposed to stochastic extinction of spatial immigration potential and deterministic extinction of standard oscillatory frameworks. Through the combination of these extinction events, we see that many natural systems may become increasingly unstable and subject to unexpected consequences with changes to the matrix environment. The combined effect of these two extinctions pathways highlights the need to maintain high immigration rates in order to offset possible ecosystem collapse in fragmented systems. The results highlight the importance of landscape conservation efforts, especially towards maintaining matrix environments with greater ecological integrity.     

Do forest plants conform to the theory of island biogeography: the case study of bog islands

Biodiversity conservation is confronted with increasing risk of extinction in isolated small-area remnants and the limitation of species to colonize recently formed habitats. We hypothesized that the equilibrium pattern of forest herb layer in long-term fragmented landscape should comply with the theory of island biogeography. Forests on mineral soil islands located in large mires of western Estonia were considered as dispersal target habitats, and forests on mainland and peninsulas in mires as sources. Species richness was the lowest in mainland forests and the effect was confounded by habitat structure, suggesting a negative effect of silvicultural management in easily accessible forests. We observed the ‘small island effect’, i.e. greater overall species richness in small-area habitats, which was determined by the habitat preference of shade tolerant generalists. The average species richness of common mainland forest specialists varied little, but capitalizing on the traditional approach and analyzing only island data, weak effects of distance and habitat quality were detected. At single species level, unexpectedly, many habitat specialists were observed to have successfully dispersed to islands, indicating insufficient knowledge of the long-distance dispersal mechanisms of forest-dwelling plants. In fragmented forest landscapes the theory of island biogeography can be applied to habitat specialist plant species, but only regarding the effect of isolation and in conditions of persistent forest structural quality. In the light of global changes, optimized conservation planning should primarily target on (i) the conservation of ancient habitat fragments independent of their current area, and (ii) the promotion of diversity of potential dispersal vectors in the landscape.     

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.     

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.     

Is the matrix a sea? Habitat specificity in a naturally fragmented landscape

Abstract.  1. Metapopulation and island biogeography theory assume that landscapes consist of habitat patches set in a matrix of non-habitat. If only a small proportion of species conform to the patch–matrix assumptions then metapopulation theory may only describe special cases rather than being of more general ecological importance.
2. As an initial step towards understanding the prevalence of metapopulation dynamics in a naturally fragmented landscape, the distribution of beetle species in three replicates of three habitat types was examined, including rainforest and eucalypt forest (the habitat patches), and buttongrass sedgeland (the matrix), in south-west Tasmania, Australia.
3. Ordination methods indicated that the buttongrass fauna was extremely divergent from the fauna of forested habitats. Permutation tests showed that the abundance of 13 of 17 commonly captured species varied significantly among habitats, with eight species confined to eucalypts or rainforest, and three species found only in buttongrass. Approximately 60% of species were confined to forested habitat implying that metapopulation theory has the potential to be very important in the forest–buttongrass landscape.
4. Although floristically the rainforest and eucalypts were extremely distinct, the beetle faunas from eucalypts and rainforests overlapped substantially. Therefore rainforest patches connected by eucalypt forest represent continuous habitat for most species.
5. Other studies report a wide range of values for the proportion of patch-specific species in fragmented landscapes. Understanding the environmental or historical conditions under which a high proportion of species become patch specialists would help to identify where spatial dynamic theory may be especially applicable, and where habitat loss and fragmentation poses the greatest threat to biodiversity.     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

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.     

Factors determining mammal species richness on habitat islands and isolates: habitat diversity, disturbance, species interactions and guild assembly rules

• 1 For over three decades the equilibrium theory of island biogeography has galvanized studies in ecological biogeography. Studies of oceanic islands and of natural habitat islands share some similarities to continental studies, particularly in developed regions where habitat fragmentation results from many land uses. Increasingly, remnant habitat is in the form of isolates created by the clearing and destruction of natural areas. Future evolution of a theory to predict patterns of species abundance may well come from the application of island biogeography to habitat fragments or isolates.
• 2 In this paper we consider four factors other than area and isolation that influence the number and type of mammal species coexisting in one place: habitat diversity, habitat disturbance, species interactions and guild assembly rules. In all examples our data derive from mainland habitat, fragmented to differing degrees, with different levels of isolation.
• 3 Habitat diversity is seen to be a good predictor of species richness. Increased levels of disturbance produce a relatively greater decrease in species richness on smaller than on larger isolates. Species interactions in the recolonization of highly disturbed sites, such as regenerating mined sites, is analogous to island colonization. Species replacement sequences in secondary successions indicate not just how many, but which species are included. Lastly, the complement of species established on islands, or in insular habitats, may be governed by guild assembly rules. These contributions may assist in taking a renewed theory into the new millennium.

     

Changing perspectives on biodiversity conservation: from species protection to regional sustainability

Biodiversity is the basis for ecosystem goods and services that provide for human survival and prosperity. With a rapidly increasing human population and its demands for natural resources, landscapes are being fragmented, habitats are being destroyed, and biodiversity is declining. How can biodiversity be effectively conserved in the face of increasing human pressures? In this paper, Ⅰ review changing perspectives on biodiversity conservation, and discuss their relevance to the practice of biodiversity conservation. The major points include The notion of balance of nature is a myth rather than a scientific concept; the theory of island biogeography is useful heuristically but flawed practically; the SLOSS debate is intriguing in theory but irrelevant in reality; the concept of minimum viable population and population viability analysis are useful, but technically inefficient and conceptually inadequate; metapopulation theory is mathematically elegant but ecologically oversimplistic; and integrative perspectives and approaches for biodiversity conservation are needed that incorporate insights from landscape ecology and sustainability science. Ⅰ further discuss some key principles for regional conservation planning, and argue that the long-term success of biodiversity conservation in any region will ultimately depend on the economic and social sustainability of that region. Both research and practice in biodiversity conservation, therefore, need to adopt a broader perspective of sustainability.