A review of the SLOSS debate: Analysis methods, theoretical mechanisms and conservation practices

doi:10.17520/biods.2025081

Abstract

Abstract:

Background & Aim: The SLOSS debate in the field of biodiversity conservation outlines whether single large protected area (SL) or several small protected areas (SS) are preferred for the conservation of species diversity, given equal total area or limited resources. This issue has been of great interest to ecologists and conservation biologists and has important implications for the design of protected areas and biodiversity conservation. As a result of the development of research methods and the rise of interdisciplinarity, the SLOSS debate has made important multi-stage advances in both analytical methods and theoretical mechanisms. The main objective of this paper is to review the rapidly research progress of the SLOSS debate and provide some targeted suggestions for future directions.

Review results: We first described the origins of the SLOSS debate and then analyzed the SLOSS literature from different aspects, including comparisons of publications between study regions and study times. Secondly, we described the analysis methods of the SLOSS debate and the strengths and limitations of each method. In addition, we reviewed the improved saturation indices proposed by previous researchers as well as the optimized indices developed by us. Thirdly, we explored several theoretical mechanisms that influence the SLOSS pattern, e.g, the theory of extinction-colonization dynamics dominated by extinction, the theory of extinction-colonization dynamics dominated by colonization, and the role of risk resistance. We then described how SLOSS debate can be applied to guide conservation management and protected area design. Lastly, we put forward some targeted suggestions for the future direction of SLOSS debate, including the importance of using flexible analytical methods, interdisciplinary collaborations to test theoretical mechanisms, more functional SLOSS and phylogenetic SLOSS research, and integrating the SLOSS debate with other fields such as restoration ecology.

Conclusion: Our review will be helpful to gain a deeper understanding of the progress of SLOSS debate, which is of great significance for promoting the development of this research field.

Key words: protected area, species diversity, SLOSS indices, SLOSS analysis, species accumulation curve, ecological models, theoretical mechanisms

Yang Zhang, Yanping Wang. A review of the SLOSS debate: Analysis methods, theoretical mechanisms and conservation practices[J]. Biodiv Sci, 2025, 33(7): 25081.

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References 152

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概念 Concept		定义 Definition		对SLOSS争论的解释或与SLOSS争论的关联 Explanation of the SLOSS debate or correlation with the SLOSS debate	发生的尺度 Scale of occurrence	参考文献 References
SLOSS争论 SLOSS debate		①在总面积不变的情况下, 一个大的保护区(SL)还是几个小的保护区(SS)能容纳更多的物种? ②当总面积恒定时, 一个大的保护区(SL)还是几个小的保护区(SS)能最大限度地延长物种的灭绝时间、种群持久性或扩大种群规模? ① Can single large protected area (SL) or several small protected areas (SS) hold more species while keeping the total area unchanged? ② Can single large protected area (SL) or several small protected areas (SS) maximize the duration of species extinction, population persistence, or population size expansion while keeping the total area constant?		SLOSS争论需要解决的基本问题。The fundamental issue that need to be addressed in SLOSS debate.	景观 Landscape	Diamond, 1975; Simberloff & Abele, 1976; Ovaskainen, 2002; Groeneveld, 2005
生境破碎化 Habitat fragmentation		一大片栖息地转变为总面积较小的若干栖息地的过程, 这些小栖息地被不同于原始栖息地的基质隔离开来。A large expanse of habitat is transformed into a number of smaller patches of smaller total area, isolated from each other by a matrix of habitats unlike the original.		SLOSS争论是生境破碎化问题的一个特例, 但由SLOSS问题得出的多个小斑块可维持更高的生物多样性的结论不等同于生境破碎化是无害的。The SLOSS debate is a special case of the habitat fragmentation problem, but the conclusion drawn from the SLOSS problem that multiple small patches can maintain higher biodiversity does not necessarily mean that habitat fragmentation is harmless.	景观 Landscape	Wilcox & Murphy, 1985; Wilcove et al, 1986
生境丧失 Habitat loss	栖息地从景观中通过不同的方式被移除。Habitats are removed from the landscape in different ways.		生境丧失导致景观中栖息地总面积(总生境数量)的减少, 然而对于SLOSS争论的探讨则必须控制总面积不变。Habitat loss leads to a decrease in the total area of habitat (total number of habitats) in the landscape, but the discussion of the SLOSS debate must maintain the total area unchanged.		景观 Landscape	Fahrig, 2003
生境破碎化本身 Habitat fragmentation per se	一个给定的区域被划分为更多的独立区域或斑块, 并且该过程与生境丧失或面积损失无关。A given area is divided into more separate areas or patches and the process is not associated with habitat loss or area loss.		SLOSS类型的研究本质上是对生境破碎化本身的检验, 例如几个小斑块更好的结果支持了生境破碎化本身的积极影响, 且在具体分析时均需对栖息地总量进行控制。The SLOSS type studies are essentially tests of habitat fragmentation per se. For example, better result of several small patches supports the positive effects of habitat fragmentation per se, and both issues require controlling the total amount of habitat when analyzing them specifically.		景观 Landscape	Fahrig, 2003, 2017
边缘效应 Edge effect	由两个生态系统被一个突然的过渡(边缘)隔开而形成, 是两个相邻生态系统之间相互作用的结果。Formed by two ecosystems separated by an abrupt transition (edge) as a result of interactions between two adjacent ecosystems.		负的边缘效应会导致小斑块中物种的灭绝风险增加, 从而预测SL > SS; 在较为破碎化的景观中, 对于给定数量的栖息地, 正的边缘效应会对一些物种的多度产生积极影响, 可能会预测SS > SL。Negative edge effects lead to the increased extinction risk for species in small patches, thus predicting SL > SS. In more fragmented landscapes, for a given number of habitats, positive edge effect will positively influence the abundance of some species, possibly predicting SS > SL.		斑块 Patch	Murcia, 1995; Fahrig, 2002; Fletcher et al, 2018
景观连通性 Landscape connectivity	景观在多大程度上阻碍或促进了物种或资源在斑块之间的移动。The extent to which the landscape impedes or facilitates the movement of species or resources between patches.		当生境总量不变时, 更破碎化的景观中小斑块的数量更多, 这些较小斑块可以作为垫脚石以提高连通性, 从而预测SS > SL。When the total amount of habitat is held constant, there are more small patches in a more fragmented landscape, and these smaller patches can be used as stepping stones to improve connectivity, thus predicting SS > SL.		景观 Landscape	Taylor et al, 1993; Tischendorf & Fahrig, 2000; Fahrig et al, 2011
灭绝债务 Extinction debt	由于栖息地破坏或环境退化导致的物种灭绝的滞后现象。Lags in species extinction due to habitat destruction or environmental degradation.		若灭绝债务比迁入亏缺偿还的速度更快, 随着时间的推移, 小斑块的生物多样性会显著下降, 对应于SL > SS策略。If extinction debt is repaid more quickly than colonization credit, biodiversity in small patches declines significantly over time, corresponding to the SL > SS strategy.		景观 Landscape	Tilman et al, 1994; Jackson & Sax, 2010
迁入亏缺 Colonization credit	适宜生境恢复或新建后物种因扩散限制或繁殖滞后的延迟迁入现象。The phenomenon of delayed immigration of species due to diffusion restrictions or delayed reproduction after the restoration or construction of suitable habitats.		当物种在斑块中的定殖率超过灭绝率时, 几个小保护区可能通过迁入亏缺逐步恢复生物多样性, 对应于SS > SL策略。When the colonization rate of species in patches exceeds the extinction rate, several small protected areas may gradually restore biodiversity through colonization credit, corresponding to the SS > SL strategy.		景观 Landscape	Jackson & Sax, 2010
SLOSS分析 SLOSS analysis	利用实证方法、理论方法以及理论机制分析SLOSS问题。Analyzing the SLOSS issue using empirical and theoretical approaches as well as theoretical mechanisms.		通过物种累积曲线法、零模型法、理论模型法等来探讨SLOSS争论; 由SLOSS分析得出的结论可用于指导保护区设计。SLOSS debate is explored through methods such as the species cumulative curve method, the null modeling method, and the theoretical modeling method; Conclusions drawn from SLOSS analysis can be used to guide protected area design.		景观 Landscape	Ovaskainen, 2002; Groeneveld, 2005; Tjørve, 2010; Fahrig et al, 2022

概念 Concept		定义 Definition		对SLOSS争论的解释或与SLOSS争论的关联 Explanation of the SLOSS debate or correlation with the SLOSS debate	发生的尺度 Scale of occurrence	参考文献 References
SLOSS争论 SLOSS debate		①在总面积不变的情况下, 一个大的保护区(SL)还是几个小的保护区(SS)能容纳更多的物种? ②当总面积恒定时, 一个大的保护区(SL)还是几个小的保护区(SS)能最大限度地延长物种的灭绝时间、种群持久性或扩大种群规模? ① Can single large protected area (SL) or several small protected areas (SS) hold more species while keeping the total area unchanged? ② Can single large protected area (SL) or several small protected areas (SS) maximize the duration of species extinction, population persistence, or population size expansion while keeping the total area constant?		SLOSS争论需要解决的基本问题。The fundamental issue that need to be addressed in SLOSS debate.	景观 Landscape	Diamond, 1975; Simberloff & Abele, 1976; Ovaskainen, 2002; Groeneveld, 2005
生境破碎化 Habitat fragmentation		一大片栖息地转变为总面积较小的若干栖息地的过程, 这些小栖息地被不同于原始栖息地的基质隔离开来。A large expanse of habitat is transformed into a number of smaller patches of smaller total area, isolated from each other by a matrix of habitats unlike the original.		SLOSS争论是生境破碎化问题的一个特例, 但由SLOSS问题得出的多个小斑块可维持更高的生物多样性的结论不等同于生境破碎化是无害的。The SLOSS debate is a special case of the habitat fragmentation problem, but the conclusion drawn from the SLOSS problem that multiple small patches can maintain higher biodiversity does not necessarily mean that habitat fragmentation is harmless.	景观 Landscape	Wilcox & Murphy, 1985; Wilcove et al, 1986
生境丧失 Habitat loss	栖息地从景观中通过不同的方式被移除。Habitats are removed from the landscape in different ways.		生境丧失导致景观中栖息地总面积(总生境数量)的减少, 然而对于SLOSS争论的探讨则必须控制总面积不变。Habitat loss leads to a decrease in the total area of habitat (total number of habitats) in the landscape, but the discussion of the SLOSS debate must maintain the total area unchanged.		景观 Landscape	Fahrig, 2003
生境破碎化本身 Habitat fragmentation per se	一个给定的区域被划分为更多的独立区域或斑块, 并且该过程与生境丧失或面积损失无关。A given area is divided into more separate areas or patches and the process is not associated with habitat loss or area loss.		SLOSS类型的研究本质上是对生境破碎化本身的检验, 例如几个小斑块更好的结果支持了生境破碎化本身的积极影响, 且在具体分析时均需对栖息地总量进行控制。The SLOSS type studies are essentially tests of habitat fragmentation per se. For example, better result of several small patches supports the positive effects of habitat fragmentation per se, and both issues require controlling the total amount of habitat when analyzing them specifically.		景观 Landscape	Fahrig, 2003, 2017
边缘效应 Edge effect	由两个生态系统被一个突然的过渡(边缘)隔开而形成, 是两个相邻生态系统之间相互作用的结果。Formed by two ecosystems separated by an abrupt transition (edge) as a result of interactions between two adjacent ecosystems.		负的边缘效应会导致小斑块中物种的灭绝风险增加, 从而预测SL > SS; 在较为破碎化的景观中, 对于给定数量的栖息地, 正的边缘效应会对一些物种的多度产生积极影响, 可能会预测SS > SL。Negative edge effects lead to the increased extinction risk for species in small patches, thus predicting SL > SS. In more fragmented landscapes, for a given number of habitats, positive edge effect will positively influence the abundance of some species, possibly predicting SS > SL.		斑块 Patch	Murcia, 1995; Fahrig, 2002; Fletcher et al, 2018
景观连通性 Landscape connectivity	景观在多大程度上阻碍或促进了物种或资源在斑块之间的移动。The extent to which the landscape impedes or facilitates the movement of species or resources between patches.		当生境总量不变时, 更破碎化的景观中小斑块的数量更多, 这些较小斑块可以作为垫脚石以提高连通性, 从而预测SS > SL。When the total amount of habitat is held constant, there are more small patches in a more fragmented landscape, and these smaller patches can be used as stepping stones to improve connectivity, thus predicting SS > SL.		景观 Landscape	Taylor et al, 1993; Tischendorf & Fahrig, 2000; Fahrig et al, 2011
灭绝债务 Extinction debt	由于栖息地破坏或环境退化导致的物种灭绝的滞后现象。Lags in species extinction due to habitat destruction or environmental degradation.		若灭绝债务比迁入亏缺偿还的速度更快, 随着时间的推移, 小斑块的生物多样性会显著下降, 对应于SL > SS策略。If extinction debt is repaid more quickly than colonization credit, biodiversity in small patches declines significantly over time, corresponding to the SL > SS strategy.		景观 Landscape	Tilman et al, 1994; Jackson & Sax, 2010
迁入亏缺 Colonization credit	适宜生境恢复或新建后物种因扩散限制或繁殖滞后的延迟迁入现象。The phenomenon of delayed immigration of species due to diffusion restrictions or delayed reproduction after the restoration or construction of suitable habitats.		当物种在斑块中的定殖率超过灭绝率时, 几个小保护区可能通过迁入亏缺逐步恢复生物多样性, 对应于SS > SL策略。When the colonization rate of species in patches exceeds the extinction rate, several small protected areas may gradually restore biodiversity through colonization credit, corresponding to the SS > SL strategy.		景观 Landscape	Jackson & Sax, 2010
SLOSS分析 SLOSS analysis	利用实证方法、理论方法以及理论机制分析SLOSS问题。Analyzing the SLOSS issue using empirical and theoretical approaches as well as theoretical mechanisms.		通过物种累积曲线法、零模型法、理论模型法等来探讨SLOSS争论; 由SLOSS分析得出的结论可用于指导保护区设计。SLOSS debate is explored through methods such as the species cumulative curve method, the null modeling method, and the theoretical modeling method; Conclusions drawn from SLOSS analysis can be used to guide protected area design.		景观 Landscape	Ovaskainen, 2002; Groeneveld, 2005; Tjørve, 2010; Fahrig et al, 2022

指数划分 Classification of indices	SLOSS相关指数 SLOSS-related indices	特征 Characteristics	参考文献 References
原始饱和指数 Original saturation index	O_SI	计算结果容易倾向于几个小斑块, 未考虑物种占用对斑块大小的依赖性。O_SI tends to skew the calculations in favor of several small patches. It does not take into account the dependence of species occupancies on patch size.	Quinn & Harrison, 1988; Deane, 2022;
改进的饱和指数 Improved saturation index	N_SI	减少了高估分析结果的情况。Improved saturation indices avoid the overestimation of analytical results.	Cook, 1995
优化的饱和指数 Optimized saturation index	I_SU、I_DI	在Mac Nally和Lake (1999)的基础上提出的新指数, 可用于计算累积物种、功能和谱系3个维度的多样性, 并能避免对计算结果的高估或低估。 New indices based on Mac Nally and Lake (1999), which can be used to calculate the diversity of cumulative species, functional and phylogenetic dimensions and can avoid overestimation or underestimation of the results.	本文提出 This study
修正的Quinn-Harrison统计量 Modified statistics of Quinn-Harrison method	ξ、η	ξ可以衡量从小到大和从大到小方向上累积多样性的差异; η能够更直观地判断SL还是SS在系统中占主导地位。ξ measures the difference in cumulative diversity in the small to large and large to small directions. η enables a more intuitive determination of whether SL or SS dominates the system.	Mac Nally & Lake, 1999
外推的SLOSS指数 Extrapolated SLOSS index	SL_EX	SL_EX减少了抽样偏差带来的影响; 还能反映生境破碎化对物种丰富度的影响。SL_EXreduces the impact of sampling bias. It can also reflect the impact of habitat fragmentation on species richness.	Boecklen,1997; Gavish et al, 2012

指数划分 Classification of indices	SLOSS相关指数 SLOSS-related indices	特征 Characteristics	参考文献 References
原始饱和指数 Original saturation index	O_SI	计算结果容易倾向于几个小斑块, 未考虑物种占用对斑块大小的依赖性。O_SI tends to skew the calculations in favor of several small patches. It does not take into account the dependence of species occupancies on patch size.	Quinn & Harrison, 1988; Deane, 2022;
改进的饱和指数 Improved saturation index	N_SI	减少了高估分析结果的情况。Improved saturation indices avoid the overestimation of analytical results.	Cook, 1995
优化的饱和指数 Optimized saturation index	I_SU、I_DI	在Mac Nally和Lake (1999)的基础上提出的新指数, 可用于计算累积物种、功能和谱系3个维度的多样性, 并能避免对计算结果的高估或低估。 New indices based on Mac Nally and Lake (1999), which can be used to calculate the diversity of cumulative species, functional and phylogenetic dimensions and can avoid overestimation or underestimation of the results.	本文提出 This study
修正的Quinn-Harrison统计量 Modified statistics of Quinn-Harrison method	ξ、η	ξ可以衡量从小到大和从大到小方向上累积多样性的差异; η能够更直观地判断SL还是SS在系统中占主导地位。ξ measures the difference in cumulative diversity in the small to large and large to small directions. η enables a more intuitive determination of whether SL or SS dominates the system.	Mac Nally & Lake, 1999
外推的SLOSS指数 Extrapolated SLOSS index	SL_EX	SL_EX减少了抽样偏差带来的影响; 还能反映生境破碎化对物种丰富度的影响。SL_EXreduces the impact of sampling bias. It can also reflect the impact of habitat fragmentation on species richness.	Boecklen,1997; Gavish et al, 2012

理论模型划分 Category of theoretical models	特征 Characteristics	目标预测 Target predictions	研究案例 Case studies
经济模型 Economic models	同时考虑生态目标和经济成本 Considering both ecological objectives and economic costs	综合生态和经济效益设计最优保护区 Integration of ecological and economic benefits in the design of optimal protected areas	Groeneveld, 2005; Helmstedt et al, 2014
空间方差结构的超种群模型 Super-population model of the spatial variance structure	可分析误差范围、物种的空间分布和经济成本 Ranges of error, spatial distribution of species and economic costs can be analyzed	最大成本限制下的最优保护区配置 Optimal protected area allocation under maximum cost constraints	Picard et al, 2010
集合种群模型 Metapopulation model	特别关注物种的扩散和迁移 Particularly concerned about the dispersal and movement of species	物种灭绝风险或种群持久性 Species extinction risk or population persistence	Ovaskainen, 2002; McCarthy et al, 2005; Zhou & Wang, 2006
随机灭绝模型 Stochastic extinction model	允许随机性过程如随机灭绝事件的发生 Allow for stochastic processes such as random extinction events	灭绝风险 Extinction risk	Wright & Hubbell, 1983; Burkey, 1989
微观实验模型 Microscopic experimental model	模式生物为原生生物且实验条件可控 Model organisms are protists and experimental conditions are controlled	物种灭绝数量 Number of extinct species	Hammill & Clements, 2020; Wolfe et al, 2022
空间隐式模型 Spatially implicit model	可识别联系种群和景观的一般模式 General patterns of linked populations and landscapes can be recognized	种群生存能力 Population viability	Pichancourt et al, 2006
计算机模拟模型 Computer simulation model	允许调整多个参数以模拟不同的生态场景 Allow adjusting multiple parameters to simulate different ecological scenarios	种群生存能力 Population viability	Liu et al, 2017; Pelletier, 2000
物种分布模拟模型 Distribution simulated model	允许改变物种分布的不同组成部分 Allow changes to the different components of species distribution	物种存活和灭绝概率 Species survival and extinction probabilities	May et al, 2019
空间分配模型 Spatial allocation model	可应用于不同类型的斑块和物种 Can be applied to different types of patches and species	种群可持续性 Population sustainability	van Langevelde et al, 2002
空间均衡模型 Spatial equilibrium model	关注生态、经济和环境政策因素 Ecological, economic, and environmental policy factors are concerned	灭绝风险 Extinction risk	Eppink & Withagen, 2005
基于个体的模型 Individual-based model	强调个体行为和运动对种群动态的影响 Emphasize the effects of individual behavior and movement on population dynamics	种群规模 Population size	Fahse et al, 1998; Nakajima & Boggs, 2015
空间显式模型 Spatially explicit model	关注物种在空间上的扩散且可与其他模型结合使用 Focus on spatial dispersal of species and can be used in conjunction with other models	种群持久性 Population persistence	Robert, 2009; Evans et al, 2013; Niebuhr et al, 2015
多斑块模型 Multi-patch model	可调整物种迁移率的变化 Can adjust the changes in species migration rate	种群规模 Population size	Elbetch et al, 2022
渔业模型 Fishery model	可应用于海洋保护区且通常涉及捕捞努力的影响 Can be applied to marine protected areas and typically involves the impact of fishing efforts	种群规模 Population size	Kritzer, 2004; Moussaoui & Auger, 2015
生态系统模型 Ecosystem model	允许在广泛的空间尺度和各种类型的情景下运行 Allow operation in a wide range of spatial scales and various types of scenarios	综合生态和经济效益设计最优保护区 Integration of ecological and economic benefits in the design of optimal protected areas	Salomon et al, 2002
景观模拟模型 Landscape simulation model	允许评估景观指标的变化且可模拟多种场景 Allow for the evaluation of changes in landscape indicators and can simulate multiple scenarios	灭绝风险和生物多样性 Extinction risk and biodiversity	Watts & Hughes, 2024