How to resolve the SLOSS debate: Lessons from species-diversity models

The SLOSS debate - whether a single large reserve will conserve more species than several small - of the 1970s and 1980s never came to a resolution. The first rule of reserve design states that one large reserve will conserve the most species, a rule which has been heavily contested. Empirical data seem to undermine the reliance on general rules, indicating that the best strategy varies from case to case. Modeling has also been deployed in this debate. We may divide the modeling approaches to the SLOSS enigma into dynamic and static approaches. Dynamic approaches, covered by the fields of island equilibrium theory of island biogeography and metapopulation theory, look at immigration, emigration, and extinction. Static approaches, such as the one in this paper, illustrate how several factors affect the number of reserves that will save the most species.This article approaches the effect of different factors by the application of species-diversity models. These models combine species-area curves for two or more reserves, correcting for the species overlap between them. Such models generate several predictions on how different factors affect the optimal number of reserves. The main predictions are: Fewer and larger reserves are favored by increased species overlap between reserves, by faster growth in number of species with reserve area increase, by higher minimum-area requirements, by spatial aggregation and by uneven species abundances. The effect of increased distance between smaller reserves depends on the two counteracting factors: decreased species density caused by isolation (which enhances minimum-area effect) and decreased overlap between isolates. The first decreases the optimal number of reserves; the second increases the optimal number. The effect of total reserve-system area depends both on the shape of the species-area curve and on whether overlap between reserves changes with scale.The approach to modeling presented here has several implications for conservational strategies. It illustrates well how the SLOSS enigma can be reduced to a question of the shape of the species-area curve that is expected or generated from reserves of different sizes and a question of overlap between isolates (or reserves).     

Nestedness, biogeographic theory, and the design of nature reserves

I examine the relationship between nested distributional patterns and the degree to which several small reserves will contain more species than would a single reserve of equal total area (SLOSS). Nestedness is a common property of species distributions on real and habitat islands. However, there is considerable variation in nestedness among species distributions, some of which is related to the physical and biological background of the archipelagoes. Nestedness does not vary according to the taxonomic group examined (with the exception of aquatic invertebrates). Nestedness does vary between real and habitat islands (with aquatic invertebrates excluded), but not between oceanic and land-bridge islands. The more a biota is nested, the more likely it is that a single large reserve would preserve more species. However, nestedness is a rather poor predictor of SLOSS, as the vast majority of archipelagoes support a strategy of several small reserves, even though almost all of them are significantly nested. Nestedness says little about optimal reserve design and management, and appears to be a weak conservation tool. Received: 30 May 1995 / Accepted: 1 April 1997     

Amazonian biodiversity and protected areas: do they meet?

Protected areas are crucial for Amazonian nature conservation. Many Amazonian reserves have been selected systematically to achieve biodiversity representativeness. We review the role natural-scientific understanding has played in reserve selection, and evaluate the theoretical potential of the existing reserves to cover a complete sample of the species diversity of the Amazonian rainforest biome. In total, 108 reserves (604,832 km²) are treated as strictly protected and Amazonian; 87 of these can be seen as systematically selected to sample species diversity (75.3% of total area). Because direct knowledge on all species distributions is unavailable, surrogates have been used to select reserves: direct information on some species distributions (15 reserves, 14.8% of total area); species distribution patterns predicted on the basis of conceptual models, mainly the Pleistocene refuge hypothesis, (5/10.3%); environmental units (46/27.3%); or a combination of distribution patterns and environmental units (21/22.9%). None of these surrogates are reliable: direct information on species distributions is inadequate; the Pleistocene refuge hypothesis is highly controversial; and environmental classifications do not capture all relevant ecological variation, and their relevance for species distribution patterns is undocumented. Hence, Amazonian reserves cannot be safely assumed to capture all Amazonian species. To improve the situation, transparency and an active dialogue with the scientific community should be integral to conservation planning. We suggest that the best currently available approach for sampling Amazonian species diversity in reserve selection is to simultaneously inventory indicator plant species and climatic and geological conditions, and to combine field studies with remote sensing.     

Connectivity, Probabilities and Persistence: Comparing Reserve Selection Strategies

Reserve selection methods are often based on information on species’ occurrence. This can be presence–absence data, or probabilities of occurrence estimated with species distribution models. However, the effect of the choice of distribution model on the outcome of a reserve selection method has been ignored. Here we test a range of species distribution models with three different reserve selection methods. The distribution models had different combinations of variables related to habitat quality and connectivity (which incorporates the effect of spatial habitat configuration on species occurrence). The reserve selection methods included (i) a minimum set approach without spatial considerations; (ii) a clustering reserve selection method; and (iii) a dynamic approach where probabilities of occurrence are re-evaluated according to the spatial pattern of selected sites. The sets of selected reserves were assessed by re-computing species probability of occurrence in reserves using the best probability model and assuming loss of non-selected habitat. The results show that particular choices of distribution model and selection method may lead to reserves that overestimate the achieved target; in other words, species may seem to be represented but the reserve network may actually not be able to support them in the long-term. Instead, the use of models that incorporated connectivity as a variable resulted in the selection of aggregated reserves with higher potential for species long-term persistence. As reserve design aims at the long-term protection of species, it is important to be aware of the uncertainties related to model and method choice and their implications.     

Nested distribution of amphibians in the Zhoushan archipelago, China: can selective extinction cause nested subsets of species?

Because of their poor dispersal ability, amphibians are well suited for testing the selective extinction theory on islands. Amphibian fauna in the Zhoushan archipelago, China, exhibit a high level of nestedness (C = 0.893), and the species number is lower on islands than on similar sized areas on the mainland. No correlation was found between island-specific species richness and the nearest distance from a larger island, distance from the mainland or density of human population. These results suggest that no amphibian colonisation has occurred in the archipelago since island isolation 7000–9000 years ago. Furthermore, the results imply that selective extinction contributes to the nestedness of amphibians in the Zhoushan archipelago. The incidence of a species on the islands is significantly correlated with log area of the smallest island occupied by the species and the number of provinces on the Chinese mainland in which the species occur. However, there is no correlation with average body length of adults and island occurrence. It is concluded that (1) the area of the smallest island occupied by a species is a good estimate of the minimum area for a viable population of the species and a good predictor of species incidence on islands, (2) species with a restricted distribution range are more vulnerable to extinction from islands than those with a wide distribution range and (3) the effect of body size on occurrence on the islands is uncertain, and may be specific to the archipelago and taxa studied. The observed nestedness of amphibian assemblages has two implications for conservation: (1) not only can all the species found in several small reserves be found on a large reserve of the same total size, but additional species can be found on the single large reserve; (2) for a reserve to maintain viable populations of all species in a region it should be at least as large as the smallest island occupied by the most vulnerable species. Received: 16 December 1996 / Accepted: 22 September 1997     

江西省自然保护区发展布局空缺分析

通过专家咨询、数量标准化、主观赋权等方法,对江西省96个县(市、区)的高等动植物种类、珍稀物种种类、天然林面积和蓄积、天然湿地面积、已建自然保护区面积等进行评分,计算96个县(市、区)的物种多样性、天然林资源、天然湿地等自然资源的综合评分值和自然保护区建设的空缺评分值,并借助GIS技术分析了江西省自然保护区空间布局特点及存在的主要问题,研究提出了自然保护区空间规划布局建议,为江西省自然保护区中远期发展规划提供科学依据。     

How Well Will Brazil's System of Atlantic Forest Reserves Maintain Viable Bird Populations?

It is crucial for biodiversity conservation that protected areas are large and effective enough to support viable populations of their original species. We used a point count distance sampling method to estimate population sizes of a range of bird species in three Atlantic forest protected areas of size 5600, 22,500, and 46,050 ha. Population sizes were generally related to reserve area, although in the mid-sized reserve, there were many rare species reflecting a high degree of habitat heterogeneity. The proportions of forest species having estimated populations >500 ranged from 55% of 210 species in the largest reserve to just 25% of 140 species in the smallest reserve. All forest species in the largest reserves had expected populations >100, but in the small reserve, 28% (38 species) had populations <100 individuals. Atlantic forest endemics were no more or less likely to have small populations than widespread species. There are 79 reserves (>1000 ha) in the Atlantic forest lowlands. However, all but three reserves in the north of the region (Espírito Santo and states north) are smaller than 10,000 ha, and we predict serious levels of local extinction from these reserves. Habitat heterogeneity within reserves may promote species richness within them, but it may also be important in determining species loss over time by suppressing populations of individual species. We suggest that most reserves in the region are so small that homogeneity in the habitat/altitude within them is beneficial for maintenance of their (comparatively small) original species compliment. A lack of protection in the north, continued detrimental human activity inside reserves, and our poor knowledge of how well the reserve system protects individual taxa, are crucial considerations in biodiversity management in the region.     

保护区的大小和数量效果的模拟研究*

对一个既包含局域种群动态,又包含集合种群侵占率的模型进行了计算机模拟,结果表明:(1)集合种群的存活时间随着保护区数目的增大先增大而后减小,即保护区的数目维持在中等大小时最有利于种群在集合种群水平上的存活。(2)这个最优值与保护区的总面积密切相关。这意味着在自然保护区的设置中,如果保护区的总面积一定,那么最佳决策不是保留一个大的完整的自然保护区,而是建立一个由数目相对较多而面积相对较小的保护区组成的网络。这对于自然保护区的设置有着重要的意义。     

Size of nature reserves: densities of large trees and dead wood indicate high value of small conservation forests in southern Sweden

The optimal size of nature reserves has been debated for some time. Although edge and core habitats are often recognized, it is commonly assumed in theory and in studies of a particular habitat type that reserves or patches of different sizes have similar habitat structure. However, for older, highly fragmented landscapes it has been suggested that small areas are of conservation interest as high-quality remnants, whereas large areas are more degraded. We studied 49 randomly selected forest reserves in the size range 5–230 ha (typical for many highly fragmented landscapes) and 3653 sites of key habitat (unprotected deciduous broadleaf forest). Structures in forest that are generally correlated with value for biodiversity were measured, and reserve objectives were examined from declaration texts. Both the density of large trees and the density of dead wood (snags, logs) decreased with increasing reserve size. The mean size of identified key habitats was very small (1.6 ha). A botanical objective for establishment of reserves was more frequently used for smaller reserves. In contrast, cultural and especially recreational objectives were more commonly used when larger reserves were established, suggesting higher values for recreation in these reserves. For vascular plants, birds and beetles, a literature review indicated that small forest patches do not contain impoverished communities, but are often rich (per unit of area). Small reserves and key habitats have several disadvantages, but they are probably important components of reserve networks for biodiversity in highly fragmented landscapes.     

Extinction debt and the role of static and dynamical fragmentation on biodiversity

The mass-extinction events caused by human-driven habitat loss are a current concern in conservation science. However, the observed number of extinctions is considerably smaller than predicted. The overestimation of extinction rates comes from the time-delay which depends on the species sensitivity to habitat changes. The standard method of predicting the effect of habitat loss on biodiversity is to use the species–area relationship and progressively following it backwards to smaller areas. The difference between the actual number of species and the one provided by the backwards species–area relationship is dubbed extinction debt. Previous studies in general adopt a static view for the spatial distribution of species. Nonetheless, a precise understanding of the problem urges us to adopt a dynamic framework to this issue since the time between disturbances of the landscape plays an active role in influencing the strength of the extinction debt. In this context, here we address two distinct approaches for this question: a static and a dynamic view of fragmentation. In the former we quantify the extinction debt in a quenched spatial distribution of species, whereas in the latter the community is let to evolve between disturbance events of the landscape. Here we show that the size of the extinction debt depends on the pattern of the fragmentation. It is found that random distributions of destroyed habitats provide larger extinction debts than those obtained for contiguous areas of fragmentation. Furthermore, in the dynamic approach it is observed that dispersal can lead to unexpected outcomes such as lower biodiversity levels than ones inferred from the backwards species–area relationship.     

Plant-species richness and extinction on isolated dunes along the rocky coast of northwestern Spain

Abstract. Data on distribution and abundance of rare plant species on isolated coastal dunes along the rocky coast of northwestern Spain were compared with the local floristic literature in order to reconstruct past distribution and local extinction of species. Species extinction was not confined to species with a narrow distribution but also included widely distributed ones. Moreover, there was no evidence of colonization. Species number was related to beach and dune area, but species differed in the minimum area required for occurrence. The largest dune reserve possible would probably be insufficient to avoid further extinctions, so also small areas containing particular species should be preserved. There was no significant relationship between abundance of species in the local dune communities and their total distribution, and there were few satellite species. The former might be attributed to low fertility and habitat diversity.     

On optimal size and number of reserves for metapopulation persistence

Habitat fragmentation is generally considered to be detrimental to the persistence of natural populations. In nature management, one therefore tends to prefer few large nature reserves over many small nature reserves having equal total area. This paper examines whether this preference is warranted in a metapopulation framework with circular reserves (patches) by formulating the dependence of metapopulation persistence on the size and number of reserves, both of which depend on reserve radius if the total area is kept constant. Two measures of metapopulation persistence are used: R(0), the number of patches colonized by an occupied patch during its lifetime as an occupied patch, and T(e), the expected time to extinction. These two measures are functions of the extinction and colonization rates of the metapopulation. Several mechanisms for the extinction and colonization processes are formulated from which the dependence of these rates on reserve radius is calculated. It turns out that T(e)generally increases with reserve radius for all mechanisms, which supports the preference of few large reserves. However, R(0)supports this preference only in the case of some special, rather unrealistic, mechanisms. In many other, more realistic, cases an intermediate reserve size exists for which metapopulation persistence measured by R(0)is optimal.     

Logic for designing nature reserves for multiple species

We examine the logic of designing nature reserves to understand better how to integrate the concepts of representativeness and persistence. Simple models of viability are used to evaluate how the expected number of species in the reserve system changes with variation in the risk of extinction among species, their rate of occurrence, and the distribution of species. The optimal size of individual reserves increased with the mean and variance of the probability of extinction among species and with the rate at which the risk of extinction declines with the cost of each reserve. In contrast, the rate of occurrence of species within reserves and their rate of accumulation with increasing reserve area had a relatively minor influence on the optimal size of reserves. Patterns of endemism were most important for the location of reserves. Including differences among species in the analysis reduced the optimal number of individual reserves (and increased the size of each) when operating under a fixed budget compared with reserve designs based on single species. A case study in the city of Melbourne, Australia, demonstrates the conservation value of small (approximately 1 ha) grassland reserves and the underrepresentation of Melbourne's volcanic plains in the region's conservation network.     

Biodiversity of Bitahai Nature Reserve in Yunnan Province, China

Bitahai Nature Reserve is located in Northwest Yunnan and is the highest elevation and the highest latitude wetland nature reserve in Yunnan Province. Bitahai Nature Reserve is a typical wetland nature reserve in very low latitude and high elevation with large areas of coniferous forest around the alpine till lakes, wetland and water area ecosystem while compared with natural reserves distributed same latitude in the world. The area of the reserve distribution is the key area of Transverse Mountain Region. It was a refuge for plant and wildlife because Transverse Mountains were not covered by ice in the Tertiary. And the area is now regarded as a center of forming new species and preserving ancient species. Species biodiversity is high and the distribution of some endemic animals and plants are limited in Transverse Mountains area. In the nature reserve, there are many first and second grade protected plants and animals listed in Lists of China National Priority Protection Fauna and Flora. All of these are precious materials of genetic diversity. The diversities of the plant community and vegetation contribute to ecosystem diversity. Thus, Bitahai Nature Reserve holds with high conservation value. However, its biodiversity is threatened by different factors and its conservation should be paid great attention.     

Would climate change drive species out of reserves? An assessment of existing reserve‐selection methods

Concern for climate change has not yet been integrated in protocols for reserve selection. However if climate changes as projected, there is a possibility that current reserve‐selection methods might provide solutions that are inadequate to ensure species' long‐term persistence within reserves. We assessed, for the first time, the ability of existing reserve‐selection methods to secure species in a climate‐change context. Six methods using a different combination of criteria (representation, suitability and reserve clustering) are compared. The assessment is carried out using European distributions of 1200 plant species and considering two extreme scenarios of response to climate change: no dispersal and universal dispersal. With our data, 6–11% of species modelled would be potentially lost from selected reserves in a 50‐year period. Measured uncertainties varied in 6% being 1–3% attributed to dispersal assumptions and 2–5% to the choice of reserve‐selection method. Suitability approaches to reserve selection performed best, while reserve clustering performed poorly. We also found that 5% of species modelled would lose their entire climatic envelope in the studied area; 2% of the species modelled would have nonoverlapping distributions; 93% of the species modelled would maintain varying levels of overlapping distributions. We conclude there are opportunities to minimize species' extinctions within reserves but new approaches are needed to account for impacts of climate change on species; especially for those projected to have temporally nonoverlapping distributions.     

基于ZONATION的岷山山系多物种保护规划

空间布局不尽合理是我国的自然保护区发展面临的问题之一,优化现有的自然保护区体系是在资源有限的情况下实现自然保护区的保护效益最大化的最佳途径。以岷山地区为主要研究区域,以25种珍稀濒危物种为主要研究对象,通过MAXENT与ZONATION模型模拟,鉴别出岷山地区珍稀濒危物种保护优先区,并提出了自然保护区空间选址的优化方案。研究结果表明,(1)在物种出现点记录较少的情况下,MAXENT模型依然能够比较准确地预测出物种的分布状况;(2)目前建立的保护区对25个物种的平均保护比例为51.8%;(3)通过ZONATION鉴定的优先区总面积为19958.7 km~2,岷山地区现有的29个自然保护区只保护了目标优先区的47.1%,九寨沟中部与南部,平武北部,平武与北川的交界处,北川西北部等地都存在保护空缺。建议在保护空缺的6个地方新建或扩建自然保护区,并在规划时考虑这些区域内部的优先次序。新提出的保护体系能将物种栖息地的保护比例提高至77.9%。研究结果对于岷山及全国其他生物多样性保护关键地区保护体系的规划都具有重要的借鉴意义。     

The importance of species range attributes and reserve configuration for the conservation of angiosperm diversity in Western Australia

In order to better understand the relationship between reserve design and the species represented by such designs, we examined the effectiveness of the Western Australian reserve system for conserving angiosperm diversity, and examined the characteristics of those species conserved. We overlayed species distribution data for 14 plant lineages with the distribution of the reserve system (8.5% of the State’s area) and identified the species that remained unprotected. We found that, depending upon the method employed, between 174 (5.7%) and 570 (18.7%) of species were not included within the reserve system. Two main unprotected regions were identified, one of which was also a centre of high diversity. Geographical range sizes of unprotected species were six times smaller than those species that were protected, while species richness of small-ranged endemic species coincided with general patterns of species richness. At the level of Western Australia’s bioregions we found that conservation effectiveness was most dependent on characteristics of the reserve system rather than characteristics (size and positioning) of species ranges. At this scale, the most effective way to conserve more species in Western Australia would be to conserve more land, while conservation would be most successful in a uniformly dispersed reserve system. Our results highlight the fact that reserve systems may take on two design approaches based on scale––at continental scales, reserves should be clustered around the hotspots of endemic species, while within regions, an evenly distributed reserve system will most adequately sample species.     

Diversity patterns in savanna grassland communities: implications for conservation strategies in a biodiversity hotspot

For any conservation strategy to be effective, it must be preceded by knowledge of how diversity is configured within the area of interest. Here, data from 40 savanna-grassland sites were used to examine how plant biodiversity and species composition varied across spatial scales at the Kalakad-Mundanthurai Tiger Reserve (KMTR), south India. Grasslands surveyed contained 278 plant species, and were characterized by high spatial variability in species association patterns. Fourteen distinct community assemblages were identified, organized primarily along an elevation gradient in the reserve. Overall, grasslands at KMTR were characterized by the dominance of a few, widespread species. The bulk of species richness, however, resulted from subordinate species with fairly restricted distributions. At low elevations, grasslands had high species richness and species composition differed greatly between sites. Mid-elevation grasslands contained about half the number of species present at low elevations, but sites were more similar in species composition. Richness of high-elevation grasslands was a third of that found at low elevations, but different sites harbored unique sets of species. Herbivore use of grasslands varied between communities and showed patterns that coincided with elevation. Herbivore use of low-elevation communities was high albeit variable, of mid-elevation sites was consistently low, and increased at higher elevations. Tall grass communities were the least utilized by herbivores at all elevations. Most species surveyed were rare and restricted in their distribution suggesting that conservation efforts must, perforce, encompass the entire reserve. However, differences in the structuring of diversity across elevations, and in herbivore use of grasslands, suggest that conservation efforts can be partitioned differentially across locations, specifically targeting low and high elevation grasslands in the reserve.     

云南省小黑山自然保护区种子植物区系研究

小黑山自然保护区有野生种子植物170科787属2195种。分析表明该区种子植物区系的地理成分复杂,联系广泛。热带性质的属有506属,占总属数的68.6%,温带性质的属有224属,占总属数的30.4%。热带性质的种890种,占总种数的40.9%,温带性质的种有1267种,占总种数的58.3%。表明该地区植物区系具有明显的亚热带性质,同时深受热带植物区系的影响。植物区系来源主要由东亚成分、热带亚洲成分(印度—马来西亚成分)、中国特有成分三部分融合而成。特有现象明显,有东亚特有科5科,中国特有属6属,中国特有种778种,占种总数的35.8%,而云南特有种有355种,占中国特有种的45.6%。保护区还拥有众多的珍稀濒危植物。     

基于生态系统服务的海南岛自然保护区体系规划

随着我国生物多样性保护与生态建设的发展,区域尺度自然保护区体系规划得到广泛的关注.以海南岛为例,探索了利用生态系统服务空间格局进行区域自然保护区规划的方法.研究中,首先从生物多样性分布、水源涵养、水资源保护、土壤保持和海岸带防护等5个方面分析了海南岛生态系统服务的空间格局,综合评价单元生态系统重要性;然后基于生态系统服务综合评价结果,并根据海南省保护自然过程与生物多样性并重的要求,识别出生态重要性高但无保护区或保护面积不足的空间单元,提出海南岛自然保护区体系规划.根据本规划,海南岛将共建成规范完善的自然保护区42个,总面积290.14万hm2.其中,陆地自然保护区面积41.26万hm2,占海南岛陆地总面积的11.79％.新建保护区4个(面积8.80万hm2),调整保护区4个(扩大面积8.01万公顷),优化整合原有保护区中的17个为6个,升级保护区4个.结果表明,基于生态系统服务的自然保护区体系规划有助于发挥自然保护区的综合效益,并具有更强的可操作性.将生态系统服务纳入自然保护区体系规划应是今后的发展方向.