Selecting networks of nature reserves: methods do affect the long-term outcome

Data on vascular plants of boreal lakes in Finland were used to compare the efficiency of reserve selection methods in representing four aspects of biodiversity over a 63 year period. These aspects included species richness, phylogenetic diversity, restricted range diversity and threatened species. Our results show that the efficiency of reserve selection methods depends on the selection criteria used and on the aspect of biodiversity under consideration. Heuristic methods and optimizing algorithms were nearly equally efficient in selecting lake networks over a small geographical range. In addition, a scoring procedure was observed to be efficient in maintaining different aspects of biodiversity over time. However, the random selection of lakes seems to be the most inefficient option for a reserve network. In general, reserve selection methods seem to favour lakes that maximize one aspect of diversity at the time of selection, but the network may not be the best option for maintaining the maximum diversity over time. The reserve selection methods do affect the long-term outcome but it is impossible to recommend one method over the others unequivocally.     

Incorporating spatial criteria in optimum reserve network selection

Considering the spatial location of sites that are to be selected for inclusion in a protected reserve network may be necessary to facilitate dispersal and long-term persistence of species in the selected sites. This paper presents an integer programming (IP) approach to the reserve network selection problem where spatial considerations based on intersite distances are taken into account when selecting reserve sites. The objective is to reduce the fragmentation of preserved sites and design a compact reserve network. Two IP formulations are developed which minimize the sum of pairwise distances and the maximum intersite distance between all sites in the reserve network, respectively, while representing all species under consideration. This approach is applied to a pond invertebrate dataset consisting of 131 sites containing 256 species in Oxfordshire, UK. The results show that significant reductions in reserve fragmentation can be achieved, compared with spatially unrestricted optimum reserve selection, at the expense of a small loss in reserve efficiency.     

Considering Future Potential Regarding Structural Diversity in Selection of Forest Reserves

A rich structural diversity in forests promotes biodiversity. Forests are dynamic and therefore it is crucial to consider future structural potential when selecting reserves, to make robust conservation decisions. We analyzed forests in boreal Sweden based on 17,599 National Forest Inventory (NFI) plots with the main aim to understand how effectiveness of reserves depends on the time dimension in the selection process, specifically by considering future structural diversity. In the study both the economic value and future values of 15 structural variables were simulated during a 100 year period. To get a net present structural value (NPSV), a single value covering both current and future values, we used four discounting alternatives: (1) only considering present values, (2) giving equal importance to values in each of the 100 years within the planning horizon, (3) applying an annual discount rate considering the risk that values could be lost, and (4) only considering the values in year 100. The four alternatives were evaluated in a reserve selection model under budget-constrained and area-constrained selections. When selecting young forests higher structural richness could be reached at a quarter of the cost over almost twice the area in a budget-constrained selection compared to an area-constrained selection. Our results point to the importance of considering future structural diversity in the selection of forest reserves and not as is done currently to base the selection on existing values. Targeting future values increases structural diversity and implies a relatively lower cost. Further, our results show that a re-orientation from old to young forests would imply savings while offering a more extensive reserve network with high structural qualities in the future. However, caution must be raised against a drastic reorientation of the current old-forest strategy since remnants of ancient forests will need to be prioritized due to their role for disturbance-sensitive species.     

Effects of species turnover on reserve site selection in a fragmented landscape

Changes in species composition is an important process in many ecosystems but rarely considered in systematic reserve site selection. To test the influence of temporal variability in species composition on the establishment of a reserve network, we compared network configurations based on species data of small mammals and frogs sampled during two consecutive years in a fragmented Atlantic Forest landscape (SE Brazil). Site selection with simulated annealing was carried out with the datasets of each single year and after merging the datasets of both years. Site selection resulted in remarkably divergent network configurations. Differences are reflected in both the identity of the selected fragments and in the amount of flexibility and irreplaceability in network configuration. Networks selected when data for both years were merged did not include all sites that were irreplaceable in one of the 2 years. Results of species number estimation revealed that significant changes in the composition of the species community occurred. Hence, temporal variability of community composition should be routinely tested and considered in systematic reserve site selection in dynamic systems.     

Selection of a minimum-boundary reserve network using integer programming

In the conservation literature, heuristic procedures have been employed to incorporate spatial considerations in reserve network selection with the presumption that computationally convenient optimization models would be too difficult or impossible to formulate. This paper extends the standard set-covering formulation to incorporate a particular spatial selection criterion, namely reducing the reserve boundary to the extent possible, when selecting a reserve network that represents a set of target species at least once. Applying the model to a dataset on the occurrence of breeding birds in Berkshire, UK, demonstrated that the technique resulted in significant reductions in reserve boundary length relative to solutions produced by the standard set-covering formulation. Computational results showed that moderately large reserve network selection problems could be solved without issue. Alternative solutions may be produced to explore trade-offs between boundary length, number of sites required or alternative criteria.     

The sensitivity of gap analysis to conservation targets

A crucial stage in systematic conservation planning is the definition of explicit conservation targets to be achieved by a network of protected areas. A wide variety of targets have been employed, including overall percentage area, uniform representation of biodiversity features, and variable targets according to conservation interest. Despite the diversity of options, most studies adopt a particular set of targets without further explanation, and few have investigated the effect of target selection on their results. Here, using a data set on the distribution of plants and terrestrial vertebrates in southern France, we investigate how variation in targets can affect both stages of a gap analysis: the assessment of the completeness of an existing reserve network, and the prioritization of areas for its expansion. Target selection had a major impact on the gap analysis results, with uniform targets (50% of each species’ range) emphasizing the representation of common species, and contrasting targets (weighted according to species’ conservation interest) concentrating attention on high conservation interest species and the areas where they occur. Systematic conservation planning exercises should thus pay close attention to the definition and justification of the representation targets employed.     

Reduced survey intensity and its consequences for marine reserve selection

There has been much interest in the potential of short-cuts in biodiversity surveys (e.g. physical surrogates, indicator groups, and lower taxonomic resolution) in systematic processes to select networks of representative marine reserves. This study tested the consequences for reserve selection of reducing survey intensity in intertidal rocky shores in southeast Australia. Using a reference data set of species' distributions based on surveys of two replicate sites in each of 15 locations, a reduction in survey intensity was simulated by randomly eliminating the data from one of the replicate sites in each location. A complementarity-based reserve selection algorithm was used to determine the number of locations required to represent all species once in a reserve network and the irreplaceability value of locations. A reduction in survey intensity led to increases in: the size of reserve networks (of between 8 and 17%); the irreplaceability value of locations; and the number of irreplaceable locations. These changes were caused by a reduction in the observed range sizes of species in the data sets simulating a reduced survey intensity.     

Conservation of Afrotropical antelopes: consequences and efficiency of using different site selection methods and diversity criteria

The consequences of different measures of biotic diversity for the selection of priority sites for conservation were investigated using a dataset on Afrotropical antelopes. Site networks were selected using species richness, taxonomic diversity and restricted-range diversity as selection criteria. Restricted-range diversity was the most efficient criterion at representing all the species in the dataset. However when only a few sites could be conserved (insufficient to include all species) restricted-range diversity was relatively poor at representing absolute numbers of species and also taxonomic diversity. Use of unweighted species richness rather than a taxonomically weighted score did not significantly reduce the amount of taxonomic diversity represented. As expected an iterative selection of sites was considerably more efficient at representing all aspects of diversity than selection of the top-scoring sites. However the efficiency of an iterative selection procedure was reduced when some areas were already part of the reserve network. Since none of the criteria for selecting reserves maximizes all aspects of biodiversity under all circumstances, it is necessary to be clear about the objectives of a reserve network when deciding on a method for site selection.     

Robustness of reserve selection procedures under temporal species turnover

Complementarity-based algorithms for the selection of reserve networks emphasize the need to represent biodiversity features efficiently, but this may not be sufficient to maintain those features in the long term. Here, we use data from the Common Birds Census in Britain as an exemplar data set to determine guidelines for the selection of reserve networks which are more robust to temporal turnover in features. The extinction patterns found over the 1981-1991 interval suggest that two such guidelines are to represent species in the best sites where they occur (higher local abundance) and to give priority to the rarer species. We tested five reserve selection strategies, one which finds the minimum representation set and others which incorporate the first or both guidelines proposed. Strategies were tested in terms of their efficiency (inversely related to the total area selected) and effectiveness (inversely related to the percentage of species lost) using data on eight pairs of ten-year intervals. The minimum set strategy was always the most efficient, but suffered higher species loss than the others, suggesting that there is a trade-off between efficiency and effectiveness. A desirable compromise can be achieved by embedding the concerns about the long-term maintenance of the biodiversity features of interest in the complementarity-based algorithms.     

The cost-effectiveness of using raptor nest sites to identify areas with high species richness of other taxa

Given the limited resources available for conservation, it is important that the areas to preserve are selected in a cost effective manner. However, the cost effectiveness of the surrogate species strategy (the use of information on one or more species to identify areas of value for other species for which there is no, or more limited, available information) has seldom been evaluated.In this study, we investigate the opportunity cost of setting aside breeding sites of two forest raptor species (the surrogate species) by evaluating their individual and combined contribution to preserve diversity of polypores (wood-decaying fungi) and birds against the contributions of previously established nature reserves. We use numeric optimization models to compare different reserve selection strategies.Site selection based on nest sites of the dominant raptor species was more cost-effective than strategies using sites of the subordinate species or those processes previously used to select nature reserves in Finland. The inclusion of both raptor species in the reserve selection model further improved its performance relative to other approaches. This indicates that the means by which Finnish reserves are selected could be enhanced by including the breeding sites of these, and maybe other species, among the criteria used to select reserves in the future.These results show that information on charismatic and well-surveyed species could be a cost-efficient add-on to help enhance conservation endeavours. Where there is inter-specific competition for biodiverse sites, and using multiple species is costly, basing reserve selection primarily on breeding sites of a dominant species may be the best strategy. However, further work is required to establish the extent to which dominant species are typically better indicators of conservation relevance.     

Conservation Targets for Viable Species Assemblages in Canada: Are Percentage Targets Appropriate?

Percentage targets for conservation have become a popular tool (advocated in both the scientific literature and the conservation community) for setting minimum goals for the amount of land to be set aside as protected areas. However, there is little literature to support a consistent percentage target that might be widely applied. Moreover, most percentage targets have not taken into account issues of species persistence. A recent study of herbivores in Kruger National Park took into account issues of representation and persistence in setting conservation targets and found that results were consistently about 50% and were unaffected by different permutations of the reserve selection process. Here, we carry out a similar analysis for representation of mammals within sites that are predicted to allow for their persistence, across eight ecologically defined regions in Canada to test whether we see similar consistent patterns emerging. We found that percentage targets varied with the different permutations of the reserve selection algorithms, both within and between the study regions. Thus, we conclude that the use of percentage targets is not an appropriate conservation strategy.     

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.     

A Test of the Higher-Taxon Approach in the Identification of Candidate Sites for Marine Reserves

Alternatives to species-level identification have been advocated as one solution to the problem of selecting marine reserves with limited information on the distribution of marine biodiversity.This study evaluated the effects on selection of candidate sites for marine reserves from using the higher-taxon approach as a surrogate for species-level identification of intertidal molluscs and rocky reef fishes. These effects were evaluated by determining the percentage of species included in candidate reserves identified from genus-, family- and order-level data by a complementarity-based reserve selection algorithm, and by testing for correlations between the irreplaceability values of locations. Candidate reserves identified from genus- and family-level data of intertidal molluscs included a similar percentage of all species as the reserves identified from species-level data. Candidate reserves selected from genus- and family-level data of rocky reef fishes included, respectively, 3–7% and 14–23% fewer species than reserves selected from species-level data. When the reserve identification process was constrained by a practical planning limit (a maximum of 20% locations able to be reserved) the reserves selected from genus- and family-level data of intertidal molluscs, and genus-level data of rocky reef fishes, included a similar percentage of species as the reserves identified from species-level data. Irreplaceability values of locations for species, genera and families of intertidal molluscs were highly correlated, and irreplaceability values of locations for species and genera of rocky reef fishes were highly correlated. This study suggests that genus- and family-level data for intertidal molluscs, and genus-level data for rocky reef fishes, are suitable surrogates for species in the identification of candidate sites for marine reserves.     

最优化设计连续的自然保护区

生境破碎是导致生物多样性损失的重要原因之一,避免生境破碎的一个有效方式是建立连续的自然保护区使物种可在保护区内自由移动。不加选择地把大片土地都转为保护区是实现连续的一个途径,但资源是有限的,应当以最优的方式分配。如何最优化设计生态上和经济上都有效的保护区成为生物保护领域一个重要议题。从一组备选地块中选择一部分组成自然保护区,这样的问题主要有两种解法:启发式方法和最优化方法。启发式方法虽然灵活且运算速度快但不能保证最优解因而可能导致稀缺资源的浪费,最优化方法保证得到的解是最优的但建模和运算存在困难。建立一个线性整数规划模型用于设计一个最小的连续保护区,用Dantzig剪切法消除循环确保形成一个连续的树,对应一个连续的保护区,检验了模型的计算效率。结果显示,模型可在合理时间内解决一个包含100个备选地块和30个物种的连续保护区设计问题,计算效率显著优于同类目的的其它方法。以美国伊利诺伊州Cache河流域11种濒危鸟类的保护区设计为例说明了该方法的应用,设计了两种情况下连续的保护区。讨论了模型的局限和数据问题。     

Experimental evidence of habitat selection and territoriality in the Amazonian whip spider Heterophrynus longicornis (Arachnida, Amblypygi)

Animals that select and defend suitable habitats against conspecifics may be favored by maximizing prey encounter rate, gaining protection, or securing matings. However, the identification of habitat selection and territoriality may be hindered in observational studies in sedentary species with low-density populations, such as the whip spider Heterophrynus longicornis. To circumvent such difficulties, we adopted an experimental field approach to evaluate if H. longicornis selects and defends habitat in Central Amazon. To evaluate whether individuals perform habitat selection, we monitored the permanence of 29 experimentally released individuals in buttressed trees with a wide variation in diameter at breast height (DBH), with and without burrows at their bases. To evaluate whether individuals are territorial, we experimentally removed 21 individuals from their previously occupied trees and monitored the recolonization of these trees. If H. longicornis were territorial, we predicted that the recolonizers would be smaller than the removed individuals. We found 12 individuals in the trees where they were released on subsequent days, none of them in trees without burrows. The individual permanence was related to the presence of burrows, and not to DBH. There was recolonization by smaller males and females on ten trees from which the amblypygids were removed. In combination, the permanence of individuals in trees with burrows, the rapid recolonization of experimentally vacated trees by smaller individuals, and the preponderance of one individual per tree, suggest that both male and female H. longicornis perform habitat selection and also that after selecting a site, they defend it against conspecifics.     

Prioritizing global marine mammal habitats using density maps in place of range maps

Despite lessons from terrestrial systems, conservation efforts in marine systems continue to focus on identifying priority sites for protection based on high species richness inferred from range maps. Range maps oversimplify spatial variability in animal distributions by assuming uniform distribution within range and de facto giving equal weight to critical and marginal habitats. We used Marxan ver. 2.43 to compare species richness‐based systematic reserve network solutions using information about marine mammal range and relative abundance. At a global scale, reserve network solutions were strongly sensitive to model inputs and assumptions. Solutions based on different input data overlapped by a third at most, with agreement as low as 10% in some cases. At a regional scale, species richness was inversely related to density, such that species richness hotspots excluded highest‐density areas for all species. Based on these findings, we caution that species‐richness estimates derived from range maps and used as input in conservation planning exercises may inadvertently lead to protection of largely marginal habitat.     

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.     

A new method for conservation planning for the persistence of multiple species

Although the aim of conservation planning is the persistence of biodiversity, current methods trade-off ecological realism at a species level in favour of including multiple species and landscape features. For conservation planning to be relevant, the impact of landscape configuration on population processes and the viability of species needs to be considered. We present a novel method for selecting reserve systems that maximize persistence across multiple species, subject to a conservation budget. We use a spatially explicit metapopulation model to estimate extinction risk, a function of the ecology of the species and the amount, quality and configuration of habitat. We compare our new method with more traditional, area-based reserve selection methods, using a ten-species case study, and find that the expected loss of species is reduced 20-fold. Unlike previous methods, we avoid designating arbitrary weightings between reserve size and configuration; rather, our method is based on population processes and is grounded in ecological theory.     

How well do the existing and proposed reserve networks represent vertebrate species in Chile?

Increasingly, biogeographical knowledge and analysis are playing a fundamental role in assessing the representativeness of biodiversity in protected areas, and in identifying critical areas for conservation. With almost 20% of the country assigned to protected areas, Chile is well above the conservation target (i.e. 10–12%) proposed by many international conservation organizations. Moreover, the Chilean government has recently proposed new conservation priority sites to improve the current protected area network. Here, we used all 653 terrestrial vertebrate species present in continental Chile to assess the performance of the existing and proposed reserve networks. Using geographical information systems, we overlaid maps of species distribution, current protected areas, and proposed conservation priority sites to assess how well each species is represented within these networks. Additionally, we performed a systematic reserve selection procedure to identify alternative conservation areas for expanding the current reserve system. Our results show that over 13% of the species are not covered by any existing protected area, and that 73% of Chilean vertebrate species can be considered partial gaps, with only a small fraction of their geographical ranges currently under protection. The coverage is also deficient for endemic (species confined to Chile) and threatened species. While the proposed priority sites do increase coverage, we found that there are still several gaps and these are not the most efficient choices. Both the gap analysis and the reserve selection analysis identified important areas to be added to the existing reserve system, mostly in northern and central Chile. This study underscores the need for a systematic conservation planning approach to redefine the conservation priority sites in order to maximize the representation of species, particularly endemic and threatened species.     

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.