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.     

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.     

Biased data reduce efficiency and effectiveness of conservation reserve networks

Complementarity-based reserve selection algorithms efficiently prioritize sites for biodiversity conservation, but they are data-intensive and most regions lack accurate distribution maps for the majority of species. We explored implications of basing conservation planning decisions on incomplete and biased data using occurrence records of the plant family Proteaceae in South Africa. Treating this high-quality database as 'complete', we introduced three realistic sampling biases characteristic of biodiversity databases: a detectability sampling bias and two forms of roads sampling bias. We then compared reserve networks constructed using complete, biased, and randomly sampled data. All forms of biased sampling performed worse than both the complete data set and equal-effort random sampling. Biased sampling failed to detect a median of 1-5% of species, and resulted in reserve networks that were 9-17% larger than those designed with complete data. Spatial congruence and the correlation of irreplaceability scores between reserve networks selected with biased and complete data were low. Thus, reserve networks based on biased data require more area to protect fewer species and identify different locations than those selected with randomly sampled or complete data.     

Effects of data characteristics on the results of reserve selection algorithms

We tested the effects of four data characteristics on the results of reserve selection algorithms. The data characteristics were nestedness of features (land types in this case), rarity of features, size variation of sites (potential reserves) and size of data sets (numbers of sites and features). We manipulated data sets to produce three levels, with replication, of each of these data characteristics while holding the other three characteristics constant. We then used an optimizing algorithm and three heuristic algorithms to select sites to solve several reservation problems. We measured efficiency as the number or total area of selected sites, indicating the relative cost of a reserve system. Higher nestedness increased the efficiency of all algorithms (reduced the total cost of new reserves). Higher rarity reduced the efficiency of all algorithms (increased the total cost of new reserves). More variation in site size increased the efficiency of all algorithms expressed in terms of total area of selected sites. We measured the suboptimality of heuristic algorithms as the percentage increase of their results over optimal (minimum possible) results. Suboptimality is a measure of the reliability of heuristics as indicative costing analyses. Higher rarity reduced the suboptimality of heuristics (increased their reliability) and there is some evidence that more size variation did the same for the total area of selected sites. We discuss the implications of these results for the use of reserve selection algorithms as indicative and real-world planning tools.     

Reserve selection and persistence: complementing the existing Atlantic Forest reserve system

This study is an exercise to check the efficiency of the existing reserve system, and to show how systematic conservation planning—using information available and the complementarity concept—can improve the basis for decisions and minimize costs. We verified the performance, in number of cells and primate species representation, of the existing Atlantic Forest (Brazil) reserve network with a quarter-degree resolution grid, with 1,884 cells. We used occurrence data of 20 endemic primate species, and the maps of 237 existing reserves. Reserve networks were selected to represent primate species first considering no pre-existing reserves in Atlantic Forest, and then, considering the existing reserve system, taking into account the minimum area for viable population of the larger species (Northern muriqui Brachyteles hypoxanthus). Reserve selection was carried out using the complementarity concept implemented by a simulated annealing algorithm. Primate species representation (at least one occurrence in the network) could be achieved with 8% of the existing reserve system (nine cells in relation to the 120 in the existing reserve system). We found that today’s reserve system represents 89% of endemic primate species, excluding the species Coimbra Filho’s titi monkey (Callicebus coimbrai) and Marcgraf’s capuchin (Cebus flavius). The networks selected without considering existing reserves contained nine cells. The networks selected considering existing reserves (120 cells), had two new cells necessary to represent all the primates. This does not mean that a viable alternative is to start from zero (i.e., nonexistent reserves). Identifying critical supplementary areas using biodiversity information to fill the gaps and then starting “conservation in practice” in these areas should be priorities.     

Complementary representation and zones of ecological transition

Minimum complementary sets of sites that represent each species at least once have been argued to provide a nominal core reserve network and the starting point for regional conservation programs. However, this approach may be inadequate if there is a tendency to represent several species at marginal areas within their ranges, which may occur if high efficiency results from preferential selection of sites in areas of ecological transition. Here we use data on the distributions of birds in South Africa and Lesotho to explore this idea. We found that for five measures that are expected to reflect the location of areas of ecological transition, complementary sets tend to select higher values of these measures than expected by chance. We recommend that methods for the identification of priority areas for conservation that incorporate viability concerns be preferred to minimum representation sets, even if this results in more costly reserve networks.     

The performance of existing networks of conservation areas in representing biodiversity

It is widely held that existing reserve systems are inadequate in representing the diversity of biological features of the regions in which they reside. Evidence for this argument has, however, derived principally from analyses of the efficiency of networks when compared with a minimum set that represents each species at least once. Here, we examine the efficiency of the system of Sites of Special Scientific Interest (SSSIs) in representing wetland plants in fen sites in the Scottish Borders, a region where reserve networks might be expected a priori to perform reasonably well in this regard. The results support the general contention that networks have been designated in an inefficient manner. However, examined in terms of effectiveness (measured as the gap between the representation target required and the one attained by the existing network), the SSSI system is actually a rather good way of representing diversity. This result is consistent when each of several very different representation targets is evaluated, and suggests that a more balanced approach to evaluating the performance of reserve networks should be employed, and that general statements based on existing analyses should be treated cautiously.     

Capturing biodiversity: selecting priority areas for conservation using different criteria

International treaties call for the protection of biodiversity in all its manifestations, including ecosystem and species diversities. The selection of most priority area networks focuses, however, primarily on species richness and occurrence. The effectiveness of this approach in capturing higher order manifestations of biodiversity, that is ecosystem and environmental diversity patterns, remains poorly understood. Using a case study of birds and environmental data from South Africa and Lesotho, we test how complementary networks that maximise species diversity perform with regard to their representation of ecosystem and environmental diversity, and vice versa. We compare these results to the performance of the existing reserve network. We conclude that focusing on any single biodiversity component alone is insufficient to protect other components. We offer explanations for this in terms of the autocorrelation of species diversity in environmental space.     

Multi-Scales Analysis of Primate Diversity and Protected Areas at a Megadiverse Region

In this paper, we address the question of what proportion of biodiversity is represented within protected areas. We assessed the effectiveness of different protected area types at multiple scales in representing primate biodiversity in the Brazilian Legal Amazon. We used point locality data and distribution data for primate species within 1°, 0.5°, and 0.25° spatial resolution grids, and computed the area of reserves within each cell. Four different approaches were used – no reserves (A), exclusively strict use reserves (B), strict and sustainable use reserves (C), and strict and sustainable use reserves and indigenous lands (D). We used the complementarity concept to select reserve networks. The proportions of cells that were classified as reserves at a grid resolution of 1° were 37%, 64%, and 88% for approaches B, C and D, respectively. Our comparison of these approaches clearly showed the effect of an increase in area on species representation. Representation was consistently higher at coarser resolutions, indicating the effect of grain size. The high number of irreplaceable cells for selected networks identified based on approach A could be attributed to the use of point locality occurrence data. Although the limited number of point occurrences for some species may have been due to a Wallacean shortfall, in some cases it may also be the result of an actual restricted geographic distribution. The existing reserve system cannot be ignored, as it has an established structure, legal protection status, and societal recognition, and undoubtedly represents important elements of biodiversity. However, we found that strict use reserves (which are exclusively dedicated to biodiversity conservation) did not effectively represent primate species. This finding may be related to historical criteria for selecting reserves based on political, economic, or social motives.     

Conservation targets for viable species assemblages?

Estimates of minimum areas required for effective biodiversity conservation differ substantially. Scientific reserve design and placement procedures indicate that between 30 and 75% of any region may be required to sample biodiversity features. These estimates do not routinely incorporate measures for sampling viable populations of species or explore the area requirements of sampling viable populations of species assemblages. To determine the area requirements for sampling viable populations of a herbivore assemblage, spatially explicit abundance data from the Kruger National Park, South Africa, were analyzed. Area requirements were consistently above 50% and were unaffected by selected target population sizes. In addition, area requirements appeared to be insensitive to selection unit size (analytical grain), habitat quality, the coarseness of the land classification system used or the presence of low-density species. Thus, traditional conservation area targets of 10–15% appear inadequate for representing viable populations of a herbivore assemblage from African savanna regions. This suggests that conservation targets of at least 50% of land classification units may represent a more appropriate conservation rule of thumb, or alternatively, that the use of data independent conservation targets may need to be abandoned.     

Priority areas for the conservation of subtropical indigenous forest in southern Africa: a case study from KwaZulu-Natal

Southern Africa's subtropical forest biome, though small and highly fragmented, supports much of the region's biodiversity. With limited resources available for conservation and the exploitative use of forest escalating, identifying a network of priority forest reserves is important. We examine the distribution of forest birds, butterflies and mammals in KwaZulu-Natal, South Africa. Using an iterative algorithm we explore the efficiency of existing protected areas, species richness and rarity hotspots, prime forest sites (selected by forest area) and complementary networks as alternative approaches to priority reserve selection, as well as the potential use of indicator taxa. Existing protected areas represent 98% of species but are relatively inefficient in terms of area. Alternative selection criteria represent a high proportion of species (86–92%) and provide efficient bases for developing fully representative reserve networks. All species are represented within a network of 22 complementary quarter degree cells. This network includes several larger forests and existing protected areas and is recommended for priority conservation. Complementary networks identified separately for birds, butterflies and mammals overlap little, but each represents a high proportion of the remaining taxa, supporting their potential as representative 'indicator' taxa. The evolutionary history of the three main forest types in KwaZulu-Natal explains observed spatial patterns of alternative reserve networks. Priority areas are concentrated in scarp and coastal forest belts, regions of comparatively recent evolutionary activity with high species richness. Afromontane forest is older and less diverse, but its inclusion in any reserve network is necessary for the full representation of forest diversity.     

Scale dependence of plant species richness in a network of protected areas

Despite the widely recognised importance of reserve networks, their effectiveness in encompassing and maintaining biodiversity is still debated. Species diversity is one of the most affordable measures of biodiversity, but it is difficult to survey such data over large scales. This research aimed to perform a sample-based assessment of species richness of groups of plants with different conservation value (alien species, protected species, and all species) within a reserve network, testing the use of partitioning as a tool for assessing diversity at different spatial scales, from the plot to the entire network. Plant diversity patterns differed for the groups of species for most of the investigated spatial scales. Despite these patterns assumed divergent tendencies when different species groups were considered, most of the species richness within the network was given by larger scale β-diversity for both alien and protected species, as well for all species. Diversity partitioning proved an effective tool to quantify the role of spatial scales in structuring the total species richness of the network, and is helpful in planning reserve networks.     

Nature reserve selection in the Transvaal, South Africa: what data should we be using?

Iterative reserve selection algorithms were applied to two mammal databases, generalized to sixteenth degree grid squares, for the Transvaal region of South Africa. Based on primary point data, 24 grid squares are required to represent all species at least once, while only 13 grid squares are required when based on distribution map data; only two of these grid squares are common to both analyses. As the number of representations per species is increased from one to five, the number of selected grid squares increased to 86 and 71 or 72 respectively, with only 17 of these common to both analyses. These differences in the selection of sites are further reflected in the degree of congruence between selected grid squares and existing conservation areas which is on average 63.3% for grid squares selected from the primary database and only 42.5% for those selected from the distribution map database. These results emphasize the importance of quality data input when evaluating regional reserve networks. Highly generalized distribution map data sets, on the one hand, are extrapolations of limited data sets and contain non-quantifiable levels of false-positives which could have significant implications if used for establishing regional reserve networks. On the other hand, although there are problems associated with the establishment of primary diversity databases, namely data currency and uneven and non-random sampling (leading to false negatives), they remain our most reliable option for assigning conservation value.     

Annelids, arthropods or molluscs are suitable as surrogate taxa for selecting conservation reserves in estuaries

The urgent need to conserve aquatic biodiversity and the lack of spatial data on biodiversity has motivated conservation planners and researchers to search for more readily obtainable information that could be used as proxies or surrogates. The surrogate taxon approach shows promise in some aquatic environments (e.g. intertidal) but not others (e.g. coral reefs, temperate rocky reefs). Estuaries are transitional environments at the land–sea junction with a unique biodiversity, but are the most threatened of aquatic environments because of high levels of human use. The comparatively small numbers of conservation reserves means that estuarine biodiversity is poorly protected. Selecting additional conservation reserves within estuaries would be facilitated by the identification of a suitable surrogate that could be used in conservation planning. In one estuary in Southeast Australia, we evaluated separately the effectiveness of annelids, arthropods, and molluscs as surrogates for predicting the species richness, abundance, assemblage variation, and summed irreplaceability of other species and for coincidentally representing other species in networks of conservation reserves selected for each surrogate. Spatial patterns in the species richness and assemblage variation (but not summed irreplaceability) of each surrogate were significantly correlated with the spatial patterns of other species. The total abundance of annelids and the total abundance of arthropods were each significantly correlated with the total abundances of other species. Networks of conservation reserves selected to represent each surrogate performed significantly better than random selection in representing other species. The greatest number of non-surrogate species was coincidentally included in reserves selected for the group of mollusc species. We conclude that annelids and arthropods are effective surrogate taxa for identifying spatial variation in several measures of conservation value (species richness, abundance, assemblage variation) in estuaries. We also conclude that spatial data on annelids, arthropods or molluscs can be used to select networks of conservation reserves in estuaries. The demonstrated effectiveness of these surrogates should facilitate future conservation planning within estuaries.     

Place prioritization for biodiversity reserve network design: a comparison of the SITES and ResNet software packages for coverage and efficiency

Abstract. The place prioritization problem in conservation biology is that of establishing a sequentially prioritized list of places on the basis of biodiversity content. Such a list can then be used to select reserve networks that are designed to be fully representative of the biodiversity of an area as efficiently as possible (for instance, with minimum area or cost). The usual goal is the representation of all chosen biodiversity surrogates up to or beyond a required target, or to the greatest available extent. The purpose of this paper is to compare the respective performances of two place prioritization software packages, SITES and ResNet, on four datasets (distributions of termite genera in Namibia, breeding bird species in the Falkland Islands/Islas Malvinas, vertebrate species in Texas and flora and fauna species that are at risk in Québec), to determine their respective merits. The two software packages implement radically different algorithms: SITES is based on a simulated annealing procedure for finding (local) optima; ResNet uses an algorithm based on rarity and complementarity. This analysis indicates that the rarity‐complementarity based algorithm of ResNet surpasses the simulated annealing approach of SITES with respect to time and completeness. SITES, however, contains other features that are useful in conservation planning. Ways in which the two packages can be used together effectively are suggested.     

Complementarity and other key criteria in the conservation of herb-rich forests in Finland

Complementarity of the nature reserve network in southernmost Finlandwas examined using a simple heuristic algorithm and occurrence data of 75characteristic herb-rich forest vascular plant species in 126 protected and 120non-protected sites. Three different minimum sets were selected to represent 1,5, or 10 occurrences of each species. In each minimum set there weresignificantly more protected than non-protected sites. Thus, although efficiencyis not maximal in the existing reserve network, the network does provide arepresentative basis for the conservation of herb-rich forest plants. However,some deficiencies were also noticed, particularly new reserves in floristicallydiverse herb-rich forests along watercourses would supplement the existingreserve network. On the other hand, the growing concern over the shortcomings ofreserve selection studies using presence/absence data is echoed here, becausemany of the protected forests not included in the minimum sets harbour importantnature conservation values. With regard to the five biological reserve selectioncriteria (e.g. occurrences of threatened species) considered here, theperformance of the existing reserve network is rather good. However, examinationof five reserve design criteria revealed some deficiencies, particularlyvulnerability of many reserves to potential edge effects.     

Priority areas for anuran conservation using biogeographical data: a comparison of greedy, rarity, and simulated annealing algorithms to define reserve networks in cerrado.

Spatial patterns in biodiversity variation at a regional scale are rarely taken into account when a natural reserve is to be established, despite many available methods for determining them. In this paper, we used dimensions of occurrence of 105 species of Anura (Amphibia) in the cerrado region of central Brazil to create a regional system of potential areas that preserves all regional diversity, using three different algorithms to establish reserve networks: "greedy", rarity, and simulated annealing algorithms. These generated networks based on complementarity with 10, 12, and 8 regions, respectively, widely distributed in the biome, and encompassing various Brazilian states. Although the purpose of these algorithms is to find a small number of regions for which all species are represented at least once, the results showed that 67.6%, 76.2%, and 69.5% of the species were represented in two or more regions in the three networks. Simulated annealing produced the smallest network, but it left out three species (one endemic). On the other hand, while the greedy algorithm produce a smaller solution, the rarity-based algorithm ensured that more species were represented more than once, which can be advantageous because it takes into consideration the high levels of habitat loss in the cerrado. Although usually coarse, these macro-scale approaches can provide overall guidelines for conservation and are useful in determining the focus for more local and effective conservation efforts, which is especially important when dealing with a taxonomic group such as anurans, for which quick and drastic population declines have been reported throughout the world.     

Using presence-absence data to establish reserve selection procedures that are robust to temporal species turnover

Previous studies suggest that a network of nature reserves with maximum efficiency (obtained by selecting the minimum area such that each species is represented once) is likely to be insufficient to maintain species in the network over time. Here, we test the performance of three selection strategies which require presence-absence data, two of them previously proposed (multiple representations and selecting an increasing percentage of each species' range) and a novel one based on selecting the site where each species has exhibited a higher permanence rate in the past. Multiple representations appear to be a safer strategy than selecting a percentage of range because the former gives priority to rarer species while the latter favours the most widespread. The most effective strategy was the one based on the permanence rate, indicating that the robustness of reserve networks can be improved by adopting reserve selection procedures that integrate information about the relative value of sites. This strategy was also very efficient, suggesting that the investment made in the monitoring schemes may be compensated for by a lower cost in reserve acquisition.     

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.     

Strengthening the Natural and National Park system of Iberia to conserve vascular plants

The diversity of the Iberian vascular flora has been investigated using WORLDMAP versions 3.08 and 3.18. Two data sets scoring plant distributions as presences within the Iberian Peninsula were compiled; one for 2133 species at 50 × 50 km grid and the other for 801 species at 10 × 10 km map grids. Patterns of biodiversity were determined using the diversity measures of species richness, range-size rarity and character richness diversity. Using the diversity measures, combined with an area selection method, maps of priority areas were calculated using iterative procedures. Near minimum sets (NMSs) for both scales were calculated. Comparison of the NMS for the 10 × 10 km grid with the near minimum set for existing reserves (NMSER) showed that at least 2% more of the land surface would be required above and beyond the existing protected area network, currently comprising 6% of the area, to ensure representation of all species at least once as listed within the present data-base. It is demonstrated that reserve systems selected on a variety of different criteria are suboptimal when compared to particular groups of target organisms with a definite goal of representation for conservation. Calculating efficiency of existing reserve systems and accounting for all taxa identifies precisely the extra required areas for the protected area system to satisfy particular goals of representation.