Should We Protect the Strong or the Weak? Risk,Resilience, and the Selection of Marine Protected Areas

Abstract: It is thought that recovery of marine habitats from uncontrollable disturbance may be faster in marine reserves than in unprotected habitats. But which marine habitats should be protected, those areas at greatest risk or those at least risk? We first defined this problem mathematically for 2 alternate conservation objectives. We then analytically solved this problem for both objectives and determined under which conditions each of the different protection strategies was optimal. If the conservation objective was to maximize the chance of having at least 1 healthy site, then the best strategy was protection of the site at lowest risk. On the other hand, if the goal was to maximize the expected number of healthy sites, the optimal strategy was more complex. If protected sites were likely to spend a significant amount of time in a degraded state, then it was best to protect low‐risk sites. Alternatively, if most areas were generally healthy then, counterintuitively, it was best to protect sites at higher risk. We applied these strategies to a situation of cyclone disturbance of coral reefs on Australia's Great Barrier Reef. With regard to the risk of cyclone disturbance, the optimal reef to protect differed dramatically, depending on the expected speed of reef recovery of both protected and unprotected reefs. An adequate consideration of risk is fundamental to all conservation actions and can indicate surprising routes to conservation success.     

Using abiotic variables to predict importance of sites for species representation

In systematic conservation planning, species distribution data for all sites in a planning area are used to prioritize each site in terms of the site's importance toward meeting the goal of species representation. But comprehensive species data are not available in most planning areas and would be expensive to acquire. As a shortcut, ecologists use surrogates, such as occurrences of birds or another well‐surveyed taxon, or land types defined from remotely sensed data, in the hope that sites that represent the surrogates also represent biodiversity. Unfortunately, surrogates have not performed reliably. We propose a new type of surrogate, predicted importance, that can be developed from species data for a q% subset of sites. With species data from this subset of sites, importance can be modeled as a function of abiotic variables available at no charge for all terrestrial areas on Earth. Predicted importance can then be used as a surrogate to prioritize all sites. We tested this surrogate with 8 sets of species data. For each data set, we used a q% subset of sites to model importance as a function of abiotic variables, used the resulting function to predict importance for all sites, and evaluated the number of species in the sites with highest predicted importance. Sites with the highest predicted importance represented species efficiently for all data sets when q = 25% and for 7 of 8 data sets when q = 20%. Predicted importance requires less survey effort than direct selection for species representation and meets representation goals well compared with other surrogates currently in use. This less expensive surrogate may be useful in those areas of the world that need it most, namely tropical regions with the highest biodiversity, greatest biodiversity loss, most severe lack of inventory data, and poorly developed protected area networks.     

The Value of Biodiversity in Reserve Selection: Representation, Species Weighting, and Benefit Functions

Abstract:  The limited availability of resources for conservation has led to the development of many quantitative methods for selecting reserves that aim to maximize the biodiversity value of reserve networks. In published analyses, species are often considered equal, although some are in much greater need of protection than others. Furthermore, representation is usually treated as a threshold: a species is either represented or not, but varying levels of representation over or under a given target level are not valued differently. We propose that a higher representation level should also have higher value. We introduce a framework for reserve selection that includes species weights and benefit functions for under- and overrepresentation (number of locations for each species). We applied the method to conservation planning for herb-rich forests in southern Finland. Our use of benefit functions and weighting changed the identity of about 50% of the selected sites at different funding levels and improved the representation of rare and threatened species. We also identified a small area of additional land that would substantially enhance the existing reserve network. We suggest that benefit functions and species weighting should be considered as standard options in reserve-selection applications.     

The Capacity of Australia's Protected‐Area System to Represent Threatened Species

Abstract: The acquisition or designation of new protected areas is usually based on criteria for representation of different ecosystems or land‐cover classes, and it is unclear how well‐threatened species are conserved within protected‐area networks. Here, we assessed how Australia's terrestrial protected‐area system (89 million ha, 11.6% of the continent) overlaps with the geographic distributions of threatened species and compared this overlap against a model that randomly placed protected areas across the continent and a spatially efficient model that placed protected areas across the continent to maximize threatened species’ representation within the protected‐area estate. We defined the minimum area needed to conserve each species on the basis of the species’ range size. We found that although the current configuration of protected areas met targets for representation of a given percentage of species’ ranges better than a random selection of areas, 166 (12.6%) threatened species occurred entirely outside protected areas and target levels of protection were met for only 259 (19.6%) species. Critically endangered species were among those with the least protection; 12 (21.1%) species occurred entirely outside protected areas. Reptiles and plants were the most poorly represented taxonomic groups, and amphibians the best represented. Spatial prioritization analyses revealed that an efficient protected‐area system of the same size as the current protected‐area system (11.6% of the area of Australia) could meet representation targets for 1272 (93.3%) threatened species. Moreover, the results of these prioritization analyses showed that by protecting 17.8% of Australia, all threatened species could reach target levels of representation, assuming all current protected areas are retained. Although this amount of area theoretically could be protected, existing land uses and the finite resources available for conservation mean land acquisition may not be possible or even effective for the recovery of threatened species. The optimal use of resources must balance acquisition of new protected areas, where processes that threaten native species are mitigated by the change in ownership or on‐ground management jurisdiction, and management of threatened species inside and outside the existing protected‐area system.     

The Effectiveness of Iberian Protected Areas in Conserving Terrestrial Biodiversity

Abstract:  The Iberian Peninsula harbors about 50% of European plant and terrestrial vertebrate species and more than 30% of European endemic species. Despite the global recognition of its importance, the selection of protected areas has been ad hoc and the effectiveness of such choices has rarely been assessed. We compiled the most comprehensive distributional data set of Iberian terrestrial plant and vertebrate species available to date and used it to assess the degree of species representation within existing protected areas. Existing protected areas in Spain and Portugal reasonably represented the plant and animal species we considered (73–98%). Nevertheless, species of some groups (amphibians, reptiles, birds, and gymnosperms) did not accumulate in protected areas at a rate higher than expected by chance ( p > 0.05). We determined that to conserve all vertebrate and plant species in the Iberian Peninsula, at least 36 additional areas are needed. Selection of additional areas for conservation would be facilitated if such areas coincided with sites of community importance (SCI) designated under the European Commission Habitats Directive. Additional areas required for full representation of the selected plant and animal species all coincide with SCI in Spain. Nevertheless, the degree of coincidence varies between 0.3% and 74.6%, and there is a possibility that important areas for conservation occur outside the SCI. Our results support the view that current SCI can be used for prioritization of areas for conservation, but a systematic reevaluation of conservation priorities in Spain and Portugal would be necessary to ensure that effective conservation of one of European's most important biodiversity regions is achieved.     

Terrestrial Reserve Networks Do Not Adequately Represent Aquatic Ecosystems

Abstract: Protected areas are a cornerstone of conservation and have been designed largely around terrestrial features. Freshwater species and ecosystems are highly imperiled, but the effectiveness of existing protected areas in representing freshwater features is poorly known. Using the inland waters of Michigan as a test case, we quantified the coverage of four key freshwater features (wetlands, riparian zones, groundwater recharge, rare species) within conservation lands and compared these with representation of terrestrial features. Wetlands were included within protected areas more often than expected by chance, but riparian zones were underrepresented across all (GAP 1–3) protected lands, particularly for headwater streams and large rivers. Nevertheless, within strictly protected lands (GAP 1–2), riparian zones were highly represented because of the contribution of the national Wild and Scenic Rivers Program. Representation of areas of groundwater recharge was generally proportional to area of the reserve network within watersheds, although a recharge hotspot associated with some of Michigan's most valued rivers is almost entirely unprotected. Species representation in protected areas differed significantly among obligate aquatic, wetland, and terrestrial species, with representation generally highest for terrestrial species and lowest for aquatic species. Our results illustrate the need to further evaluate and address the representation of freshwater features within protected areas and the value of broadening gap analysis and other protected‐areas assessments to include key ecosystem processes that are requisite to long‐term conservation of species and ecosystems. We conclude that terrestrially oriented protected‐area networks provide a weak safety net for aquatic features, which means complementary planning and management for both freshwater and terrestrial conservation targets is needed.     

Incorporating private lands in conservation planning: protected areas in Britain.

Evaluations of the effectiveness of protected areas often report their inadequate representation of regional variation in environmental conditions, land cover, and biological diversity. One frequent contributory explanation is the heavy reliance placed upon the designation of public as opposed to private lands for statutory protection. Given that protected area designation in Britain has no such constraint, and indeed that more than half of such areas are on private lands, we tested the a priori assumption that within this region the representation of environmental conditions and land cover within statutory protected areas would be more equitable. Despite the reduction in land ownership constraints on where protected areas can be established, a marked bias in protected area coverage remains. Protected areas in Britain tend toward regions of higher elevation, soils of lower economic potential, and coastal/estuarine habitat and fail adequately to represent areas of lower elevation and woodland habitats. Improving the current situation requires not only a more systematic approach to site selection, but a more equitable and diverse portfolio of incentives for private landowners to facilitate the decision to manage sites for conservation.     

Conservation priorities for mammals in megadiverse Mexico: the efficiency of reserve networks.

A major goal of conservation biologists is to identify critical areas for the conservation of biological diversity and then strategically include them in an efficient system of reserves. In general, however, reserve networks have been selected for different objectives, and most countries lack an evaluation of their reserves' ability to represent a percentage of the national diversity. This paper evaluates the effectiveness of a network of reserves to represent the species of mammals in Mexico. The focus of the analyses is on species and site level, evaluating the representation of all terrestrial mammals in the 30 most important reserves. The representation of all species, endemic species, endangered species, and species with restricted distributions in the reserves was assessed and compared. Endemic or endangered species with restricted distributions were expected to be less represented in reserves than were widespread species. The most important reserves for the conservation of mammals were determined with the use of complementarity analyses. Priority sites for the representation of all the species currently absent from the reserve network were then selected. The results have broad applications for conservation. First, 82% of the mammal species from Mexico were represented in the reserve network, which covers a small portion (3.8%) of the country. Second, this percentage is certainly larger as several reserves were not evaluated due to a lack of data. A priority for a national conservation strategy could be to conduct biological surveys in those reserves lacking inventories to evaluate their contribution to conservation. Third, in spite of its demonstrated value, Mexico's reserve network can be improved by designating complementary areas. Additional priority sites, where reserves are required to represent most gap species in the network, were identified. Finally, it is clear that this reserve network has limitations for maintaining biodiversity and ecosystem services at regional scales. A comprehensive conservation strategy has, therefore, to incorporate mechanisms that enhance the value of human-dominated landscapes for the maintenance of biodiversity.     

Area-Based Refinement for Selection of Reserve Sites with the Benefit-Function Approach

Abstract:  Optimization of resource use is necessary for efficient conservation planning. Many reserve-selection algorithms aim to identify representative but inexpensive networks, which may lead to selecting small sites due to their lower costs and collectively higher species richness. Nevertheless, larger sites would be preferable regarding species' long-term persistence. An area-based refinement can be used to overcome this problem. We used a reserve-planning framework in which continuous benefit functions valued representation (numbers of populations), and differential species weights were based on a species' local rarity and threatened status. We introduced a refinement based on the species-area relationship that provides relatively higher values for larger sites. We applied the proposed method to rich fen vegetation in southern Finland. The species-area refinement resulted in a network of significantly larger sites with minor trade-offs with representation (numbers of populations). Giving endangered species higher weights ensured that the trade-off occurred mostly between site size and representation of low-priority species. We recommend using a species-area refinement for practical, maximum-coverage conservation planning.     

Spatial Components of Bird Diversity in the Andes of Colombia: Implications for Designing a Regional Reserve System

Abstract:  Beta diversity, or the turnover in species composition among sampling sites in a region, is an important criterion for obtaining adequate representation of regional biodiversity in systems of protected areas. Recently, the additive model for partitioning regional (gamma) diversity (in opposition to the multiplicative model) has been proposed because it allows a direct measure of the contribution of beta diversity to gamma diversity. We determined avian beta diversity along latitudinal (among neighboring river drainages) and elevational axes in a 1347-km² region on the western slope of the Central Cordillera of the Colombian Andes, where a regional system of protected areas is being designed. We then compared avian beta diversity between sites based on rapid versus long-term (>1 year) inventories and between fragmented sites versus continuous forest. Overall, beta diversity represented 63.1% of gamma diversity among 16 sites. Elevational differences in species composition accounted for 43.3% of regional diversity, whereas differences among drainages accounted for 19.8%. A complementary cluster analysis showed that sites grouped by elevational zones. Rapid inventories overestimated beta diversity because of sampling effects, but the effect was biologically small. Estimators of species richness derived from species accumulation curves provided a useful alternative to compensate for undersampling in short-term surveys. Forest fragmentation increased beta diversity because of differential local extinction of populations. Nevertheless, in our region, forest fragments contributed to gamma diversity because they contained complementary sets of species. More importantly, they contained populations of special-interest species. Although the region is relatively small, our analyses indicate that spatial differentiation of the biota is an important factor for deciding number and location of protected areas in the Andean region.     

Objectives for Multiple-Species Conservation Planning

Abstract:  The first step in conservation planning is to identify objectives. Most stated objectives for conservation, such as to maximize biodiversity outcomes, are too vague to be useful within a decision-making framework. One way to clarify the issue is to define objectives in terms of the risk of extinction for multiple species. Although the assessment of extinction risk for single species is common, few researchers have formulated an objective function that combines the extinction risks of multiple species. We sought to translate the broad goal of maximizing the viability of species into explicit objectives for use in a decision-theoretic approach to conservation planning. We formulated several objective functions based on extinction risk across many species and illustrated the differences between these objectives with simple examples. Each objective function was the mathematical representation of an approach to conservation and emphasized different levels of threat. Our objectives included minimizing the joint probability of one or more extinctions, minimizing the expected number of extinctions, and minimizing the increase in risk of extinction from the best-case scenario. With objective functions based on joint probabilities of extinction across species, any correlations in extinction probabilities had to be known or the resultant decisions were potentially misleading. Additive objectives, such as the expected number of extinctions, did not produce the same anomalies. We demonstrated that the choice of objective function is central to the decision-making process because alternative objective functions can lead to a different ranking of management options. Therefore, decision makers need to think carefully in selecting and defining their conservation goals.     

A Protocol for Plant Conservation by Translocation in Threatened Lowland Fynbos

Abstract: A form of lowland, sandplain Fynbos restricted to the Cape Flats near the city of Cape Town is the South African vegetation type most threatened by urban and agricultural development. Cape Flats Fynbos remnants, totaling 4.8 km², contain 14 plant species endemic to the Cape Flats scattered through four protected areas and 11 unprotected vegetation fragments on public land. Despite their small size and management problems, the remaining protected areas are under pressure from the public and developers to function as "lifeboats," or places of safety, for attractive indigenous plant species from the unprotected areas destined for development because the popular perception is that any indigenous plant population threatened by habitat destruction should be saved by being transplanted into a protected area. The urgency for dealing with the issue of plant rescue in the Cape Town metropolitan area was the catalyst for a workshop held at the University of Cape Town in 1998 to develop protocols for Fynbos plant species translocations. We report on points raised by managers and academics at the workshop, including the selection of target taxa, individuals, and establishment sites, and methods for introduction, and present case histories illustrating possible solutions to the problems encountered.     

Using environmental DNA to assess population‐wide spatiotemporal reserve use

Scientists increasingly rely on protected areas to assist in biodiversity conservation, yet the efficacy of these areas is rarely systematically assessed, often because of underfunding. Still, adaptive management strategies to maximize conservation success often rely on understanding the temporal and spatial dynamism of populations therein. Examination of environmental DNA (eDNA) is a time and cost‐effective way to monitor species’ distribution, and quantitative polymerase chain reaction (qPCR) provides information on organismal abundance. To date, however, such techniques remain underused for population assessments in protected areas. We determined eDNA concentration of the critically endangered Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) to describe its occurrence, range, and use of the Tian e‐Zhou National Nature Reserve in Hubei, China, across seasons and hydrological depths. Despite the observation that total eDNA concentrations were highest in surface waters in summer, finless porpoise eDNA concentrations were significantly higher in deeper waters than in surface waters in summer. During the breeding season (spring), eDNA signals were site specific and restricted to the core area of the reserve. However, postbreeding eDNA concentrations were widespread across the reserve, encompassing sites previously thought to be unfrequented by the species. Our results suggest spatiotemporal idiosyncrasies in site, depth, and seasonal use of the reserve and a propensity for postbreeding population dispersal. With eDNA and qPCR we were able to assess an entire population's use of a protected area. Illuminating nuances in habitat use via eDNA could be valuable to set pragmatic conservation goals for this, and other, species.     

Two contrasting effects of predation on species richness in coral reef habitats

The species richness of sessile organisms on settlement panels on a coral reef was measured by the slope of a regression of log_e number of species against log_e area of sample. At a well illuminated site where panels were colonised by algae, the species richness of algae was 19% smaller on surfaces grazed by fishes than on protected surfaces. At a second site in a cave, the species richness of animals on grazed surfaces was 20% greater than on protected surfaces. These results are discussed in the light of differences between the sites. The contrasting effects of predation at the two sites are probably the result of more selective predation at the cave site than at the other site.     

Multispecies genetic objectives in spatial conservation planning

Growing threats to biodiversity and global alteration of habitats and species distributions make it increasingly necessary to consider evolutionary patterns in conservation decision making. Yet, there is no clear‐cut guidance on how genetic features can be incorporated into conservation‐planning processes, despite multiple molecular markers and several genetic metrics for each marker type to choose from. Genetic patterns differ between species, but the potential tradeoffs among genetic objectives for multiple species in conservation planning are currently understudied. We compared spatial conservation prioritizations derived from 2 metrics of genetic diversity (nucleotide and haplotype diversity) and 2 metrics of genetic isolation (private haplotypes and local genetic differentiation) in mitochondrial DNA of 5 marine species. We compared outcomes of conservation plans based only on habitat representation with plans based on genetic data and habitat representation. Fewer priority areas were selected for conservation plans based solely on habitat representation than on plans that included habitat and genetic data. All 4 genetic metrics selected approximately similar conservation‐priority areas, which is likely a result of prioritizing genetic patterns across a genetically diverse array of species. Largely, our results suggest that multispecies genetic conservation objectives are vital to creating protected‐area networks that appropriately preserve community‐level evolutionary patterns.     

Protected species use of a coastal marine migratory corridor connecting marine protected areas

The establishment of protected corridors linking the breeding and foraging grounds of many migratory species remains deficient, particularly in the world’s oceans. For example, Australia has recently established a network of Commonwealth Marine Reserves, supplementing existing State reserves, to protect a wide range of resident and migratory marine species; however, the routes used by mobile species to access these sites are often unknown. The flatback marine turtle (Natator depressus) is endemic to the continental shelf of Australia, yet information is not available about how this species uses the marine area. We used a geospatial approach to delineate a coastal corridor from 73 adult female flatback postnesting migratory tracks from four rookeries along the north-west coast of Australia. A core corridor of 1,150 km length and 30,800 km² area was defined, of which 52 % fell within 11 reserves, leaving 48 % (of equivalent size to several Commonwealth Reserves) of the corridor outside of the reserve network. Despite limited data being available for other marine wildlife in this region, humpback whale migratory tracks overlapped with 96 % of the core corridor, while the tracks of three other species overlapped by 5–10 % (blue whales, olive ridley turtles, whale sharks). The overlap in the distribution ranges of at least 20 other marine vertebrates (dugong, cetaceans, marine turtles, sea snakes, crocodiles, sharks) with the corridor also imply potential use. In conclusion, this study provides valuable information towards proposing new locations requiring protection, as well as identifying high-priority network linkages between existing marine protected areas.     

Software for prioritizing conservation actions based on probabilistic information

Marxan is the most common decision-support tool used to inform the design of protected-area systems. The original version of Marxan does not consider risk and uncertainty associated with threatening processes affecting protected areas, including uncertainty about the location and condition of species’ populations and habitats now and in the future. We described and examined the functionality of a modified version of Marxan, Marxan with Probability. This software explicitly considers 4 types of uncertainty: probability that a feature exists in a particular place (estimated based on species distribution models or spatially explicit population models); probability that features in a site will be lost in the future due to a threatening process, such as climate change, natural catastrophes, and uncontrolled human interventions; probability that a feature will exist in the future due to natural successional processes, such as a fire or flood; and probability the feature exists but has been degraded by threatening processes, such as overfishing or pollution, and thus cannot contribute to conservation goals. We summarized the results of 5 studies that illustrate how each type of uncertainty can be used to inform protected area design. If there were uncertainty in species or habitat distribution, users could maximize the chance that these features were represented by including uncertainty using Marxan with Probability. Similarly, if threatening processes were considered, users minimized the chance that species or habitats were lost or degraded by using Marxan with Probability. Marxan with Probability opens up substantial new avenues for systematic conservation planning research and application by agencies.     

Conservation Planning when Costs Are Uncertain

Abstract: Spatially explicit information on the financial costs of conservation actions can improve the ability of conservation planning to achieve ecological and economic objectives, but the magnitude of this improvement may depend on the accuracy of the cost estimates. Data on costs of conservation actions are inherently uncertain. For example, the cost of purchasing a property for addition to a protected‐area network depends on the individual landholder's preferences, values, and aspirations, all of which vary in space and time, and the effect of this uncertainty on the conservation priority of a site is relatively untested. We investigated the sensitivity of the conservation priority of sites to uncertainty in cost estimates. We explored scenarios for expanding (four‐fold) the protected‐area network in Queensland, Australia to represent a range of vegetation types, species, and abiotic environments, while minimizing the cost of purchasing new properties. We estimated property costs for 17, 790 10 × 10 km sites with data on unimproved land values. We systematically changed property costs and noted how these changes affected conservation priority of a site. The sensitivity of the priority of a site to changes in cost data was largely dependent on a site's importance for meeting conservation targets. Sites that were essential or unimportant for meeting targets maintained high or low priorities, respectively, regardless of cost estimates. Sites of intermediate conservation priority were sensitive to property costs and represented the best option for efficiency gains, especially if they could be purchased at a lower price than anticipated. Thus, uncertainty in cost estimates did not impede the use of cost data in conservation planning, and information on the sensitivity of the conservation priority of a site to estimates of the price of land can be used to inform strategic conservation planning before the actual price of the land is known.     

Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.     

Choice set formation for outdoor destinations: The role of motivations and preference discrimination in site selection for the management of public expenditures on protected areas

Effective public expenditure currently dominates the management focus of many protected areas. This calls for explicit modeling of constraints and motivations that, respectively, obstruct and stimulate visits to selected outdoor destinations. Choice set formation is the result of screening and/or inclusion of specific sites (alternatives) to form the set of sites considered in real choices. Evidence shows that the omission of a structural representation of choice set formation is harmful to econometric inference. Yet, the literature has largely ignored the underlying behavioral phenomenon. We show, using a discrete choice experiment involving selection among seven recreational sites in an Italian national park, that choice set formation is behaviorally relevant, even after controlling for preference discrimination. Motivations (why visit?) are important determinants of preliminary site screening for choice set inclusion, as well as site selection, justifying the additional value of such modeling extension.