Integrating larval connectivity into the marine conservation decision-making process across spatial scales

Larval dispersal connectivity is typically integrated into spatial conservation decisions at regional or national scales, but implementing agencies struggle with translating these methods to local scales. We used larval dispersal connectivity at regional (hundreds of kilometers) and local (tens of kilometers) scales to aid in design of networks of no-take reserves in Southeast Sulawesi, Indonesia. We used Marxan with Connectivity informed by biophysical larval dispersal models and remotely sensed coral reef habitat data to design marine reserve networks for 4 commercially important reef species across the region. We complemented regional spatial prioritization with decision trees that combined network-based connectivity metrics and habitat quality to design reserve boundaries locally. Decision trees were used in consensus-based workshops with stakeholders to qualitatively assess site desirability, and Marxan was used to identify areas for subsequent network expansion. Priority areas for protection and expected benefits differed among species, with little overlap in reserve network solutions. Because reef quality varied considerably across reefs, we suggest reef degradation must inform the interpretation of larval dispersal patterns and the conservation benefits achievable from protecting reefs. Our methods can be readily applied by conservation practitioners, in this region and elsewhere, to integrate connectivity data across multiple spatial scales.     

Assessing the current state of ecological connectivity in a large marine protected area system

The establishment of marine protected areas (MPAs) is a critical step in ensuring the continued persistence of marine biodiversity. Although the area protected in MPAs is growing, the movement of individuals (or larvae) among MPAs, termed connectivity, has only recently been included as an objective of many MPAs. As such, assessing connectivity is often neglected or oversimplified in the planning process. For promoting population persistence, it is important to ensure that protected areas in a system are functionally connected through dispersal or adult movement. We devised a multi-species model of larval dispersal for the Australian marine environment to evaluate how much local scale connectivity is protected in MPAs and determine whether the extensive system of MPAs truly functions as a network. We focused on non-migratory species with simplified larval behaviors (i.e., passive larval dispersal) (e.g., no explicit vertical migration) as an illustration. Of all the MPAs analyzed (approximately 2.7 million km²), outside the Great Barrier Reef and Ningaloo Reef, <50% of MPAs (46-80% of total MPA area depending on the species considered) were functionally connected. Our results suggest that Australia's MPA system cannot be referred to as a single network, but rather a collection of numerous smaller networks delineated by natural breaks in the connectivity of reef habitat. Depending on the dispersal capacity of the taxa of interest, there may be between 25 and 47 individual ecological networks distributed across the Australian marine environment. The need to first assess the underlying natural connectivity of a study system prior to implementing new MPAs represents a key research priority for strategically enlarging MPA networks. Our findings highlight the benefits of integrating multi-species connectivity into conservation planning to identify opportunities to better incorporate connectivity into the design of MPA systems and thus to increase their capacity to support long-term, sustainable biodiversity outcomes.     

Defining and Evaluating the Umbrella Species Concept for Conserving and Restoring Landscape Connectivity

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species—a bird (the umbrella), a butterfly, and a frog—inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements. Definición y Evaluación del Concepto de Especie Paraguas para Conservar y Restaurar la Conectividad de Paisajes     

Incorporating connectivity into conservation planning for the optimal representation of multiple species and ecosystem services

Conservation planning tends to focus on protecting species’ ranges or landscape connectivity but seldom both—particularly in the case of diverse taxonomic assemblages and multiple planning goals. Therefore, information on potential trade-offs between maintaining landscape connectivity and achieving other conservation objectives is lacking. We developed an optimization approach to prioritize the maximal protection of species’ ranges, ecosystem types, and forest carbon stocks, while also including habitat connectivity for range-shifting species and dispersal corridors to link protected area. We applied our approach to Sabah, Malaysia, where the state government mandated an increase in protected-area coverage of approximately 305,000 ha but did not specify where new protected areas should be. Compared with a conservation planning approach that did not incorporate the 2 connectivity features, our approach increased the protection of dispersal corridors and elevational connectivity by 13% and 21%, respectively. Coverage of vertebrate and plant species’ ranges and forest types were the same whether connectivity was included or excluded. Our approach protected 2% less forest carbon and 3% less butterfly range than when connectivity features were not included. Hence, the inclusion of connectivity into conservation planning can generate large increases in the protection of landscape connectivity with minimal loss of representation of other conservation targets.     

Effect of Planning for Connectivity on Linear Reserve Networks

Although the concept of connectivity is decades old, it remains poorly understood and defined, and some argue that habitat quality and area should take precedence in conservation planning instead. However, fragmented landscapes are often characterized by linear features that are inherently connected, such as streams and hedgerows. For these, both representation and connectivity targets may be met with little effect on the cost, area, or quality of the reserve network. We assessed how connectivity approaches affect planning outcomes for linear habitat networks by using the stock‐route network of Australia as a case study. With the objective of representing vegetation communities across the network at a minimal cost, we ran scenarios with a range of representation targets (10%, 30%, 50%, and 70%) and used 3 approaches to account for connectivity (boundary length modifier, Euclidean distance, and landscape‐value [LV]). We found that decisions regarding the target and connectivity approach used affected the spatial allocation of reserve systems. At targets ≥50%, networks designed with the Euclidean distance and LV approaches consisted of a greater number of small reserves. Hence, by maximizing both representation and connectivity, these networks compromised on larger contiguous areas. However, targets this high are rarely used in real‐world conservation planning. Approaches for incorporating connectivity into the planning of linear reserve networks that account for both the spatial arrangement of reserves and the characteristics of the intervening matrix highlight important sections that link the landscape and that may otherwise be overlooked. El Efecto de la Planeación para la Conectividad en Redes de Reservas Lineales     

Reserve design to optimize functional connectivity and animal density

Ecological distance-based spatial capture–recapture models (SCR) are a promising approach for simultaneously estimating animal density and connectivity, both of which affect spatial population processes and ultimately species persistence. We explored how SCR models can be integrated into reserve-design frameworks that explicitly acknowledge both the spatial distribution of individuals and their space use resulting from landscape structure. We formulated the design of wildlife reserves as a budget-constrained optimization problem and conducted a simulation to explore 3 different SCR-informed optimization objectives that prioritized different conservation goals by maximizing the number of protected individuals, reserve connectivity, and density-weighted connectivity. We also studied the effect on our 3 objectives of enforcing that the space-use requirements of individuals be met by the reserve for individuals to be considered conserved (referred to as home-range constraints). Maximizing local population density resulted in fragmented reserves that would likely not aid long-term population persistence, and maximizing the connectivity objective yielded reserves that protected the fewest individuals. However, maximizing density-weighted connectivity or preemptively imposing home-range constraints on reserve design yielded reserves of largely spatially compact sets of parcels covering high-density areas in the landscape with high functional connectivity between them. Our results quantify the extent to which reserve design is constrained by individual home-range requirements and highlight that accounting for individual space use in the objective and constraints can help in the design of reserves that balance abundance and connectivity in a biologically relevant manner.     

A network approach to prioritize conservation efforts for migratory birds

Habitat loss can trigger migration network collapse by isolating migratory bird breeding grounds from nonbreeding grounds. Theoretically, habitat loss can have vastly different impacts depending on the site's importance within the migratory corridor. However, migration-network connectivity and the impacts of site loss are not completely understood. We used GPS tracking data on 4 bird species in the Asian flyways to construct migration networks and proposed a framework for assessing network connectivity for migratory species. We used a node-removal process to identify stopover sites with the highest impact on connectivity. In general, migration networks with fewer stopover sites were more vulnerable to habitat loss. Node removal in order from the highest to lowest degree of habitat loss yielded an increase of network resistance similar to random removal. In contrast, resistance increased more rapidly when removing nodes in order from the highest to lowest betweenness value (quantified by the number of shortest paths passing through the specific node). We quantified the risk of migration network collapse and identified crucial sites by first selecting sites with large contributions to network connectivity and then identifying which of those sites were likely to be removed from the network (i.e., sites with habitat loss). Among these crucial sites, 42% were not designated as protected areas. Setting priorities for site protection should account for a site's position in the migration network, rather than only site-specific characteristics. Our framework for assessing migration-network connectivity enables site prioritization for conservation of migratory species.     

Linking life history to landscape for threatened species conservation in a multiuse region

Landscape-scale conservation that considers metapopulation dynamics will be essential for preventing declines of species facing multiple threats to their survival. Toward this end, we developed a novel approach that combines occurrence records, spatial–environmental data, and genetic information to model habitat, connectivity, and patterns of genetic structure and link spatial attributes to underlying ecological mechanisms. Using the threatened northern quoll (Dasyurus hallucatus) as a case study, we applied this approach to address the need for conservation decision-making tools that promote resilient metapopulations of this threatened species in the Pilbara, Western Australia, a multiuse landscape that is a hotspot for biodiversity and mining. Habitat and connectivity were predicted by different landscape characteristics. Whereas habitat suitability was overwhelmingly driven by terrain ruggedness, dispersal was facilitated by proximity to watercourses. Although there is limited evidence for major physical barriers in the Pilbara, areas with high silt and clay content (i.e., alluvial and hardpan plains) showed high resistance to dispersal. Climate subtlety shaped distributions and patterns of genetic turnover, suggesting the potential for local adaptation. By understanding these spatial–environmental associations and linking them to life-history and metapopulation dynamics, we highlight opportunities to provide targeted species management. To support this, we have created habitat, connectivity, and genetic uniqueness maps for conservation decision-making in the region. These tools have the potential to provide a more holistic approach to conservation in multiuse landscapes globally.     

Spatial,socio‐economic,and ecological implications of incorporating minimum size constraints in marine protected area network design

Marine protected areas (MPAs) are the cornerstone of most marine conservation strategies, but the effectiveness of each one partly depends on its size and distance to other MPAs in a network. Despite this, current recommendations on ideal MPA size and spacing vary widely, and data are lacking on how these constraints might influence the overall spatial characteristics, socio‐economic impacts, and connectivity of the resultant MPA networks. To address this problem, we tested the impact of applying different MPA size constraints in English waters. We used the Marxan spatial prioritization software to identify a network of MPAs that met conservation feature targets, whilst minimizing impacts on fisheries; modified the Marxan outputs with the MinPatch software to ensure each MPA met a minimum size; and used existing data on the dispersal distances of a range of species found in English waters to investigate the likely impacts of such spatial constraints on the region's biodiversity. Increasing MPA size had little effect on total network area or the location of priority areas, but as MPA size increased, fishing opportunity cost to stakeholders increased. In addition, as MPA size increased, the number of closely connected sets of MPAs in networks and the average distance between neighboring MPAs decreased, which consequently increased the proportion of the planning region that was isolated from all MPAs. These results suggest networks containing large MPAs would be more viable for the majority of the region's species that have small dispersal distances, but dispersal between MPA sets and spill‐over of individuals into unprotected areas would be reduced. These findings highlight the importance of testing the impact of applying different MPA size constraints because there are clear trade‐offs that result from the interaction of size, number, and distribution of MPAs in a network.     

Validating graph-based connectivity models with independent presence–absence and genetic data sets

Habitat connectivity is a key objective of current conservation policies and is commonly modeled by landscape graphs (i.e., sets of habitat patches [nodes] connected by potential dispersal paths [links]). These graphs are often built based on expert opinion or species distribution models (SDMs) and therefore lack empirical validation from data more closely reflecting functional connectivity. Accordingly, we tested whether landscape graphs reflect how habitat connectivity influences gene flow, which is one of the main ecoevolutionary processes. To that purpose, we modeled the habitat network of a forest bird (plumbeous warbler [Setophaga plumbea]) on Guadeloupe with graphs based on expert opinion, Jacobs’ specialization indices, and an SDM. We used genetic data (712 birds from 27 populations) to compute local genetic indices and pairwise genetic distances. Finally, we assessed the relationships between genetic distances or indices and cost distances or connectivity metrics with maximum-likelihood population-effects distance models and Spearman correlations between metrics. Overall, the landscape graphs reliably reflected the influence of connectivity on population genetic structure; validation R² was up to 0.30 and correlation coefficients were up to 0.71. Yet, the relationship among graph ecological relevance, data requirements, and construction and analysis methods was not straightforward because the graph based on the most complex construction method (species distribution modeling) sometimes had less ecological relevance than the others. Cross-validation methods and sensitivity analyzes allowed us to make the advantages and limitations of each construction method spatially explicit. We confirmed the relevance of landscape graphs for conservation modeling but recommend a case-specific consideration of the cost-effectiveness of their construction methods. We hope the replication of independent validation approaches across species and landscapes will strengthen the ecological relevance of connectivity models.     

Complex effects of habitat amount and fragmentation on functional connectivity and inbreeding in a giant panda population

The relationships between habitat amount and fragmentation level and functional connectivity and inbreeding remain unclear. Thus, we used genetic algorithms to optimize the transformation of habitat area and fragmentation variables into resistance surfaces to predict genetic structure and examined habitat area and fragmentation effects on inbreeding through a moving window and spatial autoregressive modeling approach. We applied these approaches to a wild giant panda population. The amount of habitat and its level of fragmentation had nonlinear effects on functional connectivity (gene flow) and inbreeding. Functional connectivity was highest when approximately 80% of the surrounding landscape was habitat. Although the relationship between habitat amount and inbreeding was also nonlinear, inbreeding increased as habitat increased until about 20% of the local landscape contained habitat, after which inbreeding decreased as habitat increased. Because habitat fragmentation also had nonlinear relationships with functional connectivity and inbreeding, we suggest these important responses cannot be effectively managed by minimizing or maximizing habitat or fragmentation. Our work offers insights for prioritization of protected areas.     

Effects of Weighting Schemes on the Identification of Wildlife Corridors Generated with Least‐Cost Methods

The importance of movement corridors for maintaining connectivity within metapopulations of wild animals is a cornerstone of conservation. One common approach for determining corridor locations is least‐cost corridor (LCC) modeling, which uses algorithms within a geographic information system to search for routes with the lowest cumulative resistance between target locations on a landscape. However, the presentation of multiple LCCs that connect multiple locations generally assumes all corridors contribute equally to connectivity, regardless of the likelihood that animals will use them. Thus, LCCs may overemphasize seldom‐used longer routes and underemphasize more frequently used shorter routes. We hypothesize that, depending on conservation objectives and available biological information, weighting individual corridors on the basis of species‐specific movement, dispersal, or gene flow data may better identify effective corridors. We tested whether locations of key connectivity areas, defined as the highest 75th and 90th percentile cumulative weighted value of approximately 155,000 corridors, shift under different weighting scenarios. In addition, we quantified the amount and location of private land that intersect key connectivity areas under each weighting scheme. Some areas that appeared well connected when analyzed with unweighted corridors exhibited much less connectivity compared with weighting schemes that discount corridors with large effective distances. Furthermore, the amount and location of key connectivity areas that intersected private land varied among weighting schemes. We believe biological assumptions and conservation objectives should be explicitly incorporated to weight corridors when assessing landscape connectivity. These results are highly relevant to conservation planning because on the basis of recent interest by government agencies and nongovernmental organizations in maintaining and enhancing wildlife corridors, connectivity will likely be an important criterion for prioritization of land purchases and swaps. Efectos de los Esquemas de Ponderación sobre la Identificación de Corredores para Vida Silvestre Generados con Métodos Menos Costosos     

Developing Metapopulation Connectivity Criteria from Genetic and Habitat Data to Recover the Endangered Mexican Wolf

Restoring connectivity between fragmented populations is an important tool for alleviating genetic threats to endangered species. Yet recovery plans typically lack quantitative criteria for ensuring such population connectivity. We demonstrate how models that integrate habitat, genetic, and demographic data can be used to develop connectivity criteria for the endangered Mexican wolf (Canis lupus baileyi), which is currently being restored to the wild from a captive population descended from 7 founders. We used population viability analysis that incorporated pedigree data to evaluate the relation between connectivity and persistence for a restored Mexican wolf metapopulation of 3 populations of equal size. Decreasing dispersal rates greatly increased extinction risk for small populations (<150–200), especially as dispersal rates dropped below 0.5 genetically effective migrants per generation. We compared observed migration rates in the Northern Rocky Mountains (NRM) wolf metapopulation to 2 habitat‐based effective distance metrics, least‐cost and resistance distance. We then used effective distance between potential primary core populations in a restored Mexican wolf metapopulation to evaluate potential dispersal rates. Although potential connectivity was lower in the Mexican wolf versus the NRM wolf metapopulation, a connectivity rate of >0.5 genetically effective migrants per generation may be achievable via natural dispersal under current landscape conditions. When sufficient data are available, these methods allow planners to move beyond general aspirational connectivity goals or rules of thumb to develop objective and measurable connectivity criteria that more effectively support species recovery. The shift from simple connectivity rules of thumb to species‐specific analyses parallels the previous shift from general minimum‐viable‐population thresholds to detailed viability modeling in endangered species recovery planning. Desarrollo de Criterios de Conectividad Metapoblacional a Partir de Datos Genéticos y de Hábitat para Recuperar al Lobo Mexicano en Peligro de Extinción     

Effects of land use,cover, and protection on stream and riparian ecosystem services and biodiversity

Protected areas are an important part of broader landscapes that are often used to preserve biodiversity or natural features. Some argue that protected areas may also help ensure provision of ecosystem services. However, the effect of protection on ecosystem services and whether protection affects the provision of ecosystem services is known only for a few services in a few types of landscapes. We sought to fill this gap by investigating the effect of watershed protection status and land use and land cover on biodiversity and the provision of ecosystem services. We compared the ecosystem services provided in and around streams in 4 watershed types: International Union for Conservation of Nature category II protected forests, unprotected forests, unprotected forests with recent timber harvesting, and unprotected areas with agriculture. We surveyed 28 streams distributed across these watershed types in Quebec, Canada, to quantify provisioning of clean water, carbon storage, recreation, wild foods, habitat quality, and terrestrial and aquatic biodiversity richness and abundance. The quantity and quality of ecosystem services and biodiversity were generally higher in sites with intact forest—whether protected or not—relative to those embedded in production landscapes with forestry or agriculture. Clean-water provision, carbon storage, habitat quality, and tree diversity were significantly higher in and around streams surrounded by forest. Recreation, wild foods, and aquatic biodiversity did not vary among watershed types. Although some services can be provided by both protected and unprotected areas, protection status may help secure the continued supply of services sensitive to changes in land use or land cover. Our findings provide needed information about the ecosystem service and biodiversity trade-offs and synergies that result from developing a watershed or from protecting it.     

Minimizing the cost of translocation failure with decision‐tree models that predict species’ behavioral response in translocation sites

The high number of failures is one reason why translocation is often not recommended. Considering how behavior changes during translocations may improve translocation success. To derive decision‐tree models for species’ translocation, we used data on the short‐term responses of an endangered Australian skink in 5 simulated translocations with different release conditions. We used 4 different decision‐tree algorithms (decision tree, decision‐tree parallel, decision stump, and random forest) with 4 different criteria (gain ratio, information gain, gini index, and accuracy) to investigate how environmental and behavioral parameters may affect the success of a translocation. We assumed behavioral changes that increased dispersal away from a release site would reduce translocation success. The trees became more complex when we included all behavioral parameters as attributes, but these trees yielded more detailed information about why and how dispersal occurred. According to these complex trees, there were positive associations between some behavioral parameters, such as fight and dispersal, that showed there was a higher chance, for example, of dispersal among lizards that fought than among those that did not fight. Decision trees based on parameters related to release conditions were easier to understand and could be used by managers to make translocation decisions under different circumstances. Minimizar el Costo del Fracaso de la Reubicación con Modelos de Árboles de Decisión que Predigan la Respuesta Conductual de la Especie en los Sitios de Reubicación     

Protected area planning to conserve biodiversity in an uncertain future

Protected areas are a key instrument for conservation. Despite this, they are vulnerable to risks associated with weak governance, land-use intensification, and climate change. We used a novel hierarchical optimization approach to identify priority areas for expanding the global protected area system that explicitly accounted for such risks while maximizing protection of all known terrestrial vertebrate species. To incorporate risk categories, we built on the minimum set problem, where the objective is to reach species distribution protection targets while accounting for 1 constraint, such as land cost or area. We expanded this approach to include multiple objectives accounting for risk in the problem formulation by treating each risk layer as a separate objective in the problem formulation. Reducing exposure to these risks required expanding the area of the global protected area system by 1.6% while still meeting conservation targets. Incorporating risks from weak governance drove the greatest changes in spatial priorities for protection, and incorporating risks from climate change required the largest increase (2.52%) in global protected area. Conserving wide-ranging species required countries with relatively strong governance to protect more land when they bordered nations with comparatively weak governance. Our results underscore the need for cross-jurisdictional coordination and demonstrate how risk can be efficiently incorporated into conservation planning. Planeación de las áreas protegidas para conservar la biodiversidad en un futuro incierto     

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.     

Conservation planning on China's borders with Myanmar,Laos, and Vietnam

Transboundary conservation is playing an increasingly important role in maintaining ecosystem integrity and halting biodiversity loss caused by anthropogenic activities. However, lack of information on species distributions in transboundary regions and understanding of the threats in these areas impairs conservation. We developed a spatial conservation plan for the transboundary areas between Yunnan province, southwestern China, and neighboring Myanmar, Laos, and Vietnam in the Indo-Burma biodiversity hotspot. To identify priority areas for conservation and restoration, we determined species distribution patterns and recent land-use changes and examined the spatiotemporal dynamics of the connected natural forest, which supports most species. We assessed connectivity with equivalent connected area (ECA), which is the amount of reachable habitat for a species. An ECA incorporates the presence of habitat in a patch and the amount of habitat in other patches within dispersal distance. We analyzed 197,845 locality records from specimen collections and monographs for 21,004 plant and vertebrate species. The region of Yunnan immediately adjacent to the international borders had the highest species richness, with 61% of recorded species and 56% of threatened vertebrates, which suggests high conservation value. Satellite imagery showed the area of natural forest in the border zone declined by 5.2% (13,255 km²) from 1995 to 2018 and monoculture plantations increased 92.4%, shrubland 10.1%, and other cropland 6.2%. The resulting decline in connected natural forest reduced the amount of habitat, especially for forest specialists with limited dispersal abilities. The most severe decline in connectivity was along the Sino-Vietnamese border. Many priority areas straddle international boundaries, indicating demand and potential for establishing transboundary protected areas. Our results illustrate the importance of bi- and multilateral cooperation to protect biodiversity in this region and provide guidance for future conservation planning and practice.     

Dispersal connectivity and reserve selection for marine conservation

Although larval dispersal is crucial for the persistence of most marine populations, dispersal connectivity between sites is rarely considered in designing marine protected area networks. In particular the role of structural characteristics (known as topology) for the network of larval dispersal routes in the conservation of metapopulations has not been addressed. To determine reserve site configurations that provide highest persistence values with respect to their connectivity characteristics, we model nine connectivity topological models derived from graph theory in a demographic metapopulation model. We identify reserve site configurations that provide the highest persistence values for each of the metapopulation connectivity models. Except for the minimally connected and fully connected populations, we observed two general ‘rules of thumb’ for optimising the mean life time for all topological models: firstly place the majority of reserves, so that they are neighbours of each other, on the sites where the number of connections between the populations is highest (hub), secondly when the reserves have occupied the majority of the vertices in the hub, then select another area of high connectivity and repeat. If there are no suitable hubs remaining then distribute the remaining reserves to isolated locations optimising contact with non-reserved sites.     

Accounting for enforcement costs in the spatial allocation of marine zones

Marine fish stocks are in many cases extracted above sustainable levels, but they may be protected through restricted‐use zoning systems. The effectiveness of these systems typically depends on support from coastal fishing communities. High management costs including those of enforcement may, however, deter fishers from supporting marine management. We incorporated enforcement costs into a spatial optimization model that identified how conservation targets can be met while maximizing fishers’ revenue. Our model identified the optimal allocation of the study area among different zones: no‐take, territorial user rights for fisheries (TURFs), or open access. The analysis demonstrated that enforcing no‐take and TURF zones incurs a cost, but results in higher species abundance by preventing poaching and overfishing. We analyzed how different enforcement scenarios affected fishers’ revenue. Fisher revenue was approximately 50% higher when territorial user rights were enforced than when they were not. The model preferentially allocated area to the enforced‐TURF zone over other zones, demonstrating that the financial benefits of enforcement (derived from higher species abundance) exceeded the costs. These findings were robust to increases in enforcement costs but sensitive to changes in species’ market price. We also found that revenue under the existing zoning regime in the study area was 13–30% lower than under an optimal solution. Our results highlight the importance of accounting for both the benefits and costs of enforcement in marine conservation, particularly when incurred by fishers. Justificación de los Costos de Aplicación en la Asignación Espacial de Zonas Marinas