Conservation value of small reserves

The importance of large reserves has been long maintained in the scientific literature, often leading to dismissal of the conservation potential of small reserves. However, over half the global protected-area inventory is composed of protected areas that are <100 ha, and the median size of added protected area is decreasing. Studies of the conservation value of small reserves and fragments of natural area are relatively uncommon in the literature. We reviewed SCOPUS and WOK for studies on small reserve and fragment contributions to biodiversity conservation and ecosystem services, and fifty-eight taxon-specific studies were included in the review. Small reserves harbored substantial portions (upward of 50%) of regional species diversity for many taxa (birds, plants, amphibians, and small mammals) and even some endemic, specialist bird species. Unfortunately, small reserves and fragments almost always harbored more generalist and exotic species than large reserves. Community composition depended on habitat quality, surrounding land use (agricultural vs. urban), and reserve and fragment size, which presents opportunities for management and improvement. Small reserves also provided ecosystem services, such as pollination and biological pest control, and cultural services, such as recreation and improved human health. Limitations associated with small reserves, such as extinction debt and support of area-sensitive species, necessitate a complement of larger reserves. However, we argue that small reserves can make viable and significant contributions to conservation goals directly as habitat and indirectly by increasing landscape connectivity and quality to the benefit of large reserves. To effectively conserve biodiversity for future generations in landscapes fragmented by human development, small reserves and fragments must be included in conservation planning.     

Conservation of the Marbled Murrelet under the Northwest Forest Plan

Abstract:  The Marbled Murrelet ( Brachyramphus marmoratus ) was listed as threatened in 1992, primarily because of loss of its old-forest nesting habitat. Monitoring conducted over the first 10 years following implementation of the Northwest Forest Plan shows at-sea murrelet populations appear to be stationary, but recruitment is very low and demographic models project a 4–6% annual rate of decline. Monitoring of nesting habitat indicated there were about 1.6 million ha of higher-suitability nesting habitat on all lands at the start of the plan, about half of which occurred on federal lands. Most (88%) of higher-suitability habitat on federal lands was protected within reserves. Over the past 10 years, losses of habitat due primarily to fire have totaled about 2% on federal lands. Losses have been much greater (12%) on nonfederal lands, due primarily to timber harvest. Habitat is expected to accrue within reserves as younger forest matures and attains sufficient diameter to support nesting sites. At-sea estimates of population size are strongly and positively correlated with amounts of adjacent nesting habitat at a broad scale, supporting the idea that amounts of nesting habitat are a primary driver in wide-scale murrelet population distribution. Conditions at sea, however, such as temperature regimes, prey availability, and pollutants, continue to affect murrelet populations. The system of large reserves seems to have achieved the short-term objective of conserving much of the remaining nesting habitat on federal lands. These reserves are also likely to contribute to the long-term objective of creating large, contiguous blocks of nesting habitat. The plan has a primary role in conserving and restoring nesting habitat on federal land but will succeed in this role only if land allocations calling for such protection are in place for many decades.     

A Method for Setting the Size of Plant Conservation Target Areas

Abstract: Realistic time frames in which management decisions are made often preclude the completion of the detailed analyses necessary for conservation planning. Under these circumstances, efficient alternatives may assist in approximating the results of more thorough studies that require extensive resources and time. We outline a set of concepts and formulas that may be used in lieu of detailed population viability analyses and habitat modeling exercises to estimate the protected areas required to provide desirable conservation outcomes for a suite of threatened plant species. We used expert judgment of parameters and assessment of a population size that results in a specified quasiextinction risk based on simple dynamic models. The area required to support a population of this size is adjusted to take into account deterministic and stochastic human influences, including small-scale disturbance, deterministic trends such as habitat loss, and changes in population density through processes such as predation and competition. We set targets for different disturbance regimes and geographic regions. We applied our methods to Banksia cuneata, Boronia keysii, and Parsonsia dorrigoensis, resulting in target areas for conservation of 1102, 733, and 1084 ha, respectively. These results provide guidance on target areas and priorities for conservation strategies.     

Designing the Masoala National Park in Madagascar Based on Biological and Socioeconomic Data

Abstract: Conservation biologists have developed powerful tools for reserve selection and design over the past two decades, yet seldom are protected areas actually designed on scientific grounds. Using fundamental biological and socioeconomic principles of conservation science, we designed a new protected area and its multiple-use zone on the Masoala Peninsula in the humid forest zone of Madagascar. The explicit design criteria determined the data gathered for the work, which included (1) spatial distribution and quality of habitat, (2) the areas and species at greatest risk, (3) the relationship between environmental gradients and species distributions, (4) current and predicted human settlement and land and resource use, and (5) the economic potential of natural forest management as an alternative to deforestation. We used a geographic information system to integrate these data layers and applied the design criteria to develop a park proposal that balanced human and wildlife needs. The proposal won the approval of local residents, and a national decree in 1997 designated 2100 km² of rainforest and three satellite marine reserves as the Masoala National Park, with a surrounding multiple-use zone of approximately 1000 km². The new park is Madagascar's largest protected area and protects more lowland (<400 m) humid forest habitat than the entire reserve system combined, a significant step forward in conserving a globally important ecoregion. Consideration of local needs and the national economy was a key element in gaining approval for the Masoala Park. Such an approach toward reserve design could be applied elsewhere to improve chances of establishing and maintaining protected areas over the long term.     

Effectiveness of management zoning designed for flagship species in protecting sympatric species

Flagship species have been used widely as umbrella species (i.e., species with large home range whose protection often provides protection for sympatric species) in the management of China's nature reserves. This conflation of flagship and umbrella species is best represented by the giant panda (Ailuropoda melanoleuca) and other large, endangered mammals designated as conservation targets in site selection and planning of reserves. Few empirical studies have tested the effectiveness of flagship species as surrogates for a broader range of sympatric species. Using extensive camera-trap data, we examined the effectiveness of management zones designated to protect flagship (target) species in conserving sympatric species in 4 wildlife reserves (Gutianshan, Changqing, Laohegou, and Wolong). We tested whether the progression from peripheral to core zones was associated with an increasing habitat association for both target and sympatric species. The distribution patterns of the study species across the zones in each reserve indicated a disparity between management zones and the species’ habitat requirements. Management zone was included in the final model for all target species, and most of them had higher occurrence in core zones relative to less-protected zones, but zone was not a predictor for most of the sympatric species. When management zone was associated with the occurrence of sympatric species, threatened species generally had higher detections in core zones, whereas common species had higher detections outside of the core zone. Our results suggested that reserve planning based on flagship species does not adequately protect sympatric species due to their specialized habitat requirements. We recommend re-examining the effectiveness of management zoning and urge a multispecies and reserve-wide monitoring plan to improve protection of China's wildlife.     

Improving protected area effectiveness through consideration of different human-pressure baselines

Previous assessments of the effectiveness of protected areas (PAs) focused primarily on changes in human pressure over time and did not consider the different human-pressure baselines of PAs, thereby potentially over- or underestimating PA effectiveness. We developed a framework that considers both human-pressure baseline and change in human pressure over time and assessed the effectiveness of 338 PAs in China from 2010 to 2020. The initial state of human pressure on PAs was taken as the baseline, and changes in human pressure index (HPI) were further analyzed under different baselines. We used the random forest models to identify the management measures that most improved effectiveness in resisting human pressure for the PAs with different baselines. Finally, the relationships between the changes in the HPI and the changes in natural ecosystems in PAs were analyzed with different baselines. Of PAs with low HPI baselines, medium HPI baselines, and high HPI baselines, 76.92% (n=150), 11.11% (n=12), and 22.86% (n=8) , respectively, showed positive effects in resisting human pressure. Overall, ignoring human-pressure baselines somewhat underestimated the positive effects of PAs, especially for those with low initial human pressure. For PAs with different initial human pressures, different management measures should be taken to improve effectiveness and reduce threats to natural ecosystems. We believe our framework is useful for assessing the effectiveness of PAs globally, and we recommend it be included in the Convention on Biological Diversity Post-2020 Strategy.     

Effects of anthropogenic landscapes on population maintenance of waterbirds

Anthropogenic impacts have reduced natural areas but increased the area of anthropogenic landscapes. There is debate about whether anthropogenic landscapes (e.g., farmlands, orchards, and fish ponds) provide alternatives to natural habitat and under what circumstances. We considered whether anthropogenic landscapes can mitigate population declines for waterbirds. We collected data on population trends and biological traits of 1203 populations of 579 species across the planet. Using Bayesian generalized linear mixed models, we tested whether the ability of a species to use an anthropogenic landscape can predict population trends of waterbird globally and of species of conservation concern. Anthropogenic landscapes benefited population maintenance of common but not less-common species. Conversely, the use of anthropogenic landscapes was associated with population declines for threatened species. Our findings delineate some limitations to the ability of anthropogenic landscapes to mitigate population declines, suggesting that the maintenance of global waterbird populations depends on protecting remaining natural areas and improving the habitat quality in anthropogenic landscapes. Article impact statement: Protecting natural areas and improving the quality of anthropogenic landscapes as habitat are both needed to achieve effective conservation.     

Critical factors for the recovery of marine mammals

{en} Over the past decades, much research has focused on understanding the critical factors for marine extinctions with the aim of preventing further species losses in the oceans. Although conservation and management strategies are enabling several species and populations to recover, others remain at low abundance levels or continue to decline. To understand these discrepancies, we used a published database on abundance trends of 137 populations of marine mammals worldwide and compiled data on 28 potentially critical factors for recovery. We then applied random forests and additive mixed models to determine which intrinsic and extrinsic factors are critical for the recovery of marine mammals. A mix of life‐history characteristics, ecological traits, phylogenetic relatedness, population size, geographic range, human impacts, and management efforts explained why populations recovered or not. Consistently, species with lower age at maturity and intermediate habitat area were more likely to recover, which is consistent with life‐history and ecological theory. Body size, trophic level, social interactions, dominant habitat, ocean basin, and habitat disturbance also explained some differences in recovery patterns. Overall, a variety of intrinsic and extrinsic factors were important for species’ recovery, pointing to cumulative effects. Our results provide insight for improving conservation and management strategies to enhance recoveries in the future.     

A counterfactual approach to measure the impact of wet grassland conservation on U.K. breeding bird populations

Wet grassland populations of wading birds in the United Kingdom have declined severely since 1990. To help mitigate these declines, the Royal Society for the Protection of Birds has restored and managed lowland wet grassland nature reserves to benefit these and other species. However, the impact of these reserves on bird population trends has not been evaluated experimentally due to a lack of control populations. We compared population trends from 1994 to 2018 among 5 bird species of conservation concern that breed on these nature reserves with counterfactual trends created from matched breeding bird survey observations. We compared reserve trends with 3 different counterfactuals based on different scenarios of how reserve populations could have developed in the absence of conservation. Effects of conservation interventions were positive for all 4 targeted wading bird species: Lapwing (Vanellus vanellus), Redshank (Tringa totanus), Curlew (Numenius arquata), and Snipe (Gallinago gallinago). There was no positive effect of conservation interventions on reserves for the passerine, Yellow Wagtail (Motacilla flava). Our approach using monitoring data to produce valid counterfactual controls is a broadly applicable method allowing large-scale evaluation of conservation impact.     

Temporal changes in giant panda habitat connectivity across boundaries of Wolong Nature Reserve, China.

Global biodiversity loss is largely driven by human activities such as the conversion of natural to human-dominated landscapes. A popular approach to mitigating land cover change is the designation of protected areas (e.g., nature reserves). Nature reserves are traditionally perceived as strongholds of biodiversity conservation. However, many reserves are affected by land cover changes not only within their boundaries, but also in their surrounding areas. This study analyzed the changes in habitat for the giant panda (Ailuropoda melanoleuca) inside Wolong Nature Reserve, Sichuan, China, and in a 3-km buffer area outside its boundaries, through a time series of classified satellite imagery and field observations. Habitat connectivity between the inside and the outside of the reserve diminished between 1965 and 2001 because panda habitat was steadily lost both inside and outside the reserve. However, habitat connectivity slightly increased between 1997 and 2001 due to the stabilization of some panda habitat inside and outside the reserve. This stabilization most likely occurred as a response to changes in socioeconomic activities (e.g., shifts from agricultural to nonagricultural economies). Recently implemented government policies could further mitigate the impacts of land cover change on panda habitat. The results suggest that Wolong Nature Reserve, and perhaps other nature reserves in other parts of the world, cannot be managed as an isolated entity because habitat connectivity declines with land cover changes outside the reserve even if the area inside the reserve is well protected. The findings and approaches presented in this paper may also have important implications for the management of other nature reserves across the world.     

Global protected-area coverage and human pressure on tidal flats

Tidal flats are a globally distributed coastal ecosystem important for supporting biodiversity and ecosystem services. Local to continental-scale studies have documented rapid loss of tidal habitat driven by human impacts, but assessments of progress in their conservation are lacking. With an internally consistent estimate of distribution and change, based on Landsat satellite imagery, now available for the world's tidal flats, we examined tidal flat representation in protected areas (PAs) and human pressure on tidal flats. We determined tidal flat representation and its net change in PAs by spatially overlaying tidal flat maps with the World Database of Protected Areas. Similarly, we overlaid the most recent distribution map of tidal flats (2014–2016) with the human modification map (HM_c) (range from 0, no human pressure, to 1, very high human pressure) to estimate the human pressure exerted on this ecosystem. Sixty-eight percent of the current extent of tidal flats is subject to moderate to very high human pressure (HM_c > 0.1), but 31% of tidal flat extent occurred in PAs, far exceeding PA coverage of the marine (6%) and terrestrial (13%) realms. Net change of tidal flat extent inside PAs was similar to tidal flat net change outside PAs from 1999 to 2016. Substantial shortfalls in protection of tidal flats occurred across Asia, where large intertidal extents coincided with high to very high human pressure (HM_c > 0.4–1.0) and net tidal flat losses up to 86.4 km² (95% CI 83.9–89.0) occurred inside individual PAs in the study period. Taken together, our results show substantial progress in PA designation for tidal flats globally, but that PA status alone does not prevent all habitat loss. Safeguarding the world's tidal flats will thus require deeper understanding of the factors that govern their dynamics and effective policy that promotes holistic coastal and catchment management strategies.     

The consequences of poaching and anthropogenic change for forest elephants

Poaching has devastated forest elephant populations (Loxodonta cyclotis), and their habitat is dramatically changing. The long‐term effects of poaching and other anthropogenic threats have been well studied in savannah elephants (Loxodonta africana), but the impacts of these changes for Central Africa's forest elephants have not been discussed. We examined potential repercussions of these threats and the related consequences for forest elephants in Central Africa by summarizing the lessons learned from savannah elephants and small forest elephant populations in West Africa. Forest elephant social organization is less known than the social organization of savannah elephants, but the close evolutionary history of these species suggests that they will respond to anthropogenic threats in broadly similar ways. The loss of older, experienced individuals in an elephant population disrupts ecological, social, and population parameters. Severe reduction of elephant abundance within Central Africa's forests can alter plant communities and ecosystem functions. Poaching, habitat alterations, and human population increase are probably compressing forest elephants into protected areas and increasing human–elephant conflict, which negatively affects their conservation. We encourage conservationists to look beyond documenting forest elephant population decline and address the causes of these declines when developing conversation strategies. We suggest assessing the effectiveness of the existing protected‐area networks for landscape connectivity in light of current industrial and infrastructure development. Longitudinal assessments of the effects of landscape changes on forest elephant sociality and behavior are also needed. Finally, lessons learned from West African elephant population loss and habitat fragmentation should be used to inform strategies for land‐use planning and managing human–elephant interactions.     

Bias in protected‐area location and its effects on long‐term aspirations of biodiversity conventions

To contribute to the aspirations of recent international biodiversity conventions, protected areas (PAs) must be strategically located and not simply established on economically marginal lands as they have in the past. With refined international commitments under the Convention on Biological Diversity to target protected areas in places of “importance to biodiversity,” perhaps they may now be. We analyzed location biases in PAs globally over historic (pre‐2004) and recent periods. Specifically, we examined whether the location of protected areas are more closely associated with high concentrations of threatened vertebrate species or with areas of low agricultural opportunity costs. We found that both old and new protected areas did not target places with high concentrations of threatened vertebrate species. Instead, they appeared to be established in locations that minimize conflict with agriculturally suitable lands. This entrenchment of past trends has substantial implications for the contributions these protected areas are making to international commitments to conserve biodiversity. If protected‐area growth from 2004 to 2014 had strategically targeted unrepresented threatened vertebrates, >30 times more species (3086 or 2553 potential vs. 85 actual new species represented) would have been protected for the same area or the same cost as the actual expansion. With the land available for conservation declining, nations must urgently focus new protection on places that provide for the conservation outcomes outlined in international treaties.     

Effects of landscape design of forest reserves on Saproxylic beetle diversity

Increasing the density of natural reserves in the forest landscape may provide conservation benefits for biodiversity within and beyond reserve borders. We used 2 French data sets on saproxylic beetles and landscape cover of forest reserves (LCFR) to test this hypothesis: national standardized data derived from 252 assessment plots in managed and reserve stands in 9 lowland and 5 highland forests and data from the lowland Rambouillet forest, a forested landscape where a pioneer conservation policy led to creation of a dense network of reserves. Abundance of rare and common saproxylic species and total saproxylic species richness were higher in forest reserves than in adjacent managed stands only in highland forests. In the lowland regional case study, as LCFR increased total species richness and common species abundance in reserves increased. In this case study, when there were two or more reserve patches, rare species abundance inside reserves was higher and common species richness in managed stands was higher than when there was a single large reserve. Spillover and habitat amount affected ecological processes underlying these landscape reserve effects. When LCFR positively affected species richness and abundance in reserves or managed stands, >12‐20% reserve cover led to the highest species diversity and abundance. This result is consistent with the target of 17% forested land area in reserves set at the Nagoya biodiversity summit in 2010. Therefore, to preserve biodiversity we recommend at least doubling the current proportion of forest reserves in European forested landscapes.     

Effects of climate and land-cover change on the conservation status of gibbons

Species shift their distribution in response to climate and land-cover change, which may result in a spatial mismatch between currently protected areas (PAs) and priority conservation areas (PCAs). We examined the effects of climate and land-cover change on potential range of gibbons and sought to identify PCAs that would conserve them effectively. We collected global gibbon occurrence points and modeled (ecological niche model) their current and potential 2050s ranges under climate-change and different land-cover-change scenarios. We examined change in range and PA coverage between the current and future ranges of each gibbon species. We applied spatial conservation prioritization to identify the top 30% PCAs for each species. We then determined how much of the PCAs are conserved in each country within the global range of gibbons. On average, 31% (SD 22) of each species’ current range was covered in PAs. PA coverage of the current range of 9 species was <30%. Nine species lost on average 46% (SD 29) of their potential range due to climate change. Under climate-change with an optimistic land-cover-change scenario (B1), 12 species lost 39% (SD 28) of their range. In a pessimistic land-cover-change scenario (A2), 15 species lost 36% (SD 28) of their range. Five species lost significantly more range under the A2 scenario than the B1 scenario (p = 0.01, SD 0.01), suggesting that gibbons will benefit from effective management of land cover. PA coverage of future range was <30% for 11 species. On average, 32% (SD 25) of PCAs were covered by PAs. Indonesia contained more species and PCAs and thus has the greatest responsibility for gibbon conservation. Indonesia, India, and Myanmar need to expand their PAs to fulfill their responsibility to gibbon conservation. Our results provide a baseline for global gibbon conservation, particularly for countries lacking gibbon research capacity.     

Identifying socioeconomic and biophysical factors driving forest loss in protected areas

Protected areas (PAs) are a commonly used strategy to confront forest conversion and biodiversity loss. Although determining drivers of forest loss is central to conservation success, understanding of them is limited by conventional modeling assumptions. We used random forest regression to evaluate potential drivers of deforestation in PAs in Mexico, while accounting for nonlinear relationships and higher order interactions underlying deforestation processes. Socioeconomic drivers (e.g., road density, human population density) and underlying biophysical conditions (e.g., precipitation, distance to water, elevation, slope) were stronger predictors of forest loss than PA characteristics, such as age, type, and management effectiveness. Within PA characteristics, variables reflecting collaborative and equitable management and PA size were the strongest predictors of forest loss, albeit with less explanatory power than socioeconomic and biophysical variables. In contrast to previously used methods, which typically have been based on the assumption of linear relationships, we found that the associations between most predictors and forest loss are nonlinear. Our results can inform decisions on the allocation of PA resources by strengthening management in PAs with the highest risk of deforestation and help preemptively protect key biodiversity areas that may be vulnerable to deforestation in the future.     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.     

Use of stochastic patch occupancy models in the California red-legged frog for Bayesian inference regarding past events and future persistence

Assessing causes of population decline is critically important to management of threatened species. Stochastic patch occupancy models (SPOMs) are popular tools for examining spatial and temporal dynamics of populations when presence–absence data in multiple habitat patches are available. We developed a Bayesian Markov chain method that extends existing SPOMs by focusing on past environmental changes that may have altered occupancy patterns prior to the beginning of data collection. Using occupancy data from 3 creeks, we applied the method to assess 2 hypothesized causes of population decline—in situ die-off and residual impact of past source population loss—in the California red-legged frog. Despite having no data for the 20–30 years between the hypothetical event leading to population decline and the first data collected, we were able to discriminate among hypotheses, finding evidence that in situ die-off increased in 2 of the creeks. Although the creeks had comparable numbers of occupied segments, owing to different extinction–colonization dynamics, our model predicted an 8-fold difference in persistence probabilities of their populations to 2030. Adding a source population led to a greater predicted persistence probability than did decreasing the in situ die-off, emphasizing that reversing the deleterious impacts of a disturbance may not be the most efficient management strategy. We expect our method will be useful for studying dynamics and evaluating management strategies of many species.     

Parks versus PES: Evaluating direct and incentive-based land conservation in Mexico

Protected areas (PAs) and payments for ecosystem services (PES) are the top two mechanisms available for countries to achieve international REDD agreements, yet there are few empirical comparisons of their effects. We estimate the impacts of PAs and PES on forest conservation, poverty reduction, and population change at the locality level in Mexico in the 2000s. Both policies conserved forest, generating an approximately 20–25% reduction in expected forest cover loss. PES created statistically significant but small poverty alleviation while PAs had overall neutral impacts on livelihoods. Estimates by individual policy type for the same level of deforestation risk indicate that biosphere reserves and PES balanced conservation and livelihood goals better than strict protected areas or mixed-use areas. This suggests that both direct and incentive-based instruments can be effective, and that policies combining sustainable financing, flexible zoning, and recognition of local economic goals are more likely to achieve conservation without harming livelihoods.     

Predator-prey spatial game as a tool to understand the effects of protected areas on harvester-wildlife interactions

No-take reserves are sometimes implemented for sustainable population harvesting because they offer opportunities for animals to spatially avoid harvesters, whereas harvesters can benefit in return from the reserve spillover. Here, we used the framework of predator-prey spatial games to understand how protected areas shape spatial interactions between harvesters and target species and determine animal mortality. In these spatial games, the "predator" searches for "prey" and matches their habitat use, unless it meets spatial constraints offering the opportunity for prey to avoid the mortality source. However, such prey refuges could attract predators in the surroundings, which questions the potential benefits for prey. We located, in the Geneva Basin (France), hunting dogs and wild boar Sus scrofa L. during hunting seasons with global positioning systems and very-high-frequency collars. We quantified how the proximity of the reserve shaped the matching between both habitat uses using multivariate analyses and linked these patterns to animals' mortality with a Cox regression analysis. Results showed that habitat uses by both protagonists disassociated only when hunters were spatially constrained by the reserve. In response, hunters increased hunting efforts near the reserve boundary, which induced a higher risk exposure for animals settled over the reserve. The mortality of adult wild boar decreased near the reserve as the mismatch between both habitat uses increased. However the opposite pattern was determined for younger individuals that suffered from the high level of hunting close to the reserve. The predator-prey analogy was an accurate prediction of how the protected area modified spatial relationships between harvesters and target species. Prey-searching strategies adopted by hunters around reserves strongly impacted animal mortality and the efficiency of the protected area for this harvested species. Increasing reserve sizes and/or implementing buffer areas with harvesting limitations can dampen this edge effect and helps harvesters to benefit durably from source populations of reserves. Predator-prey spatial games therefore provide a powerful theoretical background for understanding wildlife-harvester spatial interactions and developing substantial application for sustainable harvesting.