Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

Hunting and the Likelihood of Extinction of Amazonian Mammals

Species inhabiting tropical forests are thought to be on the verge of mass extinction. Much work has focused on extinction rates caused by deforestation; however, many of the recorded extinctions that have occurred since 1600 were a result of overhunting. We collected data on the relative abundance of large-bodied mammals in the northeastern Peruvian Amazon in areas with persistent hunting pressure and in areas with infrequent hunting pressure. We quantified the effects of hunting by calculating the change in abundance of species between the infrequently and persistently hunted sites. We report that in Amazonian mammals weighing more than 1 kg the degree of population declines caused by hunting is correlated with the species' intrinsic rate of natural increase (r_max  ), longevity, and generation time. Our results show that species with long-lived individuals, low rates of increase, and long generation times are more vulnerable to extinction than species with short-lived individuals, high rates of increase, and shorter generations.     

Hunting for Sustainability in Tropical Secondary Forests

Abstract:  The interaction between land-use change and the sustainability of hunting is poorly understood but is critical for sustaining hunted vertebrate populations and a protein supply for the rural poor. We investigated sustainability of hunting in an Amazonian landscape mosaic, where a small human population had access to large areas of both primary and secondary forest. Harvestable production of mammals and birds was calculated from density estimates. We compared production with offtake from three villages and used catch-per-unit-effort as an independent measure of prey abundance. Most species were hunted unsustainably in primary forest, leading to local depletion of the largest primates and birds. The estimated sustainable supply of wild meat was higher for primary (39 kg · km⁻²· yr⁻¹) than secondary forest (22 kg · km⁻²· yr⁻¹) because four species were absent and three species at low abundance in secondary forests. Production of three disturbance-tolerant mammal species was 3 times higher in secondary than in primary forest, but hunting led to overexploitation of one species. Our data suggest that an average Amazonian smallholder would require ≥3.1 km² of secondary regrowth to ensure a sustainable harvest of forest vertebrates. We conclude that secondary forests can sustainably provide only 2% of the required protein intake of Amazonian smallholders and are unlikely to be sufficient for sustainable hunting in other tropical forest regions.     

Effect of Small‐Scale Heterogeneity of Prey and Hunter Distributions on the Sustainability of Bushmeat Hunting

Abstract: Bushmeat is the main source of protein and the most important source of income for rural people in the Congo Basin, but intensive hunting of bushmeat species is also a major concern for conservationists. Although spatial heterogeneity in hunting effort and in prey populations at the landscape level plays a key role in the sustainability of hunted populations, the role of small‐scale heterogeneity within a village hunting territory in the sustainability of hunting has remained understudied. We built a spatially explicit multiagent model to capture the dynamics of a system in which hunters and preys interact within a village hunting territory. We examined the case of hunting of bay duikers (Cephalophus dorsalis) in the village of Ntsiété, northeastern Gabon. The impact of hunting on prey populations depended on the spatial heterogeneity of hunting and prey distribution at small scales within a hunting area. Within a village territory, the existence of areas hunted throughout the year, areas hunted only during certain seasons, and unhunted areas contributed to the sustainability of the system. Prey abundance and offtake per hunter were particularly sensitive to the frequency and length of hunting sessions and to the number of hunters sharing an area. Some biological parameters of the prey species, such as dispersal rate and territory size, determined their spatial distribution in a hunting area, which in turn influenced the sustainability of hunting. Detailed knowledge of species ecology and behavior, and of hunting practices are crucial to understanding the distribution of potential sinks and sources in space and time. Given the recognized failure of simple biological models to assess maximum sustainable yields, multiagent models provide an innovative path toward new approaches for the assessment of hunting sustainability, provided further research is conducted to increase knowledge of prey species’ and hunter behavior.     

Effects of Subsistence Hunting on Vertebrate Community Structure in Amazonian Forests

Abstract: Subsistence hunting affects vast tracts of tropical wilderness that otherwise remain structurally unaltered, yet distinguishing hunted from nonhunted tropical forests presents a difficult problem because this diffuse form of resource extraction leaves few visible signs of its occurrence. I used a standardized series of line-transect censuses conducted over a 10-year period to examine the effects of subsistence game harvest on the structure of vertebrate communities in 25 Amazonian forest sites subjected to varying levels of hunting pressure. Crude vertebrate biomass, which was highly correlated with hunting pressure, gradually declined from nearly 1200 kg km⁻² at nonhunted sites to less than 200 kg km⁻² at heavily hunted sites. Hunting had a negative effect on the total biomass and relative abundance of vertebrate species in different size classes at these forest sites, but it did not affect their overall density. In particular, persistent hunting markedly reduced the density of large-bodied game species (>5 kg), which contributed a large proportion of the overall community biomass at nonhunted sites (65–78%) and lightly hunted sites (55–71%). Nutrient-rich floodplain forests contained a consistently greater game biomass than nutrient-poor unflooded forests, once I controlled for the effects of hunting pressure. Conservative estimates of game yields indicate that as many as 23.5 million game vertebrates, equivalent to 89,224 tons of bushmeat with a market value of US$190.7 million, are consumed each year by the rural population of Brazilian Amazonia, which illustrates the enormous socioeconomic value of game resources in the region. My cross-site comparison documents the staggering effect of subsistence hunters on tropical forest vertebrate communities and highlights the importance of considering forest types and forest productivity in game management programs.     

Effect of Hunting in Source-Sink Systems in the Neotropics

Abstract: Previous studies of the sustainability of wildlife hunting in the Neotropics have not considered the potential dispersal of animals into hunted areas. A literature review of studies of subsistence hunting in the Neotropics suggests that hunting is often conducted in areas adjacent to relatively undisturbed habitat that may act as sources of animals for the hunted sites. We compared studies of tapir (   Tapirus terrestris ) hunting at different sites to illustrate the potential bias of sustainability evaluations based on local productivity. The limited information available suggests that dispersal could have a key role in rebuilding animal populations depleted by hunting. Thus, factors that strongly affect dispersal—such as spatial distribution and size of areas with and without hunting, population size in source areas, and social behavior—should be considered when the sustainability of hunting is evaluated in areas with heterogeneous hunting pressure. We suggest the application of two models that use spatial controls (recognizing the potential source-sink nature of some hunted systems and protecting unhunted refugia) to avoid wildlife overexploitation when biological data and enforcement capabilities to regulate harvests are limited. This approach may produce more reliable evaluations of sustainability, provide information on the dynamics of hunting systems, and help local communities and policymakers conserve key areas (including protected areas) that may act as game sources.     

Modeling the Effects of Trophy Selection and Environmental Disturbance on a Simulated Population of African Lions

Abstract:  Tanzania is a premier destination for trophy hunting of African lions (Panthera leo) and is home to the most extensive long-term study of unhunted lions. Thus, it provides a unique opportunity to apply data from a long-term field study to a conservation dilemma: How can a trophy-hunted species whose reproductive success is closely tied to social stability be harvested sustainably? We used an individually based, spatially explicit, stochastic model, parameterized with nearly 40 years of behavioral and demographic data on lions in the Serengeti, to examine the separate effects of trophy selection and environmental disturbance on the viability of a simulated lion population in response to annual harvesting. Female population size was sensitive to the harvesting of young males (≥3 years), whereas hunting represented a relatively trivial threat to population viability when the harvest was restricted to mature males (≥6 years). Overall model performance was robust to environmental disturbance and to errors in age assessment based on nose coloration as an index used to age potential trophies. Introducing an environmental disturbance did not eliminate the capacity to maintain a viable breeding population when harvesting only older males, and initially depleted populations recovered within 15–25 years after the disturbance to levels comparable to hunted populations that did not experience a catastrophic event. These results are consistent with empirical observations of lion resilience to environmental stochasticity .     

Rapid Evolution in the Wild: Changes in Body Size, Life-History Traits, and Behavior in Hunted Populations of the Japanese Mamushi Snake

Abstract:  Rapid evolution caused by human exploitation of wildlife is not usually addressed in studies of the impacts of such exploitation despite its direct relevance to population persistence. Japanese mamushi ( Gloydius blomhoffii ), an endemic venomous snake of the Japanese archipelago, has been heavily hunted by humans, and many populations appear to be declining or are already extirpated. We compared local populations that have been hunted regularly with populations that have not been hunted. Mamushi in hunted populations were smaller, had fewer vertebrae, produced more and smaller offspring, had increased reproductive effort among smaller females, and in nature fled at greater distances from an approaching human and were less defensive than mamushi in unhunted populations, as predicted from life-history theory. Heritability estimates for body size, number of vertebrae, and antipredator behavior were statistically significant, and neonates from hunted sites showed the same distribution of altered characters (compared with those from unhunted sites) as adults. Thus, distribution of the divergent trait between hunted and unhunted sites appeared in part to be genetically based, which suggests rapid evolution to human predation pressures. Trait distributions in hunted populations probably deviate from naturally (as opposed to anthropogenically) selected optima and, therefore, may have long-term negative repercussions on population persistence. Because rapid evolution affects a suite of parameters that characterize exploited populations, accurate understanding of the impacts of exploitation and effective resource management and conservation can only be achieved if evolutionary consequences are considered explicitly.     

Assessment of mammal reproduction for hunting sustainability through community‐based sampling of species in the wild

Wildlife subsistence hunting is a major source of protein for tropical rural populations and a prominent conservation issue. The intrinsic rate of natural increase. (r_max) of populations is a key reproductive parameter in the most used assessments of hunting sustainability. However, researchers face severe difficulties in obtaining reproductive data in the wild, so these assessments often rely on classic reproductive rates calculated mostly from studies of captive animals conducted 30 years ago. The result is a flaw in almost 50% of studies, which hampers management decision making. We conducted a 15‐year study in the Amazon in which we used reproductive data from the genitalia of 950 hunted female mammals. Genitalia were collected by local hunters. We examined tissue from these samples to estimate birthrates for wild populations of the 10 most hunted mammals. We compared our estimates with classic measures and considered the utility of the use of r_max in sustainability assessments. For woolly monkey (Lagothrix poeppigii) and tapir (Tapirus terrestris), wild birthrates were similar to those from captive populations, whereas birthrates for other ungulates and lowland‐paca (Cuniculus paca) were significantly lower than previous estimates. Conversely, for capuchin monkeys (Sapajus macrocephalus), agoutis (Dasyprocta sp.), and coatis (Nasua nasua), our calculated reproductive rates greatly exceeded often‐used values. Researchers could keep applying classic measures compatible with our estimates, but for other species previous estimates of r_max may not be appropriate. We suggest that data from local studies be used to set hunting quotas. Our maximum rates of population growth in the wild correlated with body weight, which suggests that our method is consistent and reliable. Integration of this method into community‐based wildlife management and the training of local hunters to record pregnancies in hunted animals could efficiently generate useful information of life histories of wild species and thus improve management of natural resources.     

Human Hunting of Nongame Birds at Zuni, New Mexico, U.S.A.

Abstract: Nongame bird hunting is a critical activity of the Zuni people of the southwestern United States. To help determine whether their current hunting practices may be negatively affecting bird populations on the Zuni Reservation, we interviewed 98 Zuni hunters. Nongame bird hunting was practiced by a large portion of Zuni males (45%); the most active age group was 20- to 49-year-olds. The rate a species was hunted was not a function of its abundance at Zuni but seemed instead to be related to its cultural demand. Five "species"—bluebirds ( Sialia currucoides , S. mexicana ), Northern Flicker ( Colaptes auratus ), woodpeckers ( Picoides villosus , P. pubescens , Melanerpes lewis ), Steller's Jay ( Cyanocitta stelleri ), and American Kestrel ( Falco sparverius )—made up 77% of all birds taken. The two most heavily hunted species were each taken in numbers> 10,000 individuals per year. Although the greatest number of hunters were active in autumn, 31% hunted in spring, despite discouragement by the tribal government. Habitats favored by hunters were coniferous forests and riparian areas. Rates of hunting of nongame birds at Zuni equaled or exceeded those reported for game birds hunted by indigenous hunters in the Neotropics and New Mexico. The Zuni believe that some heavily hunted species are decreasing in number because of hunting. Although we cannot prove this at present, we suggest measures to mitigate possible overhunting, including the creation of refugia and a reduction of spring hunting.     

Impacts of Roads, Hunting, and Habitat Alteration on Nocturnal Mammals in African Rainforests

Abstract:  Nocturnal mammals are poorly studied in Central Africa, a region experiencing dramatic increases in logging, roads, and hunting activity. In the rainforests of southern Gabon, we used spotlighting surveys to estimate abundances of nocturnal mammal species and guilds at varying distances from forest roads and between hunted and unhunted treatments (comparing a 130-km² oil concession that was nearly free of hunting, with nearby areas outside the concession that had moderate hunting pressure). At each of 12 study sites that were evenly divided between hunted and unhunted areas, we established standardized 1-km transects along road verges and at 50, 300, and 600 m from the road. We then repeatedly surveyed mammals at each site during 2006. Hunting had few apparent effects on this assemblage. Nevertheless, the species richness and often the abundance of nocturnal primates, smaller ungulates, and carnivores were significantly depressed within approximately 30 m of roads. Scansorial rodents increased in abundance in hunted forests, possibly in response to habitat changes caused by logging or nearby swidden farming. In multiple-regression models many species and guilds were significantly influenced by forest-canopy and understory cover, both of which are altered by logging and by certain abiotic variables. In general, nocturnal species, many of which are arboreal or relatively small in size (<10 kg), were less strongly influenced by hunting and more strongly affected by human-induced changes in forest structure than were larger mammal species in our study area.     

Population viability and harvest sustainability for Madagascar lemurs

Subsistence hunting presents a conservation challenge by which biodiversity preservation must be balanced with safeguarding of human livelihoods. Globally, subsistence hunting threatens primate populations, including Madagascar's endemic lemurs. We used population viability analysis to assess the sustainability of lemur hunting in Makira Natural Park, Madagascar. We identified trends in seasonal hunting of 11 Makira lemur species from household interview data, estimated local lemur densities in populations adjacent to focal villages via transect surveys, and quantified extinction vulnerability for these populations based on species-specific demographic parameters and empirically derived hunting rates. We compared stage-based Lefkovitch with periodic Leslie matrices to evaluate the impact of regional dispersal on persistence trajectories and explored the consequences of perturbations to the timing of peak hunting relative to the lemur birth pulse, under assumptions of density-dependent reproductive compensation. Lemur hunting peaked during the fruit-abundant wet season (March–June). Estimated local lemur densities were roughly inverse to body size across our study area. Life-history modeling indicated that hunting most severely threatened the species with the largest bodies (i.e., Hapalemur occidentalis, Avahi laniger, Daubentonia madagascariensis, and Indri indi), characterized by late-age reproductive onsets and long interbirth intervals. In model simulations, lemur dispersal within a regional metapopulation buffered extinction threats when a majority of local sites supported growth rates above the replacement level but drove regional extirpations when most local sites were overharvested. Hunt simulations were most detrimental when timed to overlap lemur births (a reality for D. madagascariensis and I. indri). In sum, Makira lemurs were overharvested. Regional extirpations, which may contribute to broad-scale extinctions, will be likely if current hunting rates persist. Cessation of anthropogenic lemur harvest is a conservation priority, and development programs are needed to help communities switch from wildlife consumption to domestic protein alternatives.     

Mitochondrial and microsatellite assessment of population structure of South American sea lion (<Emphasis Type="Italic">Otaria flavescens</Emphasis>) in the Southwestern Atlantic Ocean

Several major breeding areas have been defined for the South American sea lion (Otaria flavescens) along the Atlantic Ocean including the Uruguayan and Patagonian coasts. Together with a documented and severe reduction in population sizes caused by commercial hunting in the last century, these areas show opposite population trends. While Patagonian populations are recovering since hunting ceased, Uruguayan populations are declining. In this context, population genetic structure and genetic diversity were studied for the first time with both nuclear (microsatellites) and mitochondrial (control region) markers together. Alternative scenarios were found for both markers. While mitochondrial marker showed geographically structured populations, the nuclear loci showed a lack of geographical structure. These opposite patterns in genetic structure could be explained by female phylopatry and high male dispersion. The reduction in population size caused by commercial hunting did not leave a detectable footprint of bottleneck at the genetic level.     

Past,present, and future of a freshwater fish metapopulation in a threatened landscape

It is well documented that hydropower plants can affect the dynamics of fish populations through landscape alterations and the creation of new barriers. Less emphasis has been placed on the examination of the genetic consequences for fish populations of the construction of dams. The relatively few studies that focus on genetics often do not consider colonization history and even fewer tend to use this information for conservation purposes. As a case study, we used a 3‐pronged approach to study the influence of historical processes, contemporary landscape features, and potential future anthropogenic changes in landscape on the genetic diversity of a fish metapopulation. Our goal was to identify the metapopulation's main attributes, detect priority areas for conservation, and assess the consequences of the construction of hydropower plants for the persistence of the metapopulation. We used microsatellite markers and coalescent approaches to examine historical colonization processes, traditional population genetics, and simulations of future populations under alternate scenarios of population size reduction and gene flow. Historical gene flow appeared to have declined relatively recently and contemporary populations appeared highly susceptible to changes in landscape. Gene flow is critical for population persistence. We found that hydropower plants could lead to a rapid reduction in number of alleles and to population extirpation 50–80 years after their construction. More generally, our 3‐pronged approach for the analyses of empirical genetic data can provide policy makers with information on the potential impacts of landscape changes and thus lead to more robust conservation efforts.     

Impacts of Roads and Hunting on Central African Rainforest Mammals

Abstract:  Road expansion and associated increases in hunting pressure are a rapidly growing threat to African tropical wildlife. In the rainforests of southern Gabon, we compared abundances of larger (>1 kg) mammal species at varying distances from forest roads and between hunted and unhunted treatments (comparing a 130-km² oil concession that was almost entirely protected from hunting with nearby areas outside the concession that had moderate hunting pressure). At each of 12 study sites that were evenly divided between hunted and unhunted areas, we established standardized 1-km transects at five distances (50, 300, 600, 900, and 1200 m) from an unpaved road, and then repeatedly surveyed mammals during the 2004 dry and wet seasons. Hunting had the greatest impact on duikers ( Cephalophus spp. ), forest buffalo ( Syncerus caffer nanus ), and red river hogs ( Potamochoerus porcus ), which declined in abundance outside the oil concession, and lesser effects on lowland gorillas ( Gorilla gorilla gorilla ) and carnivores. Roads depressed abundances of duikers, sitatungas ( Tragelaphus spekei gratus ), and forest elephants ( Loxondonta africana cyclotis ), with avoidance of roads being stronger outside than inside the concession. Five monkey species showed little response to roads or hunting, whereas some rodents and pangolins increased in abundance outside the concession, possibly in response to greater forest disturbance. Our findings suggest that even moderate hunting pressure can markedly alter the structure of mammal communities in central Africa. Roads had the greatest impacts on large and small ungulates, with the magnitude of road avoidance increasing with local hunting pressure.     

Synergistic Effects of Subsistence Hunting and Habitat Fragmentation on Amazonian Forest Vertebrates

Abstract: Subsistence game hunting has profound negative effects on the species diversity, standing biomass, and size structure of vertebrate assemblages in Amazonian forests that otherwise remain largely undisturbed. These effects are likely to be considerably aggravated by forest fragmentation because fragments are more accessible to hunters, allow no (or very low rates of  ) recolonization from nonharvested source populations, and may provide a lower-quality resource base for the frugivore-granivore vertebrate fauna. I examined the likelihood of midsized to large-bodied bird and mammal populations persisting in Amazonian forest fragments of variable sizes whenever they continue to be harvested by subsistence hunters in the aftermath of isolation. I used data from a comprehensive compilation of game-harvest studies throughout Neotropical forests to estimate the degree to which different species and populations have been overharvested and then calculated the range of minimum forest areas required to maintain a sustainable harvest. The size distribution of 5564 Amazonian forest fragments—estimated from Landsat images of six regions of southern and eastern Brazilian Amazonia—clearly shows that these are predominantly small and rarely exceed 10 ha, suggesting that persistent overhunting is likely to drive most midsized to large vertebrate populations to local extinction in fragmented forest landscapes. Although experimental studies on this negative synergism remain largely unavailable, the prospect that increasingly fragmented Neotropical forest regions can retain their full assemblages of avian and mammalian species is unlikely.     

The Sustainability of Subsistence Hunting in the Neotropics

Hunting is an important component of native subsistence strategies in Amazonia. It is also a serious threat to biodiversity in some areas. We present data on the faunal harvests of two native Neotropical subsistence hunting peoples, Machiguenga bow-hunters and Piro shotgun hunters of Peru. The rate of annual harvest per square kilometer of catchment is estimated and compared to rates of sustainable harvest calculated by Robinson and Redford (1991). We used indicators of prey abundance to test for the depletion of species that hunters killed in numbers greater than what the model predicts to be sustainable. As predicted, we found the strongest evidence for local depletion of the large primates at the Piro site. The woolly monkey ( Lagothrix lagothricha) harvest at the Machiguenga village and the deer ( Mazama americana) and collared peccary (Tayassu tajacu) harvests at both sites were not predicted to be unsustainable and we found no evidence for depletion. Machiguenga bow hunters killed spider monkeys (Ateles paniscus) in quantities that were slightly above what the model predicted to be sustainable, yet we found no evidence for depletion. Differential species vulnerability, catchment size, and consumer population size could be important factors in determining sustainability.     

Costs and Benefits of the Presence of Leopards to the Sport‐Hunting Industry and Local Communities in Niassa National Reserve,Mozambique

Sport hunting is often proposed as a tool to support the conservation of large carnivores. However, it is challenging to provide tangible economic benefits from this activity as an incentive for local people to conserve carnivores. We assessed economic gains from sport hunting and poaching of leopards (Panthera pardus), costs of leopard depredation of livestock, and attitudes of people toward leopards in Niassa National Reserve, Mozambique. We sent questionnaires to hunting concessionaires (n = 8) to investigate the economic value of and the relative importance of leopards relative to other key trophy‐hunted species. We asked villagers (n = 158) the number of and prices for leopards poached in the reserve and the number of goats depredated by leopard. Leopards were the mainstay of the hunting industry; a single animal was worth approximately U.S.$24,000. Most safari revenues are retained at national and international levels, but poached leopard are illegally traded locally for small amounts ($83). Leopards depredated 11 goats over 2 years in 2 of 4 surveyed villages resulting in losses of $440 to 6 households. People in these households had negative attitudes toward leopards. Although leopard sport hunting generates larger gross revenues than poaching, illegal hunting provides higher economic benefits for households involved in the activity. Sport‐hunting revenues did not compensate for the economic losses of livestock at the household level. On the basis of our results, we propose that poaching be reduced by increasing the costs of apprehension and that the economic benefits from leopard sport hunting be used to improve community livelihoods and provide incentives not to poach. Costos y Beneficios de la Presencia de Leopardos para la Industria de la Caza Deportiva y las Comunidades Locales en Niassa, Reserva Nacional, Mozambique     

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.