Incorporating consumer–resource spatial interactions in reserve design

The persistence of species in reserves depends in large part on the persistence of functional ecological interactions. Despite their importance, however, ecological interactions have not yet been explicitly incorporated into conservation prioritization methods. We develop here a general method for incorporating consumer–resource interactions into spatial reserve design. This method protects spatial consumer–resource interactions by protecting areas that maintain the connectivity between the distribution of consumers and resources. We illustrate our method with a conservation planning case study of a mammalian predator, American marten (Martes americana), and its two primary prey species, Red-backed vole (Clethrionomys rutilus) and Deer mouse (Peromyscus maniculatus). The conservation goal was to identify a reserve for marten that comprised 12% of a forest management unit in the boreal forest in Québec, Canada. We compared reserves developed using analysis variants that utilized different levels of information about predator and prey habitat distributions, species-specific connectivity requirements, and interaction connectivity requirements. The inclusion of consumer–resource interactions in reserve-selection resulted in spatially aggregated reserves that maintained local habitat quality for the species. This spatial aggregation was not induced by applying a qualitative penalty for the boundary length of the reserve, but rather was a direct consequence of modelling the spatial needs of the interacting consumer and resources. Our method for maintaining connectivity between consumers and their resources within reserves can be applied even under the most extreme cases of either complete spatial overlap or complete spatial segregation of consumer–resource distributions. The method has been made available via public software.     

Fishing the line near marine reserves in single and multispecies fisheries.

Throughout the world "fishing the line" is a frequent harvesting tactic in communities where no-take marine reserves are designated. This practice of concentrating fishing effort at the boundary of a marine reserve is predicated upon the principle of spillover, the net export of stock from the marine reserve to the surrounding unprotected waters. We explore the consequences and optimality of fishing the line using a spatially explicit theoretical model. We show that fishing the line: (1) is part of the optimal effort distribution near no-take marine reserves with mobile species regardless of the cooperation level among harvesters; (2) has a significant impact on the spatial patterns of catch per unit effort (CPUE) and fish density both within and outside of the reserve; and (3) can enhance total population size and catch simultaneously under a limited set of conditions for overexploited populations. Additionally, we explore the consequences of basing the spatial distribution of fishing effort for a multispecies fishery upon the optimality of the most mobile species that exhibits the greatest spillover. Our results show that the intensity of effort allocated to fishing the line should instead be based upon more intermediate rates of mobility within the targeted community. We conclude with a comparison between model predictions and empirical findings from a density gradient study of two important game fish in the vicinity of a no-take marine-life refuge on Santa Catalina Island, California (USA). These results reveal the need for empirical studies to account for harvester behavior and suggest that the implications of spatial discontinuities such as fishing the line should be incorporated into marine-reserve design.     

Impacts of hunting on tropical forests in Southeast Asia

Although deforestation and forest degradation have long been considered the most significant threats to tropical biodiversity, across Southeast Asia (Northeast India, Indochina, Sundaland, Philippines) substantial areas of natural habitat have few wild animals (>1 kg), bar a few hunting‐tolerant species. To document hunting impacts on vertebrate populations regionally, we conducted an extensive literature review, including papers in local journals and reports of governmental and nongovernmental agencies. Evidence from multiple sites indicated animal populations declined precipitously across the region since approximately 1980, and many species are now extirpated from substantial portions of their former ranges. Hunting is by far the greatest immediate threat to the survival of most of the region's endangered vertebrates. Causes of recent overhunting include improved access to forests and markets, improved hunting technology, and escalating demand for wild meat, wildlife‐derived medicinal products, and wild animals as pets. Although hunters often take common species, such as pigs or rats, for their own consumption, they take rarer species opportunistically and sell surplus meat and commercially valuable products. There is also widespread targeted hunting of high‐value species. Consequently, as currently practiced, hunting cannot be considered sustainable anywhere in the region, and in most places enforcement of protected‐area and protected‐species legislation is weak. The international community's focus on cross‐border trade fails to address overexploitation of wildlife because hunting and the sale of wild meat is largely a local issue and most of the harvest is consumed in villages, rural towns, and nearby cities. In addition to improved enforcement, efforts to engage hunters and manage wildlife populations through sustainable hunting practices are urgently needed. Unless there is a step change in efforts to reduce wildlife exploitation to sustainable levels, the region will likely lose most of its iconic species, and many others besides, within the next few years.     

Consequences of patch reef spacing for density-dependent mortality of coral-reef fishes

The spatial configuration of habitat patches can profoundly affect a number of ecological interactions, including those between predators and prey. I examined the effects of reef spacing on predator-prey interactions within coral-reef fish assemblages in the Bahamas. Using manipulative field experiments, I determined that reef spacing influences whether and how density-dependent predation occurs. Mortality rates of juveniles of two ecologically dissimilar species (beaugregory damselfish and yellowhead wrasse) were similarly affected by reef spacing; for both species, mortality was density dependent on reef patches that were spatially isolated (separated by 50 m), and density independent on reef patches that were aggregated (separated by 5 m). A subsequent experiment with the damselfish demonstrated that a common resident predator (coney) caused a substantial proportion of the observed mortality, independent of reef spacing. Compared to isolated reefs, aggregated reefs were much more likely to be visited by transient predators (mostly yellowtail snappers), regardless of prey density, and on these reefs, mortality rates approached 100% for both prey species. Transient predators exhibited neither an aggregative response nor a type 3 functional response, and consequently were not the source of density dependence observed on the isolated reefs. These patterns suggest that resident predators caused density-dependent mortality in their prey through type 3 functional responses on all reefs, but on aggregated reefs, this density dependence was overwhelmed by high, density-independent mortality caused by transient predators. Thus, the spatial configuration of reef habitat affected both the magnitude of total predation and the existence of density-dependent mortality. The combined effects of the increasing fragmentation of coral reef habitats at numerous scales and global declines in predatory fish may have important consequences for the regulation of resident fish populations.     

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.     

Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

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.     

Effects of Landscape Features on the Distribution and Sustainability of Ungulate Hunting in Northern Congo

Abstract: Understanding the spatial dimensions of hunting and prey population dynamics is important in order to estimate the sustainability of hunting in tropical forests. We investigated how hunting offtake of vertebrates differed in mixed forest and monodominant forest (composed of Gilbertiodendron dewevrei) and over different spatial extents within the hunting catchment around the logging town of Kabo, Congo. In 9 months of recall surveys with hunters, we gathered information on over 1500 hunting trips in which ungulates were 65% of the species killed and 82% of harvested biomass. Hunters supplied information on animals killed and the hunting trip, including the area visited (i.e., hunting zone; 11 separate zones within a 506 km² catchment or commonly hunted area). Over 65% of all animals were killed in monodominant forest, which made up 28% of the hunting catchment, and zones with small amounts of monodominant forest were used most frequently by hunters. Given the large offtakes from monodominant forests, we suggest that animal dispersal may be maintaining high, localized harvests in these areas. We believe hunters preferred to hunt in monodominant forest because the understory was accessible and that areas with small amounts of monodominant forest and large amounts of mixed forest were more productive. The variation in hunting pressure we found between and within hunting zones differs from past examinations of spatial variation in hunting offtake, where entire hunting catchments were considered population sinks and areas with low to no hunting (no‐take zones) were outside hunting catchments. Future use of no‐take zones to manage hunting should incorporate variability in offtake within hunting catchments.     

Temperate marine reserves enhance targeted but not untargeted fishes in multiple no-take MPAs.

Although many papers report the effects of no-take marine protected areas (MPAs or reserves), scientifically rigorous empirical studies are rare, particularly for temperate reef fishes. We evaluated the responses of fish populations to protection from fishing in reserves by comparing densities and sizes inside and outside of five no-take reserves in southern California, USA. Our results are robust because we compared responses across multiple rocky-reef reserves in two different years and controlled for possible site differences by (a) ensuring that habitat characteristics were the same inside and outside reserves, and (b) sampling species that are not targeted, which would not be expected to have a direct response to fishing. We compared fish density and size and calculated biomass and egg production across all five sites. Fishes targeted by recreational and/or commercial fisheries consistently exhibited increases in mean density (150%), size (30%), biomass (440%), and egg production (730%) inside reserves. Reserve effects were greatest for legal-sized targeted fishes: significantly greater densities were found exclusively inside reserves for targeted species (580%), the largest size classes existed only inside reserves, and mean biomass was 1000% higher. These responses were unlikely to have been caused by habitat differences because there were no significant differences in habitat characteristics between reserve and control locations. Densities of non-targeted species did not differ between reserve and non-reserve locations, further supporting the conclusions that differences in targeted species between reserve and control locations were due to harvesting rather than site-specific effects. Although MPAs cannot replace traditional fisheries management, the concentration of increased biomass and egg production is a unique MPA benefit that serves both reserves and fisheries. Scientifically rigorous studies that include multiple reserves, such as this study, are needed to inform management and policy decisions.     

Response of predators to loss and fragmentation of prey habitat: a review of theory

Despite extensive empirical research and previous reviews, no clear patterns regarding the effects of habitat loss and fragmentation on predator-prey interactions have emerged. We suggest that this is because empirical researchers do not design their studies to test specific hypotheses arising from the theoretical literature. In fact, theoretical work is almost completely ignored by empirical researchers, perhaps because it may be inaccessible to them. The purpose of this paper is to review theoretical work on the effects of habitat loss and fragmentation on predator-prey interactions. We provide a summary of clear, testable theoretical predictions for empirical researchers. To test one or more of these predictions, an empiricist will need certain information on the predator and prey species of interest. This includes: (1) whether the predator is a specialist on one prey species or feeds on many kinds of prey (omnivore and generalist); (2) whether the predator is restricted to the same habitat type as the focal prey (specialist), can use a variety of habitats but has higher survival in the prey habitat (omnivore), or lives primarily outside of the focal prey's habitat (generalist); (3) whether prey-only patches have lower prey extinction rates than predator-prey patches; and (4) whether the prey emigrate at higher rates from predator-prey patches than from prey-only patches. Empiricists also need to be clear on whether they are testing a prediction about habitat loss or habitat fragmentation and need to conduct empirical studies at spatial scales appropriate for testing the theoretical prediction(s). We suggest that appropriate use of the theoretical predictions in future empirical research will resolve the apparent inconsistencies in the empirical literature on this topic.     

Predator diversity and trophic interactions

The recognition that predators play important roles in ecosystems has prompted research to resolve how combinations of predator species influence ecosystem functions. Interactions among predator species and their prey can lead to a host of linear and nonlinear effects. Understanding the conditions causing these effects is critical for assigning predator species to functional groups in ways that lead to predictive theory of predator diversity effects on trophic interactions. To this end, I provide a synthesis of experiments examining multiple-predator-species effects on mortality of single shared prey. I show how experimental design and experimental venue can determine the conclusion about the importance of predator diversity on trophic interactions. In addition, I link natural history insights on predator species habitat and hunting behavior with linear and nonlinear multiple-predator effects to derive a new concept of predator diversity effects on trophic interactions. This concept holds that the nature of predator diversity effects is contingent upon predator species hunting mode plus predator and prey species habitat domain (defined as the spatial extent to which a microhabitat is used by a species). This concept allows the classification of multiple-predator effects into four broad functional categories: substitutable, nonlinear due to predator species interference, nonlinear due to intraguild predation, and nonlinear due to predator species synergism. Experimental evidence so far provides ample and comparatively equal support for substitutable, interference, and intraguild effects, and equivocal support for nonlinear synergisms. The paper closes by discussing ways to further a research program aimed at using the building blocks presented here to understand predator functional diversity and trophic interactions in complex ecological systems.     

Campesino hunting and conservation in Latin America

Hunting presents a paradox for biodiversity conservation. It is both a problem and a solution to species declines and poverty. Yet, conservation scientists hold different assumptions about the significance and sustainability of hunting based on the cultures and identities of hunters. In Latin America, conservationists largely sort hunters as either indigenous or campesino. Indigenous hunters are often characterized as culturally driven stewards of wildlife sustainability. Campesino hunters, by contrast, are described as peasants—cultureless, uneducated, and uncaring toward wildlife sustainability. Although such ethnically fueled hunting discourse promotes hunting research, campesino hunters remain underrepresented in most comparative hunting reviews. Moreover, there are no targeted syntheses on the current state of knowledge about campesino hunting, nothing to guide conservation research and practice with and for the largest group of hunters in Latin America. We reviewed 334 articles published from 1937 to 2018 in English (55%) and Spanish (45%)—mostly published in 145 peer-reviewed journals—on the meanings, motivations, and sustainability of campesino hunting in Latin America. Although studies spanned 17 countries, 7 ecosystems, and >75 indigenous and nonindigenous demographics in 30 research contexts, they predominantly focused on nonindigenous campesinos for species-specific conservation and protected area management in tropical broadleaf forests of Mexico, Peru, and Colombia. Authors used 12 methods to collect campesino hunting data, primarily interviews, surveys, and questionnaires, and drew from 10 local and traditional knowledge themes about wildlife trends and uses. Eighteen drivers, 14 constraints, and 10 conflicts—mainly subsistence, income, ethics, regulations, and crop or livestock protection—shaped whether campesino hunters pursued 799 species, 70% of which were least concern species. Yet, only 25 studies (8%) empirically assessed sustainability. Our results show the need for increased interdisciplinary and geographic engagement with campesino hunting across Latin America.     

Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum

According to the weight and size of their prey, Ectatomma ruidum workers can employ different recruitment systems (solitary hunting, cooperative hunting and group hunting with recruitment) when mastering and retrieving prey items from short distances from the nest. Prey size determined the backwards entry typically adopted by this species, while prey weight determined the predatory strategy selected. After a common initial sequence (search for prey, detection, localization), predatory sequences varied in terms of the type of approach, the site of seizure, the reaction after stinging and the type of transport. Nevertheless, irrespective of prey weight and size, seizure was preferentially oriented towards the head and prey were always stung. Short-range recruitment and mass recruitment without trail laying were elicited by a large range of heavy prey (> 2.5 times the weight of an individual worker). According to the mortality risk associated with each prey, hunters exhibited a “prudent” stinging posture associated with an increase in the duration of the subsequent phase of waiting for prey immobilization. The overall time of capture was positively correlated with the weight of the prey. When collective hunting strategies were involved, E. ruidum colonies matched the number of recruited hunters to the size and weight of the prey. Compared to solitary hunting strategies, for short food–nest distances, this graded recruitment appeared to enhance the energetic benefits derived by this species from the use of recruitment systems: the higher the number of workers involved in the recruitment process, the greater the energetic benefits obtained. The exhibition or absence of trail laying behavior in the recruitment responses displayed by E. ruidum workers is discussed in relation to their involvement in scavenging or predatory behavior. Received: 27 June 1996 / Accepted after revision: 3 March 1997     

Sustainability and comanagement of subsistence hunting in an indigenous reserve in Guyana

Although hunting is a key component of subsistence strategies of many Amazonians, it is also one of the greatest threats to wildlife. Because indigenous reserves comprise over 20% of Amazonia, effective conservation often requires that conservation professionals work closely with indigenous groups to manage resource use. We used hunter‐generated harvesting data in spatially explicit biodemographic models to assess the sustainability of subsistence hunting of indigenous Waiwai in Guyana. We collected data through a hunter self‐monitoring program, systematic follows of hunters, and semistructured interviews. We used these data to predict future densities of 2 indicator species, spider monkeys (Ateles paniscus) and bearded sakis (Chiropotes sagulatus), under different scenarios of human population expansion and changing hunting technology. We used encounter rates from transect surveys and hunter catch‐per‐unit effort (CPUE) to validate model predictions. Paca (Cuniculus paca) (198 /year), Currosaw (Crax alector) (168), and spider monkey (117) were the most frequently harvested species. Predicted densities of spider monkeys were statistically indistinguishable from empirically derived transect data (Kolmogorov–Smirnov D = 0.67, p = 0.759) and CPUE (D = 0.32, p = 1.000), demonstrating the robustness of model predictions. Ateles paniscus and C. sagulatus were predicted to be extirpated from <13% of the Waiwai reserve in 20 years, even under the most intensive hunting scenarios. Our results suggest Waiwai hunting is currently sustainable, primarily due to their low population density and use of bow and arrow. Continual monitoring is necessary, however, particularly if human population increases are accompanied by a switch to shotgun‐only hunting. We suggest that hunter self‐monitoring and biodemographic modeling can be used effectively in a comanagement approach in which indigenous parabiologists continuously provide hunting data that is then used to update model parameters and validate model predictions.     

Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.

We examined the long-term effects (28 years) of habitat loss and phenotype-based selective harvest on body mass, horn size, and horn shape of mouflon (Ovis gmelini musimon) in southern France. This population has experienced habitat deterioration (loss of 50.8% of open area) since its introduction in 1956 and unrestricted selective hunting of the largest horned males since 1973. Both processes are predicted to lead to a decrease in phenotype quality by decreasing habitat quality and by reducing the reproductive contribution of individuals carrying traits that are targeted by hunters. Body mass and body size of both sexes and horn measurements of males markedly decreased (by 3.4-38.3%) in all age classes from the 1970s. Lamb body mass varied in relation to the spatiotemporal variation of habitat closure within the hunting-free reserve, suggesting that habitat closure explains part of these changes. However, the fact that there was no significant spatial variation in body mass in the early part of the study, when a decline in phenotypic quality already had occurred, provided support for the influence of selective harvesting. We also found that the allometric relationship between horn breadth and horn length changed over the study period. For a given horn length, horn breadth was lower during the second part of the study. This result, as well as changes in horn curve diameter, supports the interpretation that selective harvesting of males based on their horn configuration had evolutionary consequences for horn shape, since this phenotypic trait is less likely to be affected by changes in habitat characteristics. Moreover, males required more time (approximately four years) to develop a desirable trophy, suggesting that trophy hunting favors the reproductive contribution of animals with slow-growing horns. Managers should exploit hunters' desire for trophy males to finance management strategies which ensure a balance between the population and its environment. However, for long-term sustainable exploitation, harvest strategy should also ensure that selectively targeted males are allowed to contribute genetically to the next generations.     

Understanding the distribution of bushmeat hunting effort across landscapes by testing hypotheses about human foraging

Mitigating the massive impacts of defaunation on natural ecosystems requires understanding and predicting hunting effort across the landscape. But such understanding has been hindered by the difficulty of assessing the movement patterns of hunters in thick forests and across complex terrain. We statistically tested hypotheses about the spatial distribution of hunting with circuit theory and structural equation models. We used a data set of >7000 known kill locations in Guyana and hunter movement models to test these methods. Comparing models with different resistance layers (i.e., different estimates of how terrain and land cover influence human movement speed) showed that rivers, on average, limited movement rather than serving as transport arteries. Moreover, far more kills occurred close to villages than in remote areas. This, combined with the lack of support for structural equation models that included latent terms for prey depletion driven by past overhunting, suggests that kill locations in this system tended to be driven by where hunters were currently foraging rather than by influences of historical harvest. These analyses are generalizable to a variety of ecosystems, species, and data types, providing a powerful way of enhancing maps and predictions of hunting effort across complex landscapes.     

Effects of density-dependent dispersal behaviours on the speed and spatial patterns of range expansion in predator-prey metapopulations

Dispersal can strongly affect the spatiotemporal dynamics of a species (its spread, spatial distribution and persistence). We investigated how two dispersal behaviours, namely prey evasion (PE) and predator pursuit (PP), affect the dynamics of a predator-prey system. PE portrays the tendency of prey avoiding predators by dispersing into adjacent patches with fewer predators, while PP describes the tendency of predators to pursue the prey by moving into patches with more prey. Based on the Beddington predation model, a spatially explicit metapopulation model was built to incorporate PE and PP. Numerical simulations were run to investigate the effects of PE and PP on the rate of spread, spatial synchrony and the persistence of populations. Results show that both PE and PP can alter spatial synchrony although PP has a weaker desynchronising effect than PE. The predator-prey system without PE and PP expanded in circular waves. The effect of PE can push the prey to distribute in a circular ring front, whereas the effect of PP can change the circular waves to anisotropic expansion. Furthermore, weak PE and PP can accelerate the spread of prey while strong and disproportionate intensities slow down the range expansion. The effects of PE and PP further enhance the population size, break down the spatial synchrony and promote the persistence of populations.     

Harvesting creates ecological traps: consequences of invisible mortality risks in predator-prey metacommunities

Models of two-patch predator-prey metacommunities are used to explore how the global predator population changes in response to additional mortality in one of the patches. This could describe the dynamics of a predator in an environment that includes a refuge area where that predator is protected and a spatially distinct ("risky") area where it is harvested. The predator's movement is based on its perceived fitness in the two patches, but the risk from the additional mortality is potentially undetectable; this often occurs when the mortality is from human harvesting or from a novel type of top predator. Increases in undetected mortality in the risky area can produce an abrupt collapse of either the refuge population or of the entire predator population when the mortality rate exceeds a threshold level. This is due to the attraction of the risky patch, which has abundant prey due to its high predator mortality. Extinction of the refuge predator population does not occur when the refuge patch has a higher maximum per capita predator growth rate than the exploited patch because the refuge is then more attractive when the predator is rare. The possibility of abrupt extinction of one or both patches from high densities in response to a small increase in harvest is often associated with alternative states. In such cases, large reductions in mortality may be needed to avoid extinction in a collapsing predator population, or to reestablish an extinct population. Our analysis provides a potential explanation for sudden collapses of harvested populations, and it argues for more consideration of adaptive movement in designing protected areas.     

Comparing spatial conservation prioritization methods with site- versus spatial dependency-based connectivity

Larval dispersal is an important component of marine reserve networks. Two conceptually different approaches to incorporate dispersal connectivity into spatial planning of these networks exist, and it is an open question as to when either is most appropriate. Candidate reserve sites can be selected individually based on local properties of connectivity or on a spatial dependency-based approach of selecting clusters of strongly connected habitat patches. The first acts on individual sites, whereas the second acts on linked pairs of sites. We used a combination of larval dispersal simulations representing different seascapes and case studies of biophysical larval dispersal models in the Coral Triangle region and the province of Southeast Sulawesi, Indonesia, to compare the performance of these 2 methods in the spatial planning software Marxan. We explored the reserve design performance implications of different dispersal distances and patterns based on the equilibrium settlement of larvae in protected and unprotected areas. We further assessed different assumptions about metapopulation contributions from unprotected areas, including the case of 100% depletion and more moderate scenarios. The spatial dependency method was suitable when dispersal was limited, a high proportion of the area of interest was substantially degraded, or the target amount of habitat protected was low. Conversely, when subpopulations were well connected, the 100% depletion was relaxed, or more habitat was protected, protecting individual sites with high scores in metrics of connectivity was a better strategy. Spatial dependency methods generally produced more spatially clustered solutions with more benefits inside than outside reserves compared with site-based methods. Therefore, spatial dependency methods potentially provide better results for ecological persistence objectives over enhancing fisheries objectives, and vice versa. Different spatial prioritization methods of using connectivity are appropriate for different contexts, depending on dispersal characteristics, unprotected area contributions, habitat protection targets, and specific management objectives. Comparación entre los métodos de priorización de la conservación espacial con sitio y la conectividad espacial basada en la dependencia     

Assessment of prey vulnerability through analysis of wolf movements and kill sites.

Within predator-prey systems behavior can heavily influence spatial dynamics, and accordingly, the theoretical study of how spatial dynamics relate to stability within these systems has a rich history. However, our understanding of these behaviors in large mammalian systems is poorly developed. To address the relationship between predator selection patterns, prey density, and prey vulnerability, we quantified selection patterns for two fine-scale behaviors of a recovering wolf (Canis lupus) population in Yellowstone National Park, Wyoming, USA. Wolf spatial data were collected between November and May from 1998-1999 until 2001-2002. Over four winters, 244 aerial locations, 522 ground-based telemetry locations, 1287 km of movement data from snow tracking, and the locations of 279 wolf kill sites were recorded. There was evidence that elk (Cervus elaphus) and bison (Bison bison) densities had a weak effect on the sites where wolves traveled and made kills. Wolf movements showed a strong selection for geothermal areas, meadows, and areas near various types of habitat edges. Proximity to edge and habitat class also had a strong influence on the locations where elk were most vulnerable to predation. There was little evidence that wolf kill sites differed from the places where wolves traveled, indicating that elk vulnerability influenced where wolves selected to travel. Our results indicate that elk are more vulnerable to wolves under certain conditions and that wolves are capable of selecting for these conditions. As such, vulnerability plays a central role in predator-prey behavioral games and can potentially impact the systems to which they relate.