Aspen recovery in northern Yellowstone: A comment on Brice et al. (2021).

Aspen sapling recruitment increased as browsing by elk decreased, following the 1995-96 reintroduction of wolves in Yellowstone National Park. We address claims by Brice et al. (2021) that previous studies exaggerated recent aspen recovery. We conclude that their results actually supported previous work showing a trophic cascade benefiting aspen.

Going back to "basics": Harlow's learning set task with wolves and dogs.

To survive and reproduce, animals need to behave adaptively by adjusting their behavior to their environment, with learning facilitating some of these processes. Dogs have become a go-to model species in comparative cognition studies, making our understanding of their learning skills paramount at multiple levels, not only with regards to basic research on their cognitive skills and the effects of domestication, but also with applied purposes such as training. In order to tackle these issues, we tested similarly raised wolves and dogs in a serial learning task inspired by Harlow's "learning set." In Phase 1, different pairs of objects were presented to the animals, one of which was baited while the other was not. Both species' performance gradually improved with each new set of objects, showing that they "learnt to learn," but no differences were found between the species in their learning speed. In Phase 2, once subjects had learned the association between one of the objects and the food reward, the contingencies were reversed and the previously unrewarded object of the same pair was now rewarded. Dogs' performance in this task seemed to be better than wolves', albeit only when considering just the first session of each reversal, suggesting that the dogs might be more flexible than wolves. Further research (possibly with the aid of refined methods such as computer-based tasks) would help ascertain whether these differences between wolves and dogs are persistent across different learning tasks.

Wolves alter the trajectory of forests by shaping the central place foraging behaviour of an ecosystem engineer.

Predators can directly and indirectly alter the foraging behaviour of prey through direct predation and the risk of predation, and in doing so, initiate indirect effects that influence myriad species and ecological processes. We describe how wolves indirectly alter the trajectory of forests by constraining the distance that beavers, a central place forager and prolific ecosystem engineer, forage from water. Specifically, we demonstrate that wolves wait in ambush and kill beavers on longer feeding trails than would be expected based on the spatio-temporal availability of beavers. This pattern is driven by temporal dynamics of beaver foraging: beavers make more foraging trips and spend more time on land per trip on longer feeding trails that extend farther from water. As a result, beavers are more vulnerable on longer feeding trails than shorter ones. Wolf predation appears to be a selective evolutionary pressure propelled by consumptive and non-consumptive mechanisms that constrain the distance from water beavers forage, which in turn limits the area of forest around wetlands, lakes and rivers beavers alter through foraging. Thus, wolves appear intricately linked to boreal forest dynamics by shaping beaver foraging behaviour, a form of natural disturbance that alters the successional and ecological states of forests.

Suid alphaherpesvirus 1 of wild boar origin as a recent source of Aujeszky's disease in carnivores in Germany.

BACKGROUND: The high susceptibility of carnivores to Suid Alphaherpesvirus 1 [SuAHV1, synonymous pseudorabies virus (PrV)], renders them inadvertent sentinels for the possible occurrence of Aujeszky's disease (AD) in domestic and wild swine populations. The aim of this study was to epidemiologically analyse the occurrence of PrV infections in domestic and wild animals in Germany during the last three decades and to genetically characterise the causative PrV isolates. METHODS: PrV in dogs was detected using standard virological techniques including conventional and real time PCR, virus isolation or by immunohistochemistry. Available PrV isolates were characterized by partial sequencing of the open gC reading frame and the genetic traits were compared with those of archived PrV isolates from carnivores and domestic pigs from Germany before the elimination of AD in the domestic pig population. RESULTS: During 1995 and 2022, a total of 38 cases of AD in carnivores, e.g. dogs and red foxes, were laboratory confirmed. Sequencing and subsequent phylogenetic analysis of PrV isolates established a strong connection between AD cases in carnivores and the occurrence of PrV infections in European wild boars in the end phase of and after elimination of AD from the domestic pig population. While PrV infections occur at low numbers but regularly in hunting dogs, interestingly, PrV was not observed in grey wolves in Germany. In none of 682 dead-found grey wolves and wolf-dog hybrids tested from Germany during 2006-2022 could PrV infection be detected by molecular means. CONCLUSIONS: Although PrV has been eliminated from domestic pigs, spillover infections in domestic and wild carnivores should always be expected given the endemic presence of PrV in wild pig populations. Since detection of PrV DNA and virus in carnivores is sporadic even in areas with high seroprevalence of PrV in wild pigs, it may not reflect the full diversity of PrV.

Effect of semantics in the study of tolerance for wolves.

As conservation scholars increasingly recognize the critical role of human thought and behavior in determining the persistence of biodiversity across the globe, a growing line of inquiry regarding the validity and comparability of previous applications of core psychological concepts has emerged. Specifically, inconsistent measurement and use of terms, such as attitudes and acceptance, reveal important questions about previous approaches. Given that these concepts differ by definition, yet have been used interchangeably, we explored what drives differences in people's responses when each concept is operationalized in the context of a contested wildlife species, the gray wolf (Canis lupus). To do so, we used data from a 2014 survey of U.S. residents (n = 1287) to test how measures of six concepts (i.e., acceptance, attitudes, benefits, risks, [prior] behavior, and behavioral intentions) often employed in the conservation social sciences were related with a broad set of possible explanatory variables. Despite moderate to strong correlations between all concepts measured (| Pearson's r | = 0.39-0.65, p < 0.001), results revealed considerable variation in their respective relationships with identical explanatory variables. Specifically, although wildlife value orientation (i.e., domination or mutualism) operated fairly consistently across cognitive and behavioral concepts, the relationship between the six concepts and other factors, such as social trust, identification with various interest groups (i.e., hunter, farmer, or rancher, environmentalist, and animal rights advocate), and political ideology (i.e., liberal vs. conservative), varied considerably. Our findings underscore that differences exist in these measures, which could have serious implications for conservationists integrating social science findings in their decision-making processes if they are unaware of the theoretical underpinnings of and distinctions between core psychological concepts.

Efectos de la semántica en los estudios de tolerancia a los lobos Resumen Los académicos dedicados a la conservación reconocen cada vez más lo importantes que son el pensamiento y el comportamiento humano para definir la persistencia de la biodiversidad a nivel mundial, por lo que ha emergido una creciente línea de indagación con respecto a la validez y la comparabilidad de las aplicaciones previas de conceptos psicológicos fundamentales. Más específicamente, las medidas incompatibles y el uso de términos como actitudes y aceptación revelan preguntas importantes sobre las estrategias anteriores. Ya que estos conceptos difieren por definición y aun así se han usado indistintamente, decidimos explorar qué impulsa las diferencias en las respuestas de las personas cuando cada concepto opera en el contexto de una especie de fauna controvertida: el lobo gris (Canis lupus). Para lograr esto, usamos datos de un censo de 2014 aplicado a residentes estadunidenses (n = 1,287) para analizar cómo la medida de seis conceptos usados frecuentemente en las ciencias sociales de la conservación (aceptación, actitudes, beneficios, riesgos comportamiento [previo] e intenciones conductuales) se relacionan con un amplio conjunto de variables explicativas posibles. A pesar de las correlaciones moderadas y fuertes entre todos los conceptos medidos (| Pearson's r | = 0.39 a 0.65, p < 0.001), los resultados revelaron una variación considerable en sus respectivas relaciones con las variables explicativas idénticas. De manera más precisa, aunque la orientación del valor de la fauna (es decir, dominancia y mutualismo) operó uniformemente en los conceptos cognitivos y conductuales, la relación entre los seis conceptos y otros factores, como la confianza social, identificación con varios grupos de interés (cazador, agricultor o ranchero, ambientalista, defensor de los derechos animales) e ideología política (liberal vs conservador) variaron considerablemente. Nuestros resultados destacan las diferencias que existen en estas medidas, las cuales podrían tener repercusiones serias para los conservacionistas que integran los resultados de las ciencias sociales dentro de sus procesos de toma de decisiones si no están concientes de las teorías fundamentales y las distinciones entre los conceptos psicológicos fundamentales.

Economical defence of resources structures territorial space use in a cooperative carnivore.

Ecologists have long sought to understand space use and mechanisms underlying patterns observed in nature. We developed an optimality landscape and mechanistic territory model to understand mechanisms driving space use and compared model predictions to empirical reality. We demonstrate our approach using grey wolves (Canis lupus). In the model, simulated animals selected territories to economically acquire resources by selecting patches with greatest value, accounting for benefits, costs and trade-offs of defending and using space on the optimality landscape. Our approach successfully predicted and explained first- and second-order space use of wolves, including the population's distribution, territories of individual packs, and influences of prey density, competitor density, human-caused mortality risk and seasonality. It accomplished this using simple behavioural rules and limited data to inform the optimality landscape. Results contribute evidence that economical territory selection is a mechanistic bridge between space use and animal distribution on the landscape. This approach and resulting gains in knowledge enable predicting effects of a wide range of environmental conditions, contributing to both basic ecological understanding of natural systems and conservation. We expect this approach will demonstrate applicability across diverse habitats and species, and that its foundation can help continue to advance understanding of spatial behaviour.

Integrating basic and applied research to estimate carnivore abundance.

A clear connection between basic research and applied management is often missing or difficult to discern. We present a case study of integration of basic research with applied management for estimating abundance of gray wolves (Canis lupus) in Montana, USA. Estimating wolf abundance is a key component of wolf management but is costly and time intensive as wolf populations continue to grow. We developed a multimodel approach using an occupancy model, mechanistic territory model, and empirical group size model to improve abundance estimates while reducing monitoring effort. Whereas field-based wolf counts generally rely on costly, difficult-to-collect monitoring data, especially for larger areas or population sizes, our approach efficiently uses readily available wolf observation data and introduces models focused on biological mechanisms underlying territorial and social behavior. In a three-part process, the occupancy model first estimates the extent of wolf distribution in Montana, based on environmental covariates and wolf observations. The spatially explicit mechanistic territory model predicts territory sizes using simple behavioral rules and data on prey resources, terrain ruggedness, and human density. Together, these models predict the number of packs. An empirical pack size model based on 14 years of data demonstrates that pack sizes are positively related to local densities of packs, and negatively related to terrain ruggedness, local mortalities, and intensity of harvest management. Total abundance estimates for given areas are derived by combining estimated numbers of packs and pack sizes. We estimated the Montana wolf population to be smallest in the first year of our study, with 91 packs and 654 wolves in 2007, followed by a population peak in 2011 with 1252 wolves. The population declined ~6% thereafter, coincident with implementation of legal harvest in Montana. Recent numbers have largely stabilized at an average of 191 packs and 1141 wolves from 2016 to 2020. This new approach accounts for biologically based, spatially explicit predictions of behavior to provide more accurate estimates of carnivore abundance at finer spatial scales. By integrating basic and applied research, our approach can therefore better inform decision-making and meet management needs.

Evolution of facial muscle anatomy in dogs.

Domestication shaped wolves into dogs and transformed both their behavior and their anatomy. Here we show that, in only 33,000 y, domestication transformed the facial muscle anatomy of dogs specifically for facial communication with humans. Based on dissections of dog and wolf heads, we show that the levator anguli oculi medialis, a muscle responsible for raising the inner eyebrow intensely, is uniformly present in dogs but not in wolves. Behavioral data, collected from dogs and wolves, show that dogs produce the eyebrow movement significantly more often and with higher intensity than wolves do, with highest-intensity movements produced exclusively by dogs. Interestingly, this movement increases paedomorphism and resembles an expression that humans produce when sad, so its production in dogs may trigger a nurturing response in humans. We hypothesize that dogs with expressive eyebrows had a selection advantage and that "puppy dog eyes" are the result of selection based on humans' preferences.

Critical Periods in Science and the Science of Critical Periods: Canine Behavior in America.

This article offers a canine history of the "critical period" concept, situating its emergence within a growing, interdisciplinary network of canine behavior studies that connected eugenically minded American veterinarians, behavioral geneticists, and dog lovers with large institutional benefactors. These studies established both logistical and conceptual foundations for large-scale science with dogs while establishing a lingering interdependence between American dog science and eugenics. The article emphasizes the importance of dogs as subjects of ethological study, particularly in the United States, where some of the earliest organized efforts to analyze canine behavior began. Further, the article argues that the "critical period" is important not only for its lasting prominence in multiple fields of scientific inquiry, but also as a historiographical tool, one that invites reflection on the tendency of historians to emphasize a particular narrative structure of scientific advancement.

Evidence of economical territory selection in a cooperative carnivore.

As an outcome of natural selection, animals are probably adapted to select territories economically by maximizing benefits and minimizing costs of territory ownership. Theory and empirical precedent indicate that a primary benefit of many territories is exclusive access to food resources, and primary costs of defending and using space are associated with competition, travel and mortality risk. A recently developed mechanistic model for economical territory selection provided numerous empirically testable predictions. We tested these predictions using location data from grey wolves (Canis lupus) in Montana, USA. As predicted, territories were smaller in areas with greater densities of prey, competitors and low-use roads, and for groups of greater size. Territory size increased before decreasing curvilinearly with greater terrain ruggedness and harvest mortalities. Our study provides evidence for the economical selection of territories as a causal mechanism underlying ecological patterns observed in a cooperative carnivore. Results demonstrate how a wide range of environmental and social conditions will influence economical behaviour and resulting space use. We expect similar responses would be observed in numerous territorial species. A mechanistic approach enables understanding how and why animals select particular territories. This knowledge can be used to enhance conservation efforts and more successfully predict effects of conservation actions.

Investigating the genetic and environmental architecture of interpack aggression in North American grey wolves.

Aggression confers several fitness benefits including increased breeding opportunities and resource acquisition. Determining the relative contributions of genetic and environmental components to shaping aggression is essential for advancing our understanding of how selection affects the distribution of aggressive phenotypes in a population. In a From the Cover article in this issue of Molecular Ecology, vonHoldt et al. (2020) used RAD-seq methods to obtain genome-wide single nucleotide polymorphism (SNP) data to estimate heritability of interpack aggression of 141 North American grey wolves (Canis lupus) surveyed from 1995-2018. The authors inferred heritability using both a SNP-based genetic relationship matrix (GRM) and a consensus pedigree informed by: (a) previously obtained microsatellite data; (b) past observations of parentage; and (c) statistical reconstruction of parent-offspring pairs. SNP-based (i.e., GRM) and pedigree-based (i.e., consensus pedigree) heritability estimates were 37% and 14%, respectively, with an additional 14%-16% explained by natal pack effects. The study confirmed the previously discovered strong effects of relative pack size and breeding status on interpack aggression, illustrating how social dynamics and density-dependent factors induce variance in aggressive behaviours. Finally, the authors found associations between average individual aggression scores (IAS) and specific candidate genes (MY09A and TRAK1). In sum, vonHoldt et al. (2020) provides an unprecedented and nuanced synthesis that not only suggests gene-aggression associations, but also emphasizes how additive genetic variance and density-dependent factors interact to maintain phenotypic variance in aggression over time.

Illuminating the mysteries of wolf history.

One of the most enduring surprises about the genetic history of Late Pleistocene populations is that continuity is often disturbed by upheaval. In fact, studies that support population continuity are increasingly rare in humans, a variety of vertebrate taxa, and vascular plants (Hofreiter & Stewart 2009; Burbrink et al. 2016). Perhaps such continuity should not be expected as the Pleistocene is marked by episodes of climate change, glaciation and the invasions of humans into previously isolated areas. Although fossils are one of the primary sources for inferring population continuity, a problem with fossil material is that, even if similar morphological forms might exist in a place over time, they may not be from the same genetic lineage. There are now readily available methods to assess genetic continuity solely from DNA found in fossil material, provided the record is fairly continuous. In a From the Cover article in this issue of Molecular Ecology, Loog et al. (2020) apply some of these readily available methods to analyse mitochondrial genomes and model the demography of wolves over the last 50,000 years.

Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia.

Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that have maintained a wide geographical distribution across the Northern Hemisphere throughout the Pleistocene and Holocene. Recent genetic studies have suggested that, despite this continuous presence, major demographic changes occurred in wolf populations between the Late Pleistocene and early Holocene, and that extant wolves trace their ancestry to a single Late Pleistocene population. Both the geographical origin of this ancestral population and how it became widespread remain unknown. Here, we used a spatially and temporally explicit modelling framework to analyse a data set of 90 modern and 45 ancient mitochondrial wolf genomes from across the Northern Hemisphere, spanning the last 50,000 years. Our results suggest that contemporary wolf populations trace their ancestry to an expansion from Beringia at the end of the Last Glacial Maximum, and that this process was most likely driven by Late Pleistocene ecological fluctuations that occurred across the Northern Hemisphere. This study provides direct ancient genetic evidence that long-range migration has played an important role in the population history of a large carnivore, and provides insight into how wolves survived the wave of megafaunal extinctions at the end of the last glaciation. Moreover, because Late Pleistocene grey wolves were the likely source from which all modern dogs trace their origins, the demographic history described in this study has fundamental implications for understanding the geographical origin of the dog.

Importance of a species' socioecology: Wolves outperform dogs in a conspecific cooperation task.

A number of domestication hypotheses suggest that dogs have acquired a more tolerant temperament than wolves, promoting cooperative interactions with humans and conspecifics. This selection process has been proposed to resemble the one responsible for our own greater cooperative inclinations in comparison with our closest living relatives. However, the socioecology of wolves and dogs, with the former relying more heavily on cooperative activities, predicts that at least with conspecifics, wolves should cooperate better than dogs. Here we tested similarly raised wolves and dogs in a cooperative string-pulling task with conspecifics and found that wolves outperformed dogs, despite comparable levels of interest in the task. Whereas wolves coordinated their actions so as to simultaneously pull the rope ends, leading to success, dogs pulled the ropes in alternate moments, thereby never succeeding. Indeed in dog dyads it was also less likely that both members simultaneously engaged in other manipulative behaviors on the apparatus. Different conflict-management strategies are likely responsible for these results, with dogs' avoidance of potential competition over the apparatus constraining their capacity to coordinate actions. Wolves, in contrast, did not hesitate to manipulate the ropes simultaneously, and once cooperation was initiated, rapidly learned to coordinate in more complex conditions as well. Social dynamics (rank and affiliation) played a key role in success rates. Results call those domestication hypotheses that suggest dogs evolved greater cooperative inclinations into question, and rather support the idea that dogs' and wolves' different social ecologies played a role in affecting their capacity for conspecific cooperation and communication.

The fox tapeworm, Echinococcus multilocularis, in grey wolves and dogs in Slovakia: epidemiology and genetic analysis.

Echinococcus multilocularis, the causative agent of human alveolar echinococcosis, is an important emerging parasite in the northern hemisphere. In epidemiological studies, the highest attention is being paid to foxes as the main reservoir hosts responsible for geographic expansion from multiple focal populations and the invasion of urban habitats, but little information is available on the parasite distribution in other carnivores. Hence, the study was designed to obtain updated information about the occurrence and genetic diversity of E. multilocularis in grey wolves and dogs in Slovakia. Faecal samples of wolves were collected from three locations under a certain level of environmental protection in the central and eastern parts of the country, and the presence of the parasite DNA was detected in 35.7% of 112 samples, with the highest rate (51.2%) recorded in the Poloniny National Park in north-eastern Slovakia. Among 110 faecal dog samples, E. multilocularis was detected in three faeces from segregated Roma settlements in the eastern part of the country, which accounted for an overall positivity of 2.7%. Sequence analysis of two mitochondrial genes, 12S rRNA and NADH dehydrogenase subunit 1, revealed four haplotypes in 13 isolates from wolves and dogs originating from four sites in eastern and central Slovakia, with all samples bearing a European-type pattern of E. multilocularis. The more than one-third positivity rate of E. multilocularis in wolf faecal samples dispersed over a large part of the country has corroborated the extensive circulation of the parasite in wildlife and confirmed the need to improve intervention control strategies.

Dogs and wolves do not differ in their inhibitory control abilities in a non-social test battery.

Being able to inhibit certain behaviours is of clear advantage in various situations. In particular, it has been suggested that inhibitory control plays a role in problem-solving and cooperation. Interspecific differences in the capacity for inhibitory control have been attributed to social and ecological factors, while one additional factor, namely domestication, has received only little attention so far. Dogs are an interesting species to test the effects of socio-ecological factors and also the influence of domestication on inhibitory control abilities. While dogs might have been selected for enhanced inhibition skills during domestication, the predictions derived from their socio-ecological background are reversed. Wolves are cooperative hunters and breeders, while dogs predominately scavenge and raise their young alone, accordingly, it would be predicted that dogs show impaired inhibitory control abilities since they no longer rely on these coordinated actions. To test these hypotheses, we assessed inhibitory control abilities in dogs and wolves raised and kept under similar conditions. Moreover, considering the problem of context-specificity in inhibitory control measures, we employed a multiple-test-approach. In line with previous studies, we found that the single inhibition tests did not correlate with each other. Using an exploratory approach, we found three components that explained the variation of behaviours across tests: motivation, flexibility, and perseveration. Interestingly, these inhibition components did not differ between dogs and wolves, which contradicts the predictions based on their socio-ecological backgrounds but also suggests that at least in tasks with minimal human influence, domestication did not affect dogs' inhibitory control abilities, thus raising questions in regard to the selection processes that might have affected inhibitory control abilities during the course of domestication.

Echinococcus spp. Tapeworms in North America.

Alveolar and cystic echinococcosis are emerging and reemerging in Europe, Africa, and Asia. The expansion of Echinococcus spp. tapeworms in wildlife host reservoirs appears to be driving this emergence in some areas. Recent studies suggest a similar phenomenon may be occurring in North America. We describe the context of Echinococcus spp. research in North America, with a specific focus on the contiguous United States. Although studies were conducted in the United States throughout the 1900s on various sylvatic and domestic Echinococcus spp. tapeworm cycles, data are lacking for the past ≈30 years. We review previous research, provide analysis of more recent focal studies, and suggest that Echinococcus spp. tapeworms, in particular E. canadensis, may be underrecognized. As a result, we suggest that additional research and surveillance be conducted for these tapeworms in wildlife host reservoirs across the United States.

Nowhere to hide: Effects of linear features on predator-prey dynamics in a large mammal system.

Rapid landscape alteration associated with human activity is currently challenging the evolved dynamical stability of many predator-prey systems by forcing species to behaviourally respond to novel environmental stimuli. In many forested systems, linear features (LFs) such as roads, pipelines and resource exploration lines (i.e. seismic lines) are a ubiquitous form of landscape alteration that have been implicated in altering predator-prey dynamics. One hypothesized effect is that LFs facilitate predator movement into and within prey refugia, thereby increasing predator-prey spatial overlap. We evaluated this hypothesis in a large mammal system, focusing on the interactions between boreal woodland caribou (Rangifer tarandus caribou) and their two main predators, wolves (Canis lupus) and black bears (Ursus americanus), during the calving season of caribou. In this system, LFs extend into and occur within peatlands (i.e. bogs and nutrient-poor fens), a habitat type highly used by caribou due to its refugia effects. Using resource selection analyses, we found that LFs increased predator selection of peatlands. Female caribou appeared to respond by avoiding LFs and areas with high LF density. However, in our study area, most caribou cannot completely avoid exposure to LFs and variation in female response had demographic effects. In particular, increasing proportional use of LFs by females negatively impacted survival of their neonate calves. Collectively, these results demonstrate how LFs can reduce the efficacy of prey refugia. Mitigating such effects will require limiting or restoring LFs within prey refugia, although the effectiveness of mitigation efforts will depend upon spatial scale, which in turn will be influenced by the life-history traits of predator and prey.

Harvest and group effects on pup survival in a cooperative breeder.

Recruitment in cooperative breeders can be negatively affected by changes in group size and composition. The majority of cooperative breeding studies have not evaluated human harvest; therefore, the effects of recurring annual harvest and group characteristics on survival of young are poorly understood. We evaluated how harvest and groups affect pup survival using genetic sampling and pedigrees for grey wolves in North America. We hypothesized that harvest reduces pup survival because of (i) reduced group size, (ii) increased breeder turnover and/or (iii) reduced number of female helpers. Alternatively, harvest may increase pup survival possibly due to increased per capita food availability or it could be compensatory with other forms of mortality. Harvest appeared to be additive because it reduced both pup survival and group size. In addition to harvest, turnover of breeding males and the presence of older, non-breeding males also reduced pup survival. Large groups and breeder stability increased pup survival when there was harvest, however. Inferences about the effect of harvest on recruitment require knowledge of harvest rate of young as well as the indirect effects associated with changes in group size and composition, as we show. The number of young harvested is a poor measure of the effect of harvest on recruitment in cooperative breeders.

Cross-jurisdictional management of a trophy-hunted species.

Gray wolves (Canis lupus) are managed for competing uses in the Greater Yellowstone Ecosystem (GYE). Tourism benefits Yellowstone National Park (YNP) visitors while trophy hunting benefits hunters outside of the park. We investigate the policy scope of gray wolf management across jurisdictional boundaries by incorporating three foundations of the behavioral ecology of wolves: refuge-seeking behavior, optimal foraging group size and territoriality. Tradeoffs between and within consumptive and non-consumptive human benefits and wolf population fitness and life history indicators are quantified as a set of elasticities, providing clear implications to resource managers. Our approach highlights that hunting intensity affects the provision of consumptive and non-consumptive human benefits across jurisdictional boundaries and ought to be managed accordingly. We also show that population levels are an incomplete indicator of species fitness, which may depend on how hunting policies impact underlying group ecology. Our findings suggest traditional optimization approaches to wildlife management may lead to suboptimal policy recommendations when the boundaries on the natural system are oversimplified. Highlighting the human element of wildlife management, we show that understanding tourist and hunter responses to wildlife population abundances is critical to balancing provision of consumptive and non-consumptive human uses.