JANUARY AND FEBRUARY 2026 JOURNAL ARTICLES

Abstract

Grey wolves (Canis lupus) rely on olfaction to communicate and acquire information about their environment. Olfactory signals delivered via urine, faeces, and glandular secretions, serve critical functions in reproduction, social interactions, and territorial defence. We studied the role of olfactory signals in mediating inter-pack conflict, a leading cause of mortality in free-ranging wolves, by exposing zoo-housed wolves to the scent of an unfamiliar individual, mimicking an intruder. We predicted breeding adults to exhibit higher investigatory and territorial behaviours, accompanied by an increased patrol effort in response to this simulated intruder than other individuals due to their dependence on territorial integrity and reproductive investment. Consistent with this prediction, breeders displayed significantly higher rates of investigatory, territorial and locomotor behaviours to conspecific than heterospecific urine than non-breeders. These findings highlight the specialised role that breeders play in wolf packs, particularly in relation to territorial defence, and demonstrate the deeply conserved patterns of territorial behaviours in wolves, even in populations that have been living under human care for a long time. Our findings have implications for theories of mammalian olfactory signalling and potential applications for conservation, including scent-based bio-fencing, to reduce human–wildlife conflicts.

Abstract

Gray wolf (Canis lupus) conservation has been particularly controversial in the Northern Rocky Mountains, although reliable human dimensions data is sparse. Globally, public cognitions toward wolves are consistently positive with exceptions among rural residents, especially hunters and landowners; results vary about whether or how cognitions change over time and time-series data remains scarce. Here, we describe wolf-related cognitions in the large, rural state of Montana using three, statewide mail-back surveys spanning a decade (2012–2023; total n = 7607). Results, weighted for inference, showed high and increasing public tolerance for wolves. Tolerance was low among wolf hunters and landowners, but increasing among ungulate hunters. Among residents, a decreasing majority tolerated wolf hunting, whereas a decreasing minority tolerated wolf trapping. Tolerance of wolves was spatially correlated toward population centers and associated with respondents’ identities, values, and wolf-related experiences. We discuss these results, future work needed to explain these trends, and implications for wolf management broadly.

ABSTRACTn
Background

Understanding mortality is crucial to predict the dynamics of animal populations, especially for long-lived species subjected to widespread human pressure. This is the case for the grey wolf (Canis lupus), yet local studies provide an incomplete view of the patterns and determinants of this process.
Aims and Methods

We conducted a systematic review of the existing literature on grey wolf mortality across the species’ distribution range (i.e., most of the Northern Hemisphere) to identify global patterns and potential differences across continents. Studies were analysed based on the type of information provided, including mortality rates, proportion of dead individuals by cause, and natural and human-related determinants of mortality.
Results and Discussion

We found high global minimum annual mortality rates in wolves (0.31 ± 0.18), compared to those reported under low human persecution (< 0.2). Most deaths were linked to human activities, primarily deliberate (legal and illegal) killing (60% ± 25%). Europe reported a higher percentage of human-related deaths (86% ± 14%) than North America (66% ± 24%). This is likely due to higher anthropogenic pressures in Europe and differences in carcass detection methods between continents (opportunistic recovery in Europe vs. tracking of collared wolves in North America). However, the lack of data from areas with low human influence (e.g., the Arctic and deserts) limits our understanding of the natural dynamics of mortality in the species. We found that many individual, social and environmental variables determine mortality values, and our global perspective allowed us to disentangle long-standing debates in large carnivore ecology, including support for the hypothesis that human-caused and natural mortality are additive.

ABSTRACT

Species recovery can be influenced by a wide variety of factors, such that predicting the spatiotemporal dynamics of recovering species can be exceedingly difficult. These predictions, however, are valuable for decision makers tasked with managing species and determining their legal status. We applied a spatially explicit projection model to estimate population growth and viability of gray wolves (Canis lupus) from 2021 to 2070 in the state of Washington, USA, where wolves have been naturally recolonizing since the establishment of the first resident pack in 2008. Using this model, we predicted the effects of 12 scenarios relating to management actions (e.g., lethal removals by the state agency, translocation, recreational harvest) and system uncertainties (e.g., immigration from out of state, disease) on the probability of meeting Washington’s wolf recovery goals, the probability of extinction, and other metrics related to population status. Population recovery was defined under Washington’s Wolf Conservation and Management Plan as four breeding pairs in each of three recovery regions and three additional breeding pairs anywhere in the state. The baseline and two translocation scenarios indicated a high (> 90%) probability of wolf recovery in Washington by 2070, but scenarios related to harvest mortality (removal of 5% of the population every 6 months), increased lethal removals (removal of 8.53% of the population across the state each year), and cessation of immigration from out of state resulted in probabilities of < 0.20 (0.01, 0.04, and 0.17, respectively) of meeting recovery goals by 2070. Only two scenarios of 12 (increased harvest and lethal removals scenarios) resulted in a geometric mean of population growth ≤ 1, indicating long-term population stability or growth for most scenarios. Our results suggest that wolves will continue to recolonize Washington and that recovery goals are likely to be met so long as harvest and lethal removals are not at unsustainable levels and adjacent populations support immigration into Washington.

Abstract

The recent expansion of grey wolves (Canis lupus) in Italy has led them into new ecological contexts, including areas characterized by poor prey communities and landscape anthropization. While dietary studies are essential for predicting wolf’s ecological functions and impacts, it remains unclear whether research on wolf diet has kept pace with this expansion. We mapped the current distribution of wolves in Italy and then clusterized areas based on food resources available to wolves: domestic or wild ungulates, the coypu (Myocastor coypus), and resources associated with landscape anthropization. Finally, we checked the coverage of each cluster by dietary studies (n = 33). Areas currently inhabited by wolves in Italy include nine different food resource assemblages. However, most studies on wolf diet have focused on remote areas of the Alps, where northern Chamois and red deer are abundant, and in areas with a rich assemblage of wild ungulates. In contrast, wolf diet remains unexplored in Mediterranean ecosystems with poorer ungulate assemblages and in highly anthropized landscapes, despite these environments together accounting for most areas of ongoing wolf expansion. These gaps may preventing us from knowing if wolves in highly anthropized landscapes are exposed to toxic compounds and if predation on domestic pets can trigger conflicts. Similarly, the lack of research in areas where wolves rely almost exclusively on wild Boar hinders our ability to predict the potential impact of African Swine Fever on wolf ecology and behaviour, as well as its broader influence on human-wolf conflicts.

Abstract
Wolf-dog hybridisation (WDH) – a form of anthropogenic introgressive hybridisation – may have occasionally occurred since the times of domestication. More recently, however, despite the ongoing recovery of wolf populations in Europe, the high number of domestic dogs along with marginal ecological conditions may pose increased risks to their genetic integrity. Accordingly, WDH is currently recognised among the highest threats to European wolves, with potential cascading effects through socio-ecological systems. Based on high quality DNA from tissues of 748 wolves retrieved dead from 2020 to 2024, alongside 26 additional samples from 1993 to 2003, we assessed WDH in the peninsular Italian wolf population. Using 23 autosomal STR loci and 5 Y-linked loci for males, we detected a high proportion (46.7 %) of wolves admixed with dogs, 29.5 % of which recent hybrids and 17.2 % older backcross generations. While most of the original hybridisation events likely took place some 9–16 years ago, a few first-generation hybrids in our sample indicate that WDH is currently ongoing. This seriously threatens the genetic integrity not only of the Italian wolf population, but also of the neighbouring wolf populations through dispersal events. The situation we present, which has not been previously reported for any wolf population worldwide, serves as a cautionary example of the potential dynamics of WDH in human-dominated landscapes should the issue remain persistently unaddressed. They also highlight the importance of considering the genetic constituency of wolf populations when evaluating their conservation status, especially in light of the recent downgrading of their protection status across Europe.

18 p.

of funds for wall construction, they are repeated and erroneously upheld the Secretary of Homeland Security’s waiver authority. This authority allowed border wall construction to proceed in violation of Federal, state, and local laws – actions Fitzgerald exceeded claimsed limits and violad the nondelegation doctrine. providing comprehensive and detailed examination of the legal challenges the Endangered Species Act and the recovery of the Mexican gray wolf, this book will be of interest to legal scholars, policymakers, and conservation advocates. [Publisher’s text]

Abstract
Human intolerance is a critical factor limiting both the distributions and populations of large carnivores. Using gray wolves as a case study, we synthesize a half-century of scholarship with the aims of clarifying the conceptual foundations of “tolerance” and integrating insights from across the social sciences. Specifically, we review longitudinal studies of attitudes toward wolves and show how trends vary across the populations examined. We then identify and discuss three complementary theories that help explain variation in tolerance across individuals, social groups, and societies: (1) Risk–benefit theories illuminate how perceptions of risks, benefits, and controllability shape individuals’ tolerance of carnivores; (2) Modernization theory explains societal shifts in values and shows how reduced threats from carnivores impact tolerance at the societal level; and (3) Social Identity Theory highlights how identification with interest groups (e.g., hunters, environmentalists) shape beliefs in a manner that serves to exacerbate inter-group conflicts. Linking these theoretical perspectives provides a more holistic framework for understanding why tolerance can change within populations, and why inter-group conflicts persist even as societal attitudes have become more favorable. We conclude by outlining research priorities aimed at improving our understanding of tolerance and the conditions that allow for human–carnivore coexistence.

Abstract
Background/Objectives: Following centuries of systematic eradication, grey wolf (Canis lupus) populations across Europe have experienced a significant recovery over recent decades, which leads to concerns regarding, among others, anthropogenic hybridization. In Greece, the genetic status of the wolf population is largely unknown to date. Here, we genetically monitor and test for wolf–dog hybridization events in a recently established wolf population in the Parnitha Protected Area, in close vicinity to the capital city of Greece. Methods: One hundred and twenty-four wolf scat samples were genotyped at 20 canine-specific autosomal microsatellite loci and compared to available reference tissue samples from wolves and free-ranging dogs. Results: A minimum of 31 unique wolf individuals were identified, structured into at least three packs. No wolf–dog hybrids were detected in the study area. To validate the accuracy of the microsatellite analysis, an ancestry informative 93-SNP panel was applied to non-invasive wolf DNA samples from the study area, confirming the absence of hybrids among them. However, a possible wolf–dog hybrid was detected among reference wolf samples collected in Northern Greece, where individuals with atypical morphological traits are observed. The estimated census population size was in accordance with concurrently obtained camera trapping data, while heterozygosity values were low. Conclusions: This research represents the first systematic effort in Greece to genetically monitor wolves recently established in a protected area. It highlights the need for targeted management strategies based on genetic data to ensure balanced long-term conservation of wolves in peri-urban areas.

Abstract

Caribou (Rangifer tarandus) calf mortality during the neonatal period is commonly attributed to predation, particularly by gray wolves (Canis lupus). However, neonate mortality remains understudied in mountain caribou, despite increasing wolf reduction programs. We used an individual-based movement method to infer parturition and neonate mortality from adult female telemetry data (78 individual-years), supplemented with 3 years of camera trap data (89 cameras), to examine changes in neonate mortality rates, timing, and locations before (2012–2014) and after (2020–2021) wolf reduction, across 2 calving areas (one rugged, another gradual) for the Itcha-Ilgachuz subpopulation in British Columbia, Canada. Given the likely difference in the timing of wolf–neonate overlap between calving areas, we hypothesized that wolf predation would be additive to other mortality sources typically affecting younger neonates (e.g., grizzly bear [Ursus arctos] predation) in the rugged area, but compensatory in the gradual area. Accordingly, we predicted that reducing wolves would increase survival and lower the average mortality age for neonates in the rugged area, with smaller gains in survival and minimal change in average mortality age in the gradual area. After wolf reduction, survival increased 41% in the rugged area but did not improve in the gradual area, resulting in no overall increase in survival at the subpopulation level. Average mortality age decreased in the rugged area from approximately 14 days to 8 days, coinciding with the peak in camera detections of grizzlies and wolverines (Gulo gulo), and remained at approximately 9 days in the gradual area. Mortalities before wolf reduction (i.e., those more likely caused by wolves) were more strongly associated with anthropogenic linear features and treed valley bottoms. Our findings highlight the value of considering habitat-specific mechanisms influencing calf mortality and integrating indirect approaches to address knowledge gaps in prey–predator dynamics.

Abstract

Wolf pup Canis lupus survival is a key driver of wolf population dynamics that remains poorly understood, especially in forested systems, because wolf pups are difficult to monitor. We used a combination of pup counts at dens and remote camera observations to estimate annual survival and recruitment of wolf pups in the Greater Voyageurs Ecosystem, MN, USA, from 2019 to 2025. We estimated recruitment for 33 packs over 92 pack-years and survival for 23 litters from 13 packs. Mean annual pup recruitment was 1.27 pups per pack, and mean annual pup survival was 0.29. Annual wolf pup recruitment and survival rates were highly variable among years and packs, which is likely a result of differences in food availability and the ability of breeding animals to acquire sufficient prey to provision dependent pups. Pup survival was negatively related to litter size. Although most (71%) wolf pups born during our study did not survive their first biological year, the population remained relatively stable, suggesting that recruitment rates were sufficient to sustain the high-density wolf population over time. Our work underscores the potential of integrative monitoring approaches to advance the understanding of wolf reproductive ecology.

Abstract
The future of biodiversity conservation depends on balancing the needs of people and wildlife through collaborative, equitable approaches. While protected areas are essential, they often carry legacies of exclusion and injustice. Around the world, efforts to improve collaboration in protected areas vary in inclusivity and power-sharing—especially in cross-cultural contexts where shared decision-making is not legally mandated. Pacific Rim National Park Reserve’s (PRNPR) Wild About Wolves project exemplifies the promises and tensions of government-led collaboration. The initiative aimed to foster regional cooperation, research, and action around growing human–wolf conflict. We interviewed 32 individuals—including PRNPR staff, Nuu-chah-nulth citizens, local community members, and organization representatives—with current or past involvement in the project. Participants identified several challenges: cultural differences in views on wolf management, unclear project timelines and expectations, and concerns about managerial priorities. Yet they also highlighted opportunities: strong regional conservation values, the depth of Nuu-chah-nulth and local knowledge, and PRNPR’s growing capacity for collaboration. From these insights, we co-produced a framework to improve PRNPR-led collaboration, emphasizing institutional clarity, alignment of social and cultural priorities, and more durable models of leadership and governance. While grounded in PRNPR, the lessons offer valuable guidance for collaborative conservation worldwide.

Abstract

Reproductive output can vary widely among mammalian species. There are many drivers that affect reproductive output including evolutionary, environmental, population, social, and individual traits. Although several factors, including human-caused mortality, can affect reproductive output, we generally have a poor understanding of how such factors interact to affect reproduction, particularly in cooperative breeders. Gray wolves (Canis lupus) in Idaho, USA, are exposed to annual hunting and trapping. Thus, they are an ideal species to answer questions about how turnover within groups affects reproduction in cooperative breeders. I hypothesized that the reproductive output of wolves would be affected by individual, social, and environmental factors. Contrary to my prediction, mid-summer litter size was positively associated with wolf harvest density, suggesting a compensatory response to harvest in cooperatively breeding gray wolves. Such compensation is only partial, however, and does not fully account for all the individuals lost from harvest. At the very highest harvest densities observed, mean litter size increased nearly 28%. In contrast, mid-summer litter size was negatively associated with multiple breeding in groups, suggesting resource limitation and competition within groups. I show that characteristics associated with harvest and breeding strategies predict variations in litter size in a cooperative breeder.

Abstract

Wolves, once one of the most widely distributed carnivores on Earth, have experienced drastic population declines and range contractions due to anthropogenic pressures. We studied the population status and habitat utilization of the Indian Grey Wolf in Pench Tiger Reserve (PTR). Both direct and indirect evidence were used to assess wolf presence, and Bonferroni confidence intervals were calculated to analyze habitat utilization. Wolf distribution was found to be restricted to the Khawasa Range of PTR, with the Khawasa Beat showing the highest encounter rate (0.154/km). Most wolf signs were recorded in mixed habitats (miscellaneous), underscoring the importance of these areas for the species’ long-term conservation. A significant positive correlation was observed between tree-cutting and wolf encounter rate (r = 0.976, p = 0.024, df = 3), suggesting that habitat changes associated with tree removal may influence wolf activity.