Wolf Distribution Research Articles

Risk avoidance during dispersal: Temporal and behavioral shifts in selection by non-resident eastern wolves (Canis c.f. lycaon)

Dispersal is important for connectivity and persistence of small, disjunct wildlife populations. However, in human-altered landscapes dispersing animals are often more likely than residents to use anthropogenic features that increase mortality risk. As such, quantifying spatial and temporal responses of dispersing animals to anthropogenic features will improve our understanding of the potential for small, isolated populations to expand in human-altered landscapes. The distribution of eastern wolves (Canis c.f. lycaon; a threatened species in Canada) is limited to portions of eastern Canada within and adjacent to the population core in Algonquin Park. Recently, it was found that non-resident eastern wolves dispersing outside of Algonquin Park survive poorly, primarily due to high rates of human-caused mortality. We deployed Global Positioning System collars on 26 non-resident eastern wolves. We evaluated resource selection of dispersing individuals during different movement states to better understand their behavioral responses to anthropogenic landscape features during this critical, but risky period of their life history. Eastern wolves avoided areas with high building density during the day. However, during directed movements outside the winter period, eastern wolves selected areas closer to unpaved roads and areas with higher building densities at night. During winter, eastern wolves selected areas with high building densities at night while displaying tortuous movements. We demonstrate that harvested species may increase and relax avoidance of anthropogenic features at both daily and seasonal timescales. Failing to account for temporal and state-specific variation in resource selection patterns of dispersing animals may underestimate their ability to move through human-altered landscapes.

Wolf population density: prey biomass limits via intrinsic factors

Whether gray wolf Canis lupus population density is limited intrinsically or extrinsically has occupied researchers for decades. Since 1983, prey biomass, an extrinsic factor, has been accepted as limiting, but that conclusion was recently challenged for unusually high wolf densities which were then said to be limited by behavioral, or intrinsic, factors such as intraspecific strife, dispersal, etc., below prey‐biomass constraints. That contention has been disputed, so disagreement remains. This article discusses the models involved, presents revised models that shed new light, and offers evidence that supports the conclusion that at least at densities below 79 wolves 1000 km−2, wolf numbers per area are limited by prey biomass. As wolf density increases, behavioral factors limit the density to the number of wolves the prey biomass will support. Extra wolves disperse, and wolf range expands. Thus, while wolf density is self‐limiting, the population is not.

Wolves foraging on berries is likely a widespread behavior in southern boreal ecosystems

AbstractWolves are opportunistic generalists that can respond quickly to new and unique food sources. Wolves in some ecosystems will consume berries and other fruits when they are abundant and available; however, many aspects of this behavior remain unknown. In the Greater Voyageurs Ecosystem (GVE), Minnesota, USA, wolves consistently consume berries, particularly blueberries, when they are available. We deployed remote cameras in blueberry patches to record wolves foraging on berries over several years. We captured footage of wolves of all age classes, social statuses, and sex foraging on blueberries alone or with other wolves. Our observations indicate berry consumption by wolves is a widespread behavior in the GVE and likely in similar southern boreal ecosystems. We hope our work spurs researchers across wolf range to examine whether berry consumption by wolves is a widespread and ubiquitous behavior for wolves.

Changing the Tolerance of the Intolerant: Does Large Carnivore Policy Matter?

Success in large carnivore conservation often hinges on local residents' tolerance towards those species. Feelings of powerlessness and frustration with wildlife policies can lead to intolerance of the species. In extreme cases, intolerance may manifest in poaching. Thus, changes in policy may influence the tolerance of wildlife. To examine the connections between policy and tolerance, we examined how policy scenarios influenced anticipated changes in tolerance to wolves Canis lupus. We administered a survey in 2015-2016 in the core wolf range within northern Wisconsin, USA. Using hierarchical cluster analysis, we clustered respondents into groups based on their current tolerance of wolves. We evaluated the behavioral intentions of the clusters and examined the influence of policy scenarios on respondents' anticipated changes in tolerance. Finally, using an information-theoretic model selection framework, we assessed the effects of tolerance clusters and demographic factors. The respondents were clustered into three clusters relative to their current tolerance towards wolves: positive, ambivalent, and negative. Each cluster exhibited significantly different behavioral intentions and anticipated changes in tolerance for all scenarios. In all scenarios, respondents who already held positive attitudes towards wolves were significantly less likely to report expected changes in tolerance toward wolves following changes in wolf management. However, respondents who held ambivalent or negative attitudes towards wolves were significantly more likely to report expected changes in tolerance towards wolves following changes in wolf management. Regarding a regulated wolf hunting and trapping season, we observed a Simpson's Paradox, wherein, when examined in aggregate, no clear pattern emerged, but when examined at the cluster level, important and intuitive patterns emerged. Our demographic model results suggest that policy changes resulting in greater state management authority over wolves, especially authority to implement certain forms of legal killing of wolves, could result in significant increases in tolerance for individuals who identify as hunters, have lost livestock to a predator, or are currently ambivalent or negative towards wolves. Our work elucidates the nuanced relationship between tolerance of wildlife and wildlife policy and identifies a potential ecological fallacy.

When enough is enough: Optimising monitoring effort for large-scale wolf population size estimation in the Italian Alps.

The ongoing expansion of wolf (Canis lupus) populations in Europe has led to a growing demand for up-to-date abundance estimates. Non-invasive genetic sampling (NGS) is now widely used to monitor wolves, as it allows individual identification and abundance estimation without physically capturing individuals. However, NGS is resource-intensive, partly due to the elusive behaviour and wide distribution of wolves, as well as the cost of DNA analyses. Optimisation of sampling strategies is therefore a requirement for the long-term sustainability of wolf monitoring programs. Using data from the 2020-2021 Italian Alpine wolf monitoring, we investigate how (i) reducing the number of samples genotyped, (ii) reducing the number of transects, and (iii) reducing the number of repetitions of each search transect impacted spatial capture-recapture population size estimates. Our study revealed that a 25% reduction in the number of transects or, alternatively, a 50% reduction in the maximum number of repetitions yielded abundance estimates comparable to those obtained using the entire dataset. These modifications would result in a 2046 km reduction in total transect length and 19,628 km reduction in total distance searched. Further reducing the number of transects resulted in up to 15% lower and up to 17% less precise abundance estimates. Reducing only the number of genotyped samples led to higher (5%) and less precise (20%) abundance estimates. Randomly subsampling genotyped samples reduced the number of detections per individual, whereas subsampling search transects resulted in a less pronounced decrease in both the total number of detections and individuals detected. Our work shows how it is possible to optimise wolf monitoring by reducing search effort while maintaining the quality of abundance estimates, by adopting a modelling framework that uses a first survey dataset. We further provide general guidelines on how to optimise sampling effort when using spatial capture-recapture in large-scale monitoring programmes.

Home range and habitat selection of wolves recolonising central European human‐dominated landscapes

Decades of persecution has resulted in the long‐term absence of grey wolves Canis lupus from most European countries. However, recent changes in both legislation and public attitudes toward wolves has eased the pressure, allowing wolves to rapidly re‐establish territories in their previous central European habitats over the last 20 years. Unfortunately, these habitats are now heavily altered by humans. Understanding the spatial ecology of wolves in such highly modified environments is crucial, given the high potential for conflict and the need to reconcile their return with multiple human concerns. We equipped 20 wolves, originating from seven packs in six central European regions, with GPS collars, allowing us to calculate monthly average home range sizes for 14 of the animals of 213.3 km2 using autocorrelated kernel density estimation. We then used ESA WorldCover data to assess the mosaic of available habitats used within each home range. Our data confirmed a general seasonal pattern for breeding individuals, with smaller apparent home ranges during the reproduction phase, and no specific pattern for non‐breeders. Predictably, our wolves showed a general preference for remote areas, and especially forests, though some wolves within military training areas also showed a broader preference for grassland, possibly influenced by local land use and high availability of prey. Our results provide a comprehensive insight into the ecology of wolves during their re‐colonisation of central Europe. Though wolves are spreading relatively quickly across central European landscapes, their permanent reoccupation remains uncertain due to conflicts with the human population. To secure the restoration of European wolf populations, further robust biological data, including data on spatial ecology, will be needed to clearly identify any management implications.

Estimating distribution and abundance of wide-ranging species with integrated spatial models: Opportunities revealed by the first wolf assessment in south-central Italy.

Estimating demographic parameters for wide-ranging and elusive species living at low density is challenging, especially at the scale of an entire country. To produce wolf distribution and abundance estimates for the whole south-central portion of the Italian wolf population, we developed an integrated spatial model, based on the data collected during a 7-month sampling campaign in 2020-2021. Data collection comprised an extensive survey of wolf presence signs, and an intensive survey in 13 sampling areas, aimed at collecting non-invasive genetic samples (NGS). The model comprised (i) a single-season, multiple data-source, multi-event occupancy model and (ii) a spatially explicit capture-recapture model. The information about species' absence was used to inform local density estimates. We also performed a simulation-based assessment, to estimate the best conditions for optimizing sub-sampling and population modelling in the future. The integrated spatial model estimated that 74.2% of the study area in south-central Italy (95% CIs = 70.5% to 77.9%) was occupied by wolves, for a total extent of the wolf distribution of 108,534 km2 (95% CIs = 103,200 to 114,000). The estimate of total population size for the Apennine wolf population was of 2557 individuals (SD = 171.5; 95% CIs = 2127 to 2844). Simulations suggested that the integrated spatial model was associated with an average tendency to slightly underestimate population size. Also, the main contribution of the integrated approach was to increase precision in the abundance estimates, whereas it did not affect accuracy significantly. In the future, the area subject to NGS should be increased to at least 30%, while at least a similar proportion should be sampled for presence-absence data, to further improve the accuracy of population size estimates and avoid the risk of underestimation. This approach could be applied to other wide-ranging species and in other geographical areas, but specific a priori evaluations of model requirements and expected performance should be made.

Changes in Wolf Occupancy and Feeding Habits in the Northern Apennines: Results of Long-Term Predator-Prey Monitoring.

The comprehension of the factors that have influenced the recent changes in wolf (Canis lupus) range and diet that have occurred in our study area, characterized by a highly heterogeneous landscape, can shed light on their current process of expansion toward the plain. Wolf presence was monitored using a standardized protocol from 2007 to 2022 by carrying out eight monitoring sessions organized in seasonal surveys, during which, we collected wolf presence data. To model wolf range dynamics, we used dynamic occupancy models considering land cover types and wild ungulate abundances as covariates. Moreover, we studied the wolf diet through scat analysis, identifying the consumed items from undigested remains. Wolf occupancy in the study area progressed from mountains to lower hills gradually; the observed range dynamics were driven by prey abundance and human presence: in particular, the probability of colonization increased with roe deer (Capreolus capreolus) abundance, whereas the probability of extinction increased with urban areas. The wolf diet showed a gradual shift from the prevalent consumption of wild boar (2007-2008 and 2011-2012) to the prevalent consumption of roe deer (continuously increasing from 2015 onward). Our results might be related to a specific adaptation of the predator to the local ecology of the most consumed species: the roe deer.

The last and endangered maned wolves (Chrysocyon brachyurus) from southern Brazil and Uruguay: new records and spatial distribution modeling

ABSTRACTThe biggest canid in the neotropics, the maned wolf (Chrysocyon brachyurus), is endangered in many parts of its distribution. Only eight records have been reported for Uruguay and the southern border of Brazil over the last 30 years. After a multi-year effort, we present nine new records for this area and characterize Uruguay and southern Brazil’s habitat suitability. Our records confirm the current presence of the species in Uruguay, the Brazilian Pampa, and the Atlantic highlands of southern Brazil, although no local population could be identified. Individuals appear to move in an erratic pattern, possibly when searching for other members of the species. Only three regions presented multiple records (all in the Atlantic highlands), indicating the possible existence of local populations. Although most of southern Brazil and Uruguay are highly suitable for the species, none of the 21 surveyed areas (with 50,816 camera trap nights) within putative distribution areas have confirmed the species occurrence, as all records reported here came from opportunistic encounters. In light of this, we believe that very few individuals still occur in the southern limit of the maned wolf distribution, and thus, the species can certainly be classified as critically endangered in the Brazilian Pampa/Uruguayan Savanna.

Habitat selection of resident and non-resident gray wolves: implications for habitat connectivity

Habitat selection studies facilitate assessing and predicting species distributions and habitat connectivity, but habitat selection can vary temporally and among individuals, which is often ignored. We used GPS telemetry data from 96 Gray wolves (Canis lupus) in the western Great Lakes region of the USA to assess differences in habitat selection while wolves exhibited resident (territorial) or non-resident (dispersing or floating) movements and discuss implications for habitat connectivity. We used a step-selection function (SSF) to assess habitat selection by wolves exhibiting resident or non-resident movements, and modeled circuit connectivity throughout the western Great Lakes region. Wolves selected for natural land cover and against areas with high road densities, with no differences in selection among wolves when resident, dispersing, or floating. Similar habitat selection between resident and non-resident wolves may be due to similarity in environmental conditions, when non-resident movements occur largely within established wolf range rather than near the periphery or beyond the species range. Alternatively, non-resident wolves may travel through occupied territories because higher food availability or lower human disturbance outweighs risks posed by conspecifics. Finally, an absence of differences in habitat selection between resident and non-resident wolf movements may be due to other unknown reasons. We recommend considering context-dependency when evaluating differences in movements and habitat use between resident and non-resident individuals. Our results also provide independent validation of a previous species distribution model and connectivity analysis suggesting most potential wolf habitat in the western Great Lakes region is occupied, with limited connectivity to unoccupied habitat.

Preliminary Status, and distribution of Indian grey wolf (Canis lupus pallipes) in the Bhal region of Gujarat

Preliminary Status, and distribution of Indian grey wolf (Canis lupus pallipes) in the Bhal region of Gujarat

Genomic analysis of wolves from Pakistan clarifies boundaries among three divergent wolf lineages.

Among the three main divergent lineages of gray wolf (Canis lupus), the Holarctic lineage is the most widespread and best studied, particularly in North America and Europe. Less is known about Tibetan (also called Himalayan) and Indian wolf lineages in southern Asia, especially in areas surrounding Pakistan where all three lineages are thought to meet. Given the endangered status of the Indian wolf in neighboring India and unclear southwestern boundary of the Tibetan wolf range, we conducted mitochondrial and genome-wide sequencing of wolves from Pakistan and Kyrgyzstan. Sequences of the mitochondrial D-loop region of 81 wolves from Pakistan indicated contact zones between Holarctic and Indian lineages across the northern and western mountains of Pakistan. Reduced-representation genome sequencing of eight wolves indicated an east-to-west cline of Indian to Holarctic ancestry, consistent with a contact zone between these two lineages in Pakistan. The western boundary of the Tibetan lineage corresponded to the Ladakh region of India's Himalayas with a narrow zone of admixture spanning this boundary from the Karakoram Mountains of northern Pakistan into Ladakh, India. Our results highlight the conservation significance of Pakistan's wolf populations, especially the remaining populations in Sindh and Southern Punjab that represent the highly endangered Indian lineage.

The population bottleneck of the Iberian wolf impacted genetic diversity but not admixture with domestic dogs: A temporal genomic approach.

After decades of intense persecution, the Iberian wolf subspecies faced a severe bottleneck in the 1970s that considerably reduced its range and population size, nearly leading to its extinction in central and southern Iberian Peninsula. Such population decline could have impacted the genetic diversity of Iberian wolves through different processes, namely genetic drift and dynamics of hybridization with domestic dogs. By contrasting the genomes of 68 contemporary with 54 historical samples spanning the periods before and immediately after the 1970s bottleneck, we found evidence of its impact on genetic diversity and dynamics of wolf-dog hybridization. Our genome-wide assessment revealed that wolves and dogs form two well-differentiated genetic groups in Iberia and that hybridization rates did not increase during the bottleneck. However, an increased number of hybrid individuals was found over time during the population re-expansion, particularly at the edge of the wolf range. We estimated a low percentage of dog ancestry (~1.4%) in historical samples, suggesting that dog introgression was not a key driver for wolf extinction in central and southern Iberia. Our findings also unveil a significant decline in genetic diversity in contemporary samples, with the highest proportion of homozygous segments in the genome being recently inherited. Overall, our study provides unprecedented insight into the impact of a sharp decline on the Iberian wolf genome and refines our understanding of the ecological and evolutionary drivers of wolf-dog hybridization in the wild.

Environmental favorability models as a tool to support trophic rewilding initiatives

Trophic rewilding is considered a conservation measure that aims to restore the trophic interactions. In north‐central Portugal, the Iberian wolf (Canis lupus signatus) is endangered mainly due to human persecution, triggered by wolf depredation on livestock. Several initiatives have occurred in this area to increase wolf wild prey availability by reintroducing/reinforcing roe deer (Capreolus capreolus) populations. However, the success of trophic rewilding initiatives relies on up‐to‐date scientific evidence, considering both ecological and social settings. Favorability models can be used to infer the most suitable habitats for roe deer settlement and expansion by analyzing the environmental variables known to influence species ecology and behavior. In this study, we infer how the prey habitat favorability can contribute to plan prey populations reinforcement processes as a tool for trophic chain restoration. Using 330 camera traps, we monitored the roe deer population over 8,940 km2 in north‐central Portugal. Field data were used to evaluate the environmental determinants of roe deer presence. By using the favorability function, we created a spatially explicit gradient of habitat favorability that allowed us to define key areas for the application of management measures. The selected sites are strategically positioned to promote the natural expansion of established roe deer populations and overlap the current distribution of Iberian wolf. These results are an important step toward the restoration of the trophic chain and to reduce human–wolf conflicts. Our proposal paves the way for future rewilding initiatives, especially where prey species are central to the survival of top predators.

A shifting carnivore’s community: habitat modeling suggests increased overlap between the golden jackal and the Eurasian lynx in Europe

IntroductionThe European large carnivore community is recently showing positive trends. Beyond those, other species are expanding in Europe. In recent decades, the golden jackal (Canis aureus), a medium-sized canid, has rapidly expanded to western and northern Europe, after being historically confined to the Balkans. The Eurasian lynx (Lynx lynx), an apex predator once widely distributed throughout Europe, was nearly eradicated in the 19th century. Recent reintroductions, legal protection, and its natural expansion brought the lynx to recover, though some populations still face extinction. As the carnivores’ community across Europe is rearranging, ecological interactions are changing as well. Indeed, recent observations reported the golden jackal to kleptoparasitize the lynx. Our research aims to predict the potentially suitable habitats for the golden jackal and the Eurasian lynx, detailing the areas where those are expected to coexist soon.MethodsWe model their distribution in Europe for current and future scenarios, using both citizen-science and literature data, using the latter to buffer the possible biases of the former. For this aim, we included not only climatic variables, but also several biotic and abiotic predictors relevant to both species. Moreover, given his strong influence on the golden jackal, we compare our results with the predicted distribution of the grey wolf (Canis lupus).ResultsOur predictions show an increase in habitat suitability for the golden jackal, while an opposite trend is predicted for the Eurasian lynx. Also, we predict the target species to share a large portion of their range in the future, with most of these areas suitable for wolves, too.DiscussionOur results may have several impacts in the context of large-scale management, helping to predict further expansion of the golden jackal, and to identify critical areas for lynx conservation, supporting the management of possible reintroductions, also shedding light on the large carnivores’ changing community

Dog in sheep’s clothing: livestock depredation by free-ranging dogs may pose new challenges to wolf conservation

Livestock depredation is a common cause of human-carnivore conflicts. In Portugal, free-ranging dogs are increasingly abundant and overlap endangered Iberian wolf territories, with reports of livestock depredation. However, the lack of awareness about dogs’ possible role as predators leads to bias against wolves in cases of damages. Our goal was to assess and compare wolf and free-ranging dog’s diet composition at southern wolf range in Portugal, to offer insights on dogs’ predatory role on livestock and its implications for the conservation of an endangered wolf subpopulation. We assessed diet composition from 107 to 95 genetically confirmed wolf and dog scats, respectively, and complemented the analysis with data from 40 attacks on livestock with successful genetic predator assignment. Scat analysis highlighted goats as the most consumed dog prey in all analysed regions, with lagomorphs, small mammals, and wild boars as second most consumed in each region, respectively. Wolves mainly relied on goats and wild boars in the west, whereas in the central region they mostly fed on birds. The dietary overlap between both canids was very high (Pianka’s index O = 0.93), showing potential for competition. Additionally, we found that dogs were the sole predators detected in most attacks (62%). Our findings highlight dogs’ role as predators of livestock, and possibly also wild species, posing a further challenge to wolf conservation. Alongside adequate husbandry practices, we emphasise the need for a stronger enforcement of the legislation on dog ownership and an effective management of the stray population to reduce human-wolf conflict.

Insights into the dynamics of wolf occupancy in human-dominated landscapes

Among large carnivores, wolves show a remarkable capability to persist in human-dominated landscapes. However, the temporal dynamics of variation in spatial use of these landscapes remains poorly understood. Considering the relevance of spatio-temporal variations of territorial marking on wolf behaviour, either to defend territory boundaries and core areas or to expand into new areas, the location of wolf signs should reflect the dynamics of spatial use. Taking advantage of a long-term non-invasive wolf monitoring dataset spanning from 2005 to 2022 we fit a dynamic occupancy model to investigate the effects of environmental and anthropogenic factors on the dynamics of wolf spatial use in human-dominated landscapes. We focused on two dynamic parameters – colonization and extinction – and developed a wolf habitat suitability map for Iberia. Colonization probability increased with higher altitude, livestock density, and unpaved road density, and with the decrease of burned areas, national-regional, and local road densities. Extinction probability decreased with higher unpaved road density.In addition, we evaluated the wolf range dynamics in Iberia to understand if the ecological traits explained the expansion, stagnation or extinction sites observed since the beginning of the 2000s. Our results contribute to a sound understanding of wolf spatial use in human-dominated landscapes and its ability to adapt to these heterogeneous environments, allowing us to support adequate mitigation measures and conservation actions. The strong influence of livestock on the dynamics of wolf occupancy highlights the need to assess social factors, human dimensions, and direct wolf mortality causes for conflict management associated with livestock depredation.

Habitat Selection Differences of Two Sympatric Large Carnivores in the Southwestern Mountains of China

Large terrestrial carnivores play a crucial role in the top–down control of terrestrial ecosystems by maintaining ecosystem stability and biodiversity. However, intense interspecific competition typically occurs among large sympatric carnivores, leading to population reduction or extinction. Spatial partitioning through divergent habitat selection mitigates such competition. In this study, we analyzed the main environmental factors influencing the habitat selection and fragmentation of suitable habitats in Xinlong County, Sichuan Province, using 410 infrared cameras from 2015 to 2023. By employing generalized linear and maximum entropy models, we developed an ensemble model to predict the suitable habitat distribution of leopards (Panthera pardus) and wolves (Canis lupus). The results revealed significant disparities in suitable habitat distributions of leopards and wolves as coexisting large carnivores. Leopards prefer understory, whereas wolves prefer high-altitude meadows. Wolves spatially avoid leopards, who secure relatively superior resources and relegate wolves to inferior habitats. Although suitable habitat patches for both species cluster intensely, habitat connectivity remains low owing to pronounced anthropogenic disturbances, which is especially evident in the higher fragmentation of wolf habitats. These results suggest that sympatric large carnivores can reduce spatial competition intensity and promote spatial partitioning by selecting divergently suitable habitats, thereby facilitating species coexistence.

Gray wolf range in the western Great Lakes region under forecasted land use and climate change

AbstractLand use and climate change alter species distributions worldwide, and detecting and understanding how species ranges shift can facilitate conservation planning and action. Following extirpation from most of the contiguous United States, gray wolves (Canis lupus) have partially recolonized former range in the western Great Lakes region, but it is unknown how land use and climate change may alter amounts of wolf habitat. Using wolf observation data collected during winters 2017–2020 in Minnesota, Wisconsin, and Michigan, we created ensemble models to predict how land use and climate change may affect the amount of wolf habitat within these states. A projection model for the western Great Lakes region suggested three of four scenarios of land use and climate change will lead to 9%–35% increases in wolf habitat, while a solely climate‐based projection model supported our expectation that changes in climate, in isolation, will have limited effect on current wolf range. Our results support stable or increasing amounts of wolf habitat in the western Great Lakes region during the 21st century, suggesting limited or no adverse effects on the current distribution or further recolonization of wolves. Our findings can inform policy development regarding wolf conservation and identify areas where recolonization is plausible, thus where promoting human–wolf coexistence is most pertinent.

Вовк (Canis lupus) у степовій зоні України на зламі ХХ–ХХІ ст. та динаміка його чисельності

The article presents data on the extirpation of the wolf in southern Ukraine in the 18th to the 20th century, which was financially supported by landowners and the state. This contributed to keeping the wolf population at a low level and then led to a reduction in its range. In the 1970s, in the steppe and forest-steppe zones, wolves permanently lived only in Moldova; in the north of Odesa Oblast; in the Pavlohrad district of Dnipropetrovsk Oblast; in Kharkiv and Donetsk oblasts, as well as in the north of Poltava Oblast. From here, they spread to neighbouring territories, where they were quickly extirpated. The level of wolf decline was so significant, that in 1970 only 18 wolves were recorded in the steppe zone, and 270 in the whole of Ukraine. This contributed to the dispersal of the roe deer, wild boar, and elk to the south. After the reduction in the size of wolf hunting premiums, hunting pressure on wolves also decreased. Since there is a significant positive correlation between wolf population and hunting pressure (r = 0.93), the wolf's range and population began to grow rapidly. While in 1970 it was found in 14 regions, in 1976 it was already recorded in 18, and in 1981 in 21; in 2000, the wolf occurred in all regions of mainland Ukraine, and in 2003 it entered the Crimea. The process of restoring the wolf's range was accompanied by the emergence of wolf–dog hybrids, some of which are still occurring. Despite the harvest of 616.8 ± 102.68 wolves per year in 1970–1981, the wolf population increased by 6.5 times in Ukraine, and by more than 10 times in the steppe zone. A significant increase in its numbers was facilitated by a decrease in hunting pressure due to the adoption of the USSR Law ‘On the Protection and Use of Wildlife’ in 1980, which prohibited the use of inhumane methods in hunting on wild animals. In 1982–1992, the volume of wolf culling in Ukraine reached 71.1% (45.3–81.5), and in 1993–1999 was equal to 43.1% (34.9–49.3) per year. Naturally, due to the decrease in wolf hunting, there was a sharp increase in the wolf population, which in the steppe zone increased by 1.51 times from 2000 to 2010, and in our country in general by 1.03 times. The process of intensive growth of its population stabilised only at the beginning of the 21st century, when the annual take increased from 34.9% (2000) to 84.0% (2009).