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Why do wolves eat livestock?
... In other words, a dispersal strategy punctuated by breeding events can occur only if the ecological requirements typical of stable wolf packs are met in some areas of ecological corridors. Wolf requirements in areas occupied by stable packs have been extensively studied, both in natural and in human-modified landscapes, and involve (i) good prey availability, and (ii) shelter areas that can minimize the risk of encounters with humans (Llaneza et al., 2012;Imbert et al., 2016). ...
... Wild boar and roe deer occurrence data were collected during the systematic monitoring along routes in 2017 and 2018. Relative abundance was estimated at the route level, by calculating the Index of Kilometric Abundance (IKA) (number of detected signs divided by the route length, Meriggi et al., 2015;Imbert et al., 2016) for each species. IKA values used in this study were obtained by averaging the values of the IKA associated with winter and spring monitoring periods (for details see Dondina et al., 2022). ...
... In the Ticino Natural Park, wolves have actively selected areas with low anthropogenic disturbance (i.e., away from roads and with low arable land cover), shelter accessibility (areas with high shrub cover), and high prey availability (positive relationship with roe deer IKA). These preferences match the resource selection behavior of resident wolves, which typically settle in areas with low anthropic disturbance and high prey abundance Imbert et al., 2016;Rio-Maior et al., 2019;Meriggi et al., 2020). In particular, these findings align with the expansion pattern of the species observed in the northern Apennines (Lombardy), where the colonization, which occurred from the mountains to the hills bordering the Po Plain, has been driven by roe deer abundance . ...
A dispersal strategy punctuated by breeding events along corridors is the most effective for sustaining recolo-nization and viable metapopulations of large mammals. Consequently, functional ecological corridors not only have to offer dispersal routes, but they must also provide suitable breeding conditions. This study investigated wolf spatio-temporal behavior within the Ticino Natural Park, an ecological corridor crossing a highly human-modified area in northern Italy. Wolves were systematically monitored from 2017 to 2023. Kernel and resource selection analyses showed that core areas characterized by high naturalness attracted transient individuals due to their suitability for potential settlement. Wolves selected core areas with low anthropogenic disturbance, high shrub cover, and prey abundance, aligning with behaviors observed in resident wolves. Despite the availability of favorable habitat conditions and the nocturnal behavior adopted by wolves, anthropogenic disturbance, including culling activities, human-induced mortality, and traffic roads, have been the primary limitation to pack establishment in the park so far. However, the observations of a stable wolf pair in the most recent core area in 2023-2024 and the consequent implementation of conservation management actions by park managers have permitted the very recent settlement of the first wolf pack within the protected area. This event will reinforce the ecological connectivity between Apennines and Alps wolf sub-populations and between the Italian and European populations. This study underscores the importance of actively protecting natural corridors able to support both dispersal and settlement to ensure recolonization of formerly occupied areas and the long-term large carnivore conservation in a landscape of coexistence.
... In other words, a dispersal strategy punctuated by breeding events can occur only if the ecological requirements typical of stable wolf packs are met in some areas of ecological corridors. Wolf requirements in areas occupied by stable packs have been extensively studied, both in natural and in human-modified landscapes, and involve (i) good prey availability, and (ii) shelter areas that can minimize the risk of encounters with humans (Llaneza et al., 2012;Imbert et al., 2016). ...
... Wild boar and roe deer occurrence data were collected during the systematic monitoring along routes in 2017 and 2018. Relative abundance was estimated at the route level, by calculating the Index of Kilometric Abundance (IKA) (number of detected signs divided by the route length, Meriggi et al., 2015;Imbert et al., 2016) for each species. IKA values used in this study were obtained by averaging the values of the IKA associated with winter and spring monitoring periods (for details see Dondina et al., 2022). ...
... In the Ticino Natural Park, wolves have actively selected areas with low anthropogenic disturbance (i.e., away from roads and with low arable land cover), shelter accessibility (areas with high shrub cover), and high prey availability (positive relationship with roe deer IKA). These preferences match the resource selection behavior of resident wolves, which typically settle in areas with low anthropic disturbance and high prey abundance Imbert et al., 2016;Rio-Maior et al., 2019;Meriggi et al., 2020). In particular, these findings align with the expansion pattern of the species observed in the northern Apennines (Lombardy), where the colonization, which occurred from the mountains to the hills bordering the Po Plain, has been driven by roe deer abundance . ...
... Their role as an ecosystem regulator through trophic cascades has been well documented (Ripple et al., 2001;Ripple & Beschta, 2004;Halofsky & Ripple, 2008). Wolves were once hunted to near extinction, however, over a period of time, increased public awareness towards the species has help garnering strong legal protection, favorable media coverage and furthered ecological research (Mech, 1995;Ripple & Beschta, 2007;Chapron, Andren & Liberg, 2008a;Imbert et al., 2016). Yet, persecution remains as one of the biggest obstacles to wolf recovery around the world (Newsome et al., 2016), including the Himalayan wolf. ...
... Wolves are known to choose their prey based on wild prey abundance, vulnerability, pack stability, dispersal nature, habitat accessibility as well as husbandry regime in human-dominated landscapes (Imbert et al., 2016). Patterns of wolf diet across these regions show that wolves consumed mostly large and domestic prey (40 % and 48.21% respectively Consequences of conflict due to prey un-availability can also hinder key conservation goals. ...
... Their role as an ecosystem regulator through trophic cascades has been well documented (Ripple et al., 2001;Ripple & Beschta, 2004;Halofsky & Ripple, 2008). Wolves were once hunted to near extinction; however, over a period of time, increased public awareness toward the species has help garnering strong legal protection, favorable media coverage and furthered ecological research (Mech, 1995;Ripple & Beschta, 2007;Chapron, Andren & Liberg, 2008;Imbert et al., 2016). Yet, persecution remains as one of the biggest obstacles to wolf recovery around the world (Newsome et al., 2016), including the Himalayan wolf. ...
... Consequently, wolf predation on livestock can lead to social conflicts between conservationists, farmers and other stakeholder groups (Bautista et al., 2019). Livestock damage is often restricted to few farms (Gazzola et al., 2008), and depends on landscape structure and availability of natural prey (Treves et al., 2004;Suryawanshi et al., 2013;Imbert et al., 2016). Additionally, the social status or family history of individual wolves can affect livestock depredation rates. ...
... Additionally, the social status or family history of individual wolves can affect livestock depredation rates. For example, in Northern Italy dispersing wolves killed more livestock compared to resident pairs and packs (Imbert et al., 2016). Sometimes single individuals or packs are responsible for disproportionately high livestock damage, which often results in public pressure for such "problem individuals" to be culled. ...
... Moreover, we cannot exclude the possibility that lack of familiarity with an area might influence the propensity of non-residents to attack livestock, as shown in other areas and species (Mizutani, 1993;Linnell et al., 1999). Independent of the proximate mechanism, increased predation on livestock in areas with reduced availability of natural prey and increased availability of livestock was also shown in other areas (Suryawanshi et al., 2013;Imbert et al., 2016). We have no clear explanation why livestock depredation rates by resident wolves were higher during winter and lower during summer, but it might relate to prey switching related to the presence of dependent offspring. ...
Attacks by large predators on livestock are an important driver of conflicts. Consequently, knowledge about where predators occur, where livestock depredation takes place and what factors influence it will aid the mitigation of stakeholder conflicts. Following legal protection, wolves (Canis lupus) in Central Europe are recently spreading to areas dominated by agriculture, bringing them in closer contact with livestock. Here, we analyzed habitat selection and livestock depredation rates of 43 wolves identified by genotyping on the Jutland peninsula, consisting of mainland Denmark and the northernmost German federal state Schleswig-Holstein. Occupancy by resident wolves correlated positively with forest and other non-forested semi-natural land cover (habitat for natural ungulate prey), whereas occupancy by non-resident wolves correlated with increasing forest cover and sheep density. The latter effect likely reflected increased sampling probability of highly mobile dispersers killing livestock. We recorded 565 livestock depredation events (85 in Denmark and 480 in Schleswig-Holstein), of which 42% (55 in DK and 185 in SH) could be assigned to 27 individual wolves based on DNA evidence. Livestock (mostly sheep) were killed by wolves in 16% of the study area. Our results indicate that wolves mostly killed livestock as a context-dependent response, i.e., being dispersers in agricultural areas with low availability of wild ungulate prey and high livestock densities, and not because of behavioral preferences for sheep. Moreover, the livestock depredation was lower in areas with livestock protection measures (implemented in areas with established pairs/packs). We conclude that while wolf attacks on livestock in established wolf territories generally can be reduced through improvement of fences, livestock depredation by non-resident wolves in agricultural areas constitutes a bigger challenge. Albeit technically possible, the economic costs of implementing predator-proof fences and other preventive measures in such pastoral areas infrequently visited by wolves will be considerable. Experiences so far further indicate that lethal removal of identified “problem wolves” may be inefficient in practice.
... Therefore, whether a parent pack kills sheep may predict what a loner would prefer and predict whether the loner would become problematic towards sheep. This prediction differs from the common assumption that sheep are easy prey [8,[34][35][36]. Similarly, whether loners will be in proximity to a human environment may also relate to a habit of their parent packs. ...
... These two wolves could be the tip of an iceberg and representatives of a larger subpopulation of unobserved wolves: wolves that did enter the Netherlands, but did not kill farm animals and went unobserved. If so, it is remarkable that a subpopulation of loners enters and crosses the Netherlands without killing farm animals; sheep in particular, as sheep are considered easy prey [8,[34][35][36]. In this respect, there are three significant observations. ...
... Thus, the availability of sheep cannot explain why there are loners that do not kill sheep. Other researchers also showed that prey preference cannot be fully explained by availability [8,35,[101][102][103][104][105][106]. Therefore, prey preference is apparently not merely a matter of stochastic laws. ...
Simple Summary
Conflicts with wolves arise because wolves kill farm animals, especially sheep, or approach humans. It is expected that young wolves learn from their parent pack (PP) what their prey is and if it is safe to be near humans. To confirm this, we researched whether the behavior of young migrating wolves (loners), after they leave the pack, resembles PP behavior. Fourteen loners entering the Netherlands between 2015 and 2019 could be identified and genetically linked to their PPs. Loner and PP behavior was similar in 10 out of 14 cases. Like their PPs, some young wolves killed sheep and were near humans, others killed sheep and did not approach humans, while two loners were unproblematic, they did not kill sheep nor were they in proximity to humans. Thus, the PP behavior did predict loner’s behavior and conflicts may be similar between young wolves and their PPs. However, conflicts need not arise. To achieve that, new prevention methods are proposed to teach wolves in the PP not to approach sheep and humans. As a result, new generations may not be problematic when leaving the PP.
Abstract
Transmission of experience about prey and habitat supports the survival of next generation of wolves. Thus, the parent pack (PP) can affect whether young migrating wolves (loners) kill farm animals or choose to be in human environments, which generates human–wolf conflicts. Therefore, we researched whether the behavior of loners resembles PP behavior. After being extinct, 22 loners had entered the Netherlands between 2015 and 2019. Among them, 14 could be DNA-identified and linked with their PPs in Germany. Some loners were siblings. We assessed the behavior of each individual and PP through a structured Google search. PP behavior was determined for the loner’s rearing period. Similarity between loner and PP behavior was significant (p = 0.022) and applied to 10 of 14 cases: like their PPs, three loners killed sheep and were near humans, five killed sheep and did not approach humans, while two loners were unproblematic, they did not kill sheep, nor were they near humans. Siblings behaved similarly. Thus, sheep killing and proximity to humans may develop during early-life experiences in the PP. However, by negative reinforcement that can be prevented. New methods are suggested to achieve that. As a result, new generations may not be problematic when leaving PPs.
... This is because individual and pack characteristics can influence depredation behavior in different ways. For example, studies showed that dispersing wolves were less likely (Fabbri et al. 2018), as likely (Fritts et al. 1992) or more likely (Imbert et al. 2016) than resident wolves to prey upon livestock. Several studies concluded that depredation risks increased with pack size (Treves et al. 2002, Bradley et al. 2015, whereas others found packs involved in depredations were medium-sized (Wydeven et al. 2004). ...
... In Yellowstone National Park (YNP) in the USA, hunting with other pack members is required to successfully take down large prey such as elk Cervus elaphus (MacNulty et al. 2012) or bison Bison bison . In southern Europe, wolves rely on smaller and more vulnerable prey, such as juvenile wild boar S. scrofa or roe deer Capreolus capreolus (Mattioli et al. 2004, 2011, Anceau et al. 2015, Imbert et al. 2016, Lagos and Bárcena 2018. Therefore, a wolf 's need to hunt in a group may be reduced in this situation (IPRA 2020), as suggested by the lower mean pack size in southern Europe (e.g. ...
The efficiency of the management of predations on livestock by gray wolves (Canis lupus) through culling is under debate. Evaluating wolf culling efficiency requires to simultaneously analyze the effects of culling on the wolf population and the repercussions of these population changes on livestock predation. This protocol is technically difficult to implement in the field. To properly assess culling efficiency, we provided an integrated and flexible individual‐based model that simulated interactions between wolf population dynamics, predation behavior and culling management. We considered many social processes in wolves. We calibrated the model to match the Western Alps as a case study. By considering the prey community in this area and the opportunistic nature of wolf predation, we assumed that predation on livestock at the wolf territory level increased with pack's food needs. Under this assumption and considering livestock availability as high and livestock vulnerability as uniform in space and time, culling maintained wolf population size and predation risks at low levels. Contrary to what was expected, culling decreased the mean annual proportions of dispersing wolves in our simulations, by speeding settlement. This population‐level mechanism compensated for the high mortality and the pack instability caused by culling. Compensation was however dependent on the selectivity and the timing of culling. When executed before the natural mortality module in our model, the selective culling could undermine replacement of lost breeders and therefore decrease wolf population resilience to culling. Our model gives insights about culling effects in one specific simulated context, but we do not expect that our assumption about predation behavior necessarily holds in other ecological contexts and we therefore encourage further explorations of the model.
... According to Kruuk and Parish (1981) to each consumed species and food category a volumetric class (<1%; 1-5%; 6-25%; 26-50%; 51-75%;76-95%; >95%) was assigned and then converted in a final percentage volume: 0.5%; 2.5%; 15.5%; 38%; 63%; 85.6% and 98% respectively (Meriggi et al., 2015). The mean percent volume (MV%) of each category and species was calculated (Imbert et al., 2016). Frequency of occurrence (FO%) of each food category was calculated as number of scats in which each prey species was detected within the total quantity of scats analyzed (Imbert et al., 2016;Torretta et al., 2017). ...
... The mean percent volume (MV%) of each category and species was calculated (Imbert et al., 2016). Frequency of occurrence (FO%) of each food category was calculated as number of scats in which each prey species was detected within the total quantity of scats analyzed (Imbert et al., 2016;Torretta et al., 2017). ...
A medium-sized mammals selection as food diet for pup performed by adult wolves has been pre- viously hypothesized. In this work the diet of wolf pups living in an anthropic area in Italy has been investigated to assess which prey potentially played a key role in their sustenance. Although wild ungulates were the main prey category for both pups and adults a significantly high occurrence of medium-size mammals (hare and coypu) and birds in pup diet has been recorded. Such result could confirm a selective food provision by adult wolves. Further investigations to assess the importance of this kind of prey for pup survival are desirable.
... From the analysis of wolf scats, it is evident that wolves in Germany, like their conspecifics in several other parts of Europe, feed predominantly on wild ungulates (Jędrzejewski et al. 2012;Wagner et al. 2012;Imbert et al. 2016;Ståhlberg et al. 2017;Mysłajek et al. 2018;2022;Mengüllüoğlu et al. 2019;Sin et al. 2019;Figueiredo et al. 2020;Valdmann and Saarma 2020;Reinhardt et al. 2021). In this context, roe deer are the main prey species (MPS) in most wolf territories in Germany. ...
... Due to its defensive behavior (greater body size of adult wild boar and fangs), the wolf usually avoids this species (Okarma 1995), even in areas with very high densities (Jędrzejewski et al. 2000).Wild boar has been identified as MPS in Italy, Anatolia and South-East Europe (Mattioli et al. 2011;Zlatanova et al. 2014;Ciucci et al. 2018;Mengüllüoğlu et al. 2019;Sin et al. 2019;Trbojević et al 2020). The subspecies Sus scrofa majori found in Italy is relatively small (Niethammer and Krapp 1986;Mattioli et al. 2011;Imbert et al. 2016) and may be easier to catch than the Sus scrofa scrofa subspecies found in Germany. In Anatolia, the wild boar is often the only wild ungulate species that occurs in high densities. ...
In Germany, wolves live in a human-dominated landscape. The return of the wolf to Germany more than 20 years ago is a prime example of successful nature conservation; however, it has also led to conflict due to the high degree of landscape fragmentation and the feeding habits of the wolf. Here, the wolf diet composition was studied based on an analysis of 11,225 scats. Wolves in Germany mainly feed on wild ungulates, which comprise more than 94% of their diet. The predominant species are roe deer and wild boar, whereby in most territories roe deer dominate the diet composition. Depending on availability, red deer and fallow deer may also make up a high proportion of the diet. Seasonal and territorial variation in diet composition has been noted even in directly neighboring territories. Wolves preferentially hunt juveniles of the main prey species. With the exception of the mouflon, which has all but disappeared locally, no other ungulate species has vanished from the wolf’s diet. This study provides an overview of the varied diet of wolves in Germany since recolonization more than 20 years ago.
... Wolves also make extraterritorial excursions (i.e., predispersal movements) of varying distance and duration 27,28 . Greater use of human-disturbed areas by wolves when dispersing or floating could result from avoiding existing wolf territories in less humandisturbed areas or decreased site familiarity that reduces their ability to avoid human disturbances, compared to when they are resident of a territory 8,29 . Alternatively, disturbances such as roads may facilitate efficient travel for non-residents 30 , while areas with high livestock abundance may provide food when lower site familiarity or prey abundance limits acquisition of wild prey 29 . ...
... Greater use of human-disturbed areas by wolves when dispersing or floating could result from avoiding existing wolf territories in less humandisturbed areas or decreased site familiarity that reduces their ability to avoid human disturbances, compared to when they are resident of a territory 8,29 . Alternatively, disturbances such as roads may facilitate efficient travel for non-residents 30 , while areas with high livestock abundance may provide food when lower site familiarity or prey abundance limits acquisition of wild prey 29 . ...
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.
... Throughout history, the wolf has faced threats of extinction and today the increase in the number of heads of this species represents an ecological recovery. The wolf recolonization started naturally in the 1980s first in northern Apennines [1] and then from the 1990s in the Alps [2,3]. Now the species is expanding and is present in all the Alps, coming from both the Apennines-western Alps and from the northeast. ...
... Locally high wolf predation on horses is reported in many parts of southern Europe. In Italy wolf predations on horses are observed in most part of the Italian territory, such as in the Abruzzo region [4], in the Northeastern Apennines [1], in Northern Italy [2,3] and in Southern Apennines [5][6][7] where horses can locally reach 40% of wolf diet [4]. According to the ISPRA (Italian Institute for Environmental Protection and Research, 2022) survey [8,9] a number of around 950 specimens is estimated in the Alpine regions, while there are almost 2400 those distributed along the rest of the peninsula, for a total of about 3300 wolves in Italy. ...
Although wolf predation on horses is generally low, it can be significant where horses are grazed extensively and there are low densities of wild prey, such as in Southern Europe. The economic damage of wolf predation on horses can have a major impact in certain situations involving free-ranging horse populations and the topic needs more attention. This is the case of Spain, Portugal and Italy where it is possible to find many examples of wolf predations on feral horses. The main factor predisposing horses to wolf pre¬dation worldwide is the free-ranging management system. In Italy special attention is given to the horses bred in a particular area of the Southern Apennines: the Pentro horses; they represent a rare endangered equine population of Molise Region with its owns genetic characterisation. The losses of horses caused by wolves predations increase exponentially from year to year. Considering the very low number of Pentro Horses, any prey represents a considerable loss from a morphological and genetical point of view. Consequently, the predation phenomena represent a considerable limit to the selection and maintenance of these horses. This is why it was decided to create a special database with the aim of monitoring, analyzing and preventing predatory phenomena and consequently to keep under control the number of predatory events by the wolf. The database was processed by collection of data concerning: when the predation take place, the age of the preyed equines, the type of injury and the parts of the body injured or removed by the wolf, the body location of the damage and the recognition and value of money compensation. The data collected evidenced how there is a need for innovative approaches to prop¬erly mitigate predation and support the traditional free-ranging husbandry system with its high ecologi¬cal and cultural value.
... No comprehensive study on the number of wolves killed or poisoned per year exists, but human-caused mortality of wolves is confirmed (40 poisoned animals: 2000-2016Ntemiri et al., 2018). Wolf persecution to mitigate livestock damage seems to be frequent in Greece as in other European countries (Imbert et al., 2016;Boitani et al., 2022). Although the country lacks a national action plan for the conservation of grey wolves, the problem of increased human-caused wolf mortality in Greece was recognized in the Prioritized Action Framework (PAF) for Natura 2000 (2021-2027) and measures were suggested to enhance wolf-human coexistence (PAF, 2020). ...
... Reducing livestock losses from wolf depredation is challenging. Restoring wild ungulate populations to decrease wolf dependence on livestock and implementing preventive measures are synergistic actions that would reduce wolf retaliation by livestock farmers and substantially enhance species conservation (Imbert et al., 2016;Janeiro-Otero et al., 2020). There is a need for better quantitative assessment of the efficacy of preventive measures for protecting sheep/goats and cattle. ...
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Abstract
Livestock depredation is the primary driver of wolf-human conflict worldwide, threatening wolf conservation and impacting human livelihoods. Most countries implement relevant compensation programs, which are however rarely accompanied by proactive husbandry practices vetted with scientific research. We investigated the influence of husbandry practices on wolf depredation losses for 70 sheep/goat and 68 cattle herds with quantitative modeling of data from semi-structured interviews of livestock farmers along a livestock damage gradient in NW Greece. Sheep/goat herds were better protected than cattle herds in seven preventive measures and annual losses of sheep/goat livestock units were three times lower than losses of cattle livestock units in our study area. Furthermore, according to national compensation data from Greece, costs paid for cattle have doubled in recent years, whereas they have been cut in half for sheep/goats. Our modeling identified three core preventive measures that significantly reduced wolf depredation risk for both herd types, namely increased shepherd surveillance, systematic night confinement, and an adequate number of livestock guardian dogs (optimal ratio was 3 Greek guardian dogs per 100 sheep/goats and 7 guardian dogs per 100 cattle). Keeping young livestock in enclosures and not abandoning livestock carcasses in pastures were additional effective preventive measures for cattle herds. Our study provides evidence to inform the subsidizing policy put forth in the Common Agricultural Policy of the European Union. It can also serve to inform or revise wildlife-livestock conflict mitigation policy in countries challenged with the competing goals of conserving large carnivores while maintaining traditional grazing regimes.
... Thus, cooperative hunting of the larger prey might increase hunting success, but also might offset the risk of injury to some degree (Mech 1970). One other very interesting aspect, especially in the light of wolves returning all over Europe is that one study, examining 1,457 scats between 2008 to 2013 in Italy, calculated a model that showed the presence of stable packs, instead of dispersing wolves, reduced predation on livestock (Imbert et al. 2016). ...
... There are also some studies indicating that wolves might prefer wildlife over livestock consumption. For example, one study showed that while the presence of domestic ungulates on high-altitude pastures during summer (May-October) influenced wolf diet (summer 19.0%, winter 0.3%) in the Italian Alps, wolves still preferred wild ungulates despite the higher density of domestic livestock (Gazzola et al. 2005;Imbert et al. 2016; see also Meriggi and Lovari 1996). Other studies have found that consumption of livestock by grey wolves decreased over time, coincident with an increase in relying on wild ungulates in southern Italy (Newsome et al. 2016). ...
This short review summarizes aspects of the socio-ecology of wolves that might be relevant to understand dog-wolf differences in behaviour and cognition. It highlights the cooperative nature of wolves that usually live in family packs, raise their pups together, and jointly participate in hunting, as well as defending their territories and carcasses. However, the size and stability of family packs and the dynamics of their cooperative interactions, while still under investigation, are thought to be influenced by a number of socio-ecological factors such as the degree of saturated habitats, prey species availability, habitat disturbance, as well as kin selection and territory inheritance.
... Wolves are pack hunters and are known to feed on a variety of different food items. They choose their prey based on availability, abundance, pack stability, season, and habitat accessibility in the human-dominated landscapes (Imbert et al., 2016;Lyngdoh et al., 2020). Our study revealed that the diet of Woolly wolf from the Himalayan region consisted of 20 different food items from small birds, reptiles to large mammals and domestic animals such as cattle and yak. ...
... The Woolly wolf also consumed a sufficient proportion of the large-sized prey (32.32%) and medium-sized prey (33.87%) with a considerable quantity of small prey (24.18%) consumption in their diet (Figure 5). Various studies across the wolf distribution range confirm that large prey forms the major part of the wolf diet (Imbert et al., 2016;Mengulluoglu et al., 2019;Petridou et al., 2019;Sin et al., 2019). The dependency on small prey might be because of the scarcity of the large and medium-sized animals to avoid interactions with humans or to gain and fulfill energy requirements in the harsh climatic condition. ...
Geographical isolation can often lead to speciation, and two disconnected populations of the same species living in drastically different bioclimatic regions provide an opportunity to understand the process of speciation. The Woolly wolf is found in the cold-arid, Trans-Himalayan landscape, while the Indian wolf inhabits the semi-arid grasslands of Central India. Both the lineages of wolves from India have generated scientific debate on their taxonomic status in recent years. In this study, we collected data and reviewed published literature to document the ecological and behavioral differences between the Woolly wolf and the Indian wolf. Most studies have used genetic data; hence we discuss variation in spatial ecology, habitat preferences, vocalization, diet diversity and cranial measurements of these two subspecies. The spatial ecology of two lineages was compared from the data on three Woolly and ten Indian wolves tagged with GPS collars. The telemetry data shows that there has been no difference in the day-night movement of Woolly wolves, whereas Indian wolves show significant high displacement during the night. The BBMM method indicated that Woolly wolf home ranges were three times larger than the Indian wolf. The Woolly wolf diet is comprised of 20 different types of food items, whereas the Indian wolf diet consists of 17 types. The Woolly and Indian wolf largely depend upon domestic prey base, i.e., 48.44 and 40.34%, respectively. We found no differences in the howling parameters of these subspecies. Moreover, the Woolly wolf skull was significantly longer and broader than the Indian wolf. Wolves of India are ancient and diverged from the main clade about 200,000–1,000,000 years ago. Their genetic and ecological evolution in different bioclimatic zones has resulted in considerable differences as distinct subspecies. The present study is a step in understanding ecological differences between two important, genetically unique subspecies of wolves.
... Wolves shift to wild boar in winter, when most attacks on dogs occur, after the vulnerable roe deer fawn age class diminishes (i.e., 71% in summer scats) and vulnerability of young wild boar increases, a seasonal pattern also observed in other similar studies from Europe [98,102,103]. Apart from wild boar, hunting dogs may represent an alternative, locally abundant, and highly vulnerable prey during winter, also given the positive trend in their numbers in the last 10 years, as mentioned. ...
... Although recovery of wild boar and roe deer is ongoing in the study area, trophic stability of wolf populations and the reduction of wolf-human conflicts are achieved by increasing not only density, but also the diversity of wild ungulates [3,103]. Feasibility of reintroducing additional ungulates species previously native in the study area can be considered. ...
Hunting dog depredation by wolves triggers retaliatory killing, with negative impacts on wildlife conservation. In the wider area of the Dadia-Lefkimi-Soufli Forest National Park, reports on such incidents have increased lately. To investigate this conflict, we interviewed 56 affected hunters, conducted wolf trophic analysis, analyzed trends for 2010–2020, applied MAXENT models for risk-map creation, and GLMs to explore factors related to depredation levels. Losses averaged approximately one dog per decade and hunter showing a positive trend, while livestock depredations showed a negative trend. Wolves preyed mainly on wild prey, with dogs consisting of 5.1% of the winter diet. Low altitude areas, with low to medium livestock availability favoring wolf prey and game species, were the riskiest. Dogs were more vulnerable during hare hunting and attacks more frequent during wolf post-weaning season or in wolf territories with reproduction. Hunter experience and group hunting reduced losses. Wolves avoided larger breeds or older dogs. Making noise or closely keeping dogs reduced attack severity. Protective dog vests, risk maps, and enhancing wolf natural prey availability are further measures to be considered, along with a proper verification system to confirm and effectively separate wolf attacks from wild boar attacks, which were also common.
... To reduce livestock depredation by large carnivores and mitigate social conflicts between conservationists, farmers, and other stakeholder groups, we need to evaluate when, where, and why large carnivores depredate livestock (Bautista et al., 2019;Mayer et al., 2022;Singer et al., 2023), as well as address other factors that contribute to livestock losses. Livestock damage by large carnivores is often restricted to few farms (Gazzola et al., 2008), and depends on landscape structure and availability of natural prey (Imbert et al., 2016;Suryawanshi et al., 2013;Treves et al., 2004). Although several mitigation measures have been implemented in various European regions, such as zoning, predator-proof fencing, guard dogs, the use of specific breeds or largerbodied livestock (Bautista et al., 2019;Lorand et al., 2022), a contextdependent evaluation of their efficiency is often lacking. ...
Large carnivore populations have increased in many areas of Europe, resulting in socio-political conflicts, especially in areas with extensive outfield grazing systems. In Norway, the occurrence of large carnivores, especially brown bears (Ursus arctos) and grey wolves (Canis lupus), are largely separated from outfield grazing areas via carnivore management zones. However, the overall drivers of livestock losses in the outfields and the contribution of large carnivores are not well understood. Here, we investigated (1) correlates of cattle and sheep losses across organized grazing areas of Norway to evaluate which factors, including large carnivore densities, livestock density, the size of the grazing area, tick density, landscape characteristics, and climatic variables, are responsible for livestock losses in the outfields, and (2) spatio-temporal correlates of cattle losses suspected of being attacked by large carnivores. Based on our analyses investigating cattle and sheep losses in organized outfield grazing areas, we found no evidence that large carnivore densities correlated with cattle losses, but the proportion of sheep losses increased with increasing large carnivore densities, especially of bears and wolverines (Gulo gulo). Importantly, factors other than large carnivores, such as tick density and landscape composition, were related to higher probabilities of cattle losses. Our study provides important information concerning the overall causes of livestock losses in the outfields, indicating that large carnivores contribute little to total cattle losses.
... The wild boar is often one of the main prey species for wolves at places where it is presented (Salvador & Abad, 1987;Kübarsepp & Valdmann, 2003;Andersone & Ozoliņš, 2004;Capitani et al., 2004;Imbert et al., 2016;Ståhlberg et al., 2017). In the Kaluzhskie Zaseki SNR, this species had been the main prey species for several decades (Hernandez-Blanco & Litvinova, 2003). ...
Canis lupus (hereinafter – wolf) is a top predator specialised on large ungulates. In a community of multiple ungulate species, carnivores may exhibit a preference for one. Due to the decreasing availability of preferred species, interactions between wolves and other ungulates can change. We conducted our research from 2014 to 2019, which included an abrupt decline in the population of the wolf’s main prey, Sus scrofa, in the Kaluzhskie Zaseki State Nature Reserve, Russia. We ascertained the wolf diet (scats content analysis) and evaluated preferences using Jacobs’ index. The proportion in the diet of two ungulate species, Capreolus capreolus and Alces alces, increased after the Sus scrofa population declined, but the main prey became a non-ungulate species, namely Castor fiber. Wolves did not hunt Bison bonasus, and changes in Sus scrofa populations did not affect interactions between them. The presence of this species in the wolf diet is negligible and related to scavenging. Wolves showed a strong preference for Sus scrofa after decreasing its abundance. Dynamic interactions estimated from camera-trap data, using the coefficient of sociality, partly confirmed with analysis of preferences but also depending on other factors. The coefficient of sociality showed attraction species hardly presented in the diet only in the peripheral part of the wolf home range, where ungulates more frequently encounter subadult wolves. The decreasing population of wolf’s main prey influenced interactions with other ungulates to some extent since wolves increased the proportion of them in their diet to substitute Sus scrofa. In the absence of Sus scrofa, wolves tend to prefer middle- and small-sized prey.
... This is due to the comparatively small structured farms, the complex topography, the lack of personnel as well as the high influx of hiking tourism (Pfister, 2010;Hackländer et al., 2019;Moser et al., 2019) and disturbed wildlife movement (Jakes et al., 2018). The reduced application of protection measures of grazing areas in Alpine regions has led to a notable rise in predation numbers of livestock caused by wolves, as compared to non-alpine regions (Capitani et al., 2004;Gazzola et al., 2005;Imbert et al., 2016). To compensate farmers for the resulting financial losses, EU regulations govern compensation payments in the event of damage to livestock by protected predators. ...
In the Alpine region, the use of herd protection measures is assumed to not be reasonable in many places due to geographical, technical and economic limitations. In line with EU regulations , securing compensation for livestock damage or losses necessitates evidence of the feasibility of these measures. The aim of this study was to establish a scientifically validated classification framework for assessing the viability of employing herd protection measures (livestock guarding fences and guarding dog utilization) in alpine pastures. To achieve this, five representative Alps in the Italian alpine province of South Tyrol were considered using a devised classification framework and geospatial data. The resultant outcomes were validated through on-site inspections. Six categories were identified for the use of livestock guarding fences (proximity to roads, herding situation, topography, tourism, pasture geometry and proportion of not fencable area) and, in addition to these, two categories (herd homogeneity and number of herded livestock) for the use of livestock guarding dog utilization. The validation showed that the basic assumptions are plausible, but that the geodata partly show inaccuracies in the demarcation of hazard zones, hiking trails and pasture geometry. Despite these limitations, the developed classification framework was suitable for classifying the practicability of herd protection measures on alpine pastures. Therefore, it provides the necessary factual basis and logical reasoning for legally secure compensation payments. Additionally , our classification framework sets the stage for considering strategic predator management approaches.
... For example, conflict commonly occurs where carnivores have access to anthropogenic food sources, exemplified by wolves preying on livestock in rural areas (Janeiro-Otero et al., 2020), coyotes feeding on pets in suburban neighborhoods (Poessel et al., 2013), or bears foraging on garbage in cities and crops in agricultural lands (Scotson et al., 2014). Additionally, studies have repeatedly shown that human-carnivore conflicts increase when natural (non-anthropogenic) foods consumed by carnivores are limited (Acosta-Jamett et al., 2016;Artelle et al., 2016;Imbert et al., 2016;Johnson et al., 2015). These patterns have advanced understanding of generalizable factors associated with human-carnivore conflict (hereafter, "causes"), and improved the ability to predict where, when, and to what extent human-carnivore conflict versus coexistence might be expected. ...
Human‐carnivore conflicts pose significant challenges in the management and conservation of carnivores across the globe. Abundant research has led to generalizable insights into the causes of such conflicts. For example, conflicts predictably occur when carnivores have access to human food resources, particularly when their natural foods are scarce. However, similar insights into the effectiveness of interventions aimed at coexistence remains comparatively scarce. We hypothesized that this disparity might be reflected in a bias toward research focused on causes of conflict rather than interventions to address it. To test our hypothesis, we evaluated the content of studies on human–carnivore conflicts and coexistence in Canada and the United States from 2010 to 2021. We found that studies disproportionately focused on causes of conflict, with that discrepancy increasing through our study period. We also found a disproportionate focus on black bears and wolves and western jurisdictions, and a disproportionate use of observational (vs. experimental) approaches. Studies on conflict interventions were primarily directed at the carnivores themselves (e.g., lethal approaches) versus human elements (e.g., attractant management, policies), despite evidence that the latter are more effective. We expect that a shift in focus toward solutions‐oriented research, integrating insights across geographies, taxa, social contexts, and disciplines, would facilitate effective interventions and foster coexistence, improving outcomes for people and carnivores alike.
... As expected, the risk of an attack on livestock decreased with the maximum distance to a wolf territory. However, if wolves roam through an area not yet occupied by a wolf pack, there is increased pressure on the livestock from these dispersed wolves (Imbert et al., 2016;Mech and Boitani, 2010;Packard, 2003). In newly occupied regions, the risk of a wolf attack on livestock was considerable (Müller, 2020;Reinhardt and Kluth, 2007). ...
... Rooting is a trophic behavior of wild boars in the forest environment that results in soil renewal, leading to its aeration, as well as improved oxygen penetration and the mixing of nutrients and organic matter [26,27]. Several studies have highlighted numerous positive effects of this species, which exerts a natural plowing effect on the soil [6,8,28]. These studies have rehabilitated the functional role of the wild boar, which appears to be ecologically beneficial for both fauna and flora, and especially for the soils of the environments they colonize [29]. ...
Wild boars use a wide range of habitats. Their invasive nature is gaining attention due to the complexity of its impact. The goal of this research is to analyze the impact of the wild boar on the chemical properties of soils in a natural and a post-fire forest in the Edough Forest Massif in Algeria. This study compares the impact of wild boar rooting on soil parameters to determine the functional role of the wild boar. The research was conducted during the winter of 2022. The study sites included a natural forest and a post-fire area. Rooting tracks were geolocated and soil samples were collected. The results show significant differences between rooted and control patches in the chemical parameters measured in the two environments. However, in the natural environment, significant differences were only noted for the calcium content and electrical conductivity. But in the post-fire environment, strong significant differences were observed for all measured parameters, suggesting that wild boars do not exert a noticeable soil homogenization effect on the soil properties. This research highlights the importance of understanding and managing the impact of wild boars in natural and post-fire forests on soil formation processes, the diversity of soil properties, and their magnitude.
... Such unprotected livestock creates easily accessible prey that in some regions comprises an important food source for wolves (Lagos & Bárcena, 2018;Vos, 2000). The availability of wild relative to domestic prey is an important factor determining the proportion of livestock in the wolf diet (Imbert et al., 2016;Meriggi & Lovari, 1996). Moreover, there is a growing interest in rewilding projects in Europe aiming to restore trophic interactions (Bakker & Svenning, 2018). ...
The wolf (Canis lupus) is arguably the most successful species at recolonizing its now human‐dominated former ranges in Europe and North America. Over the centuries while the wolf was absent, humans have transformed ecosystems to a large extent. In this paper, we highlight key aspects of these human‐modified ecosystems that include changes to (meso)carnivore communities, wolves themselves (genetics, behaviour), woody plant communities and the playing field for predator–prey interactions (landscape structure).
We argue that the recognition of the novelty of human‐modified ecosystems logically leads to novel pathways of how wolves can influence ecosystem functioning. Thus far, the ecological impacts of wolves in human‐dominated systems have largely been predicted based on the documented effects they have on prey species or lower trophic levels in well‐preserved systems with low human impact. However, wolves in human‐modified ecosystems will engage in an array of novel interactions and potential novel trophic cascades that do not occur in more natural ecosystems with lower human impact.
This should encourage us to re‐assess the questions we ask about wolf impacts in novel systems. A promising direction for future studies is exploring what novel interactions establish and under what conditions wolves can exert their ecosystem impacts (context dependence) in the human‐modified ecosystems wolves are recolonizing.
Policy implications. Understanding these novel interactions and the context dependence of ecosystem impacts could guide us to act to improve conditions to enable wolves to exert their ecosystem impacts again. These novel interactions may be the true ecological and societal value of having wolves returning to human‐modified landscapes.
... Specifically, during their expansion, wolves not only benefited from such landscape changes, but also mutually constituted them, possibly becoming a more-than-human force reproducing structural inequities. Although wolves largely reoriented their feeding ecology towards wild prey in the closing decades of the twentieth century Meriggi et al., 2020), they also preyed on unguarded livestock that they increasingly encountered during their return (Ciucci and Boitani, 1998;Imbert et al., 2016;Gervasi et al., 2022). Husbandry practices, following local extirpations of wolves, were no longer adapted to the presence of this predator (Boitani, 1992). ...
There is increasing interest in human-wildlife coexistence. In Europe, coexistence with expanding large carnivore populations is a pressing issue. Seeking to inform contemporary management with an historical perspective, this study investigates the formation of human-wolf relations in Italy during critical periods of wolf decline (since the nineteenth century) and comeback (since the closing decades of the twentieth century). Specifically, this study adopts a 'more-than-human political ecology' approach, focused on exploring the entangled influence of non-human agency and wider political economies in the (co)production of human-wildlife relations. Such analysis is used to fill knowledge gaps on these key historical events in carnivore management, which are currently explained from either a more rigid structural angle or depoliticised lenses. The analyses of this study bring together the pivotal work of Italian historians and scholars on topics of capitalist development and wolf ecology, ethology and management in Italy. This work highlights how the onset of capitalism in Italy around the end of the 1700s inadvertently affected the rise of particularly problematic wolf behaviours in that period, which were in turn managed through an intensification of wolf persecution. Wolves co-shaped the need of and desire for their conservation near the end of the 1900s, which alongside the rise of a postmodern capitalist regime promoted the return of wolves in this period. The (re)alignment of wolf conservation with capitalist logics, however, displaced the costs of the wolf's return onto local communities, exacerbating conflict. This study comes with two main implications: first, it problematises fixed representations of non-humans, highlighting instead their adaptive capacities and alterity; second, it further conceives non-humans as in a constant co-becoming with human practices and wider political economies, emphasising the need for structural change for conviviality. These analyses may inform wolf management in Europe and beyond, as well as other contexts of coexistence.
... Nomadic wolves moving in unfamiliar landscapes ("floaters", sensu Fuller et al., 2003) are probably more prone to become exposed to ARs. By not hunting large prey in groups (MacNulty et al., 2012), floaters usually shift to easier prey, such as livestock (Imbert et al., 2016). ...
Second-generation Anticoagulant Rodenticides (ARs) can be critical for carnivores, due to their widespread use and impacts. However, although many studies explored the impacts of ARs on small and mesocarnivores, none assessed the extent to which they could contaminate large carnivores in anthropized landscapes.
We filled this gap by exploring spatiotemporal trends in grey wolf (Canis lupus) exposure to ARs in central and northern Italy, by subjecting a large sample of dead wolves (n = 186) to the LC-MS/MS method.
Most wolves (n = 115/186, 61.8%) tested positive for ARs (1 compound, n = 36; 2 compounds, n = 47; 3 compounds, n = 16; 4 or more compounds, n = 16). Bromadiolone, brodifacoum and difenacoum, were the most common compounds, with nrodifacoum and nromadiolone being the ARs that co-occurred the most (n = 61).
Both the probability of testing positive for multiple ARs and the concentration of brodifacoum, and bromadiolone in the liver, systematically increased in wolves that were found at more anthropized sites. Moreover, wolves became more likely to test positive for ARs through time, particularly after 2020.
Our results underline that rodent control, based on ARs, increases the risks of unintentional poisoning of non-target wildlife. However, this risk does not only involve small and mesocarnivores, but also large carnivores at the top of the food chain, such as wolves. Therefore, rodent control is adding one further conservation threat to endangered large carnivores in anthropized landscapes of Europe, whose severity could increase over time and be far higher than previously thought. Large-scale monitoring schemes for ARs in European large carnivores should be devised as soon as possible.
... Το κόκκινο ελάφι έχει σχεδόν εξαφανιστεί από όλη την έκταση της εξάπλωσής του, οι πληθυσμοί του αγριόγιδου έχουν συρρικνωθεί, το πλατώνι έχει εξαφανιστεί εντελώς από την ηπειρωτική Ελλάδα, oι πληθυσμοί τoυ ζαρκαδιού έχουν υποστεί σημαντική μείωση, ενώ και οι πληθυσμοί του αγριόχοιρου έχουν επίσης μειωθεί (Papaioannou, 2010). Κατά τη διάρκεια των τελευταίων δεκαετιών, οι πληθυσμοί των αγριόχοιρου και του ζαρκαδιού φαίνεται να έχουν αυξηθεί σε ορισμένες περιοχές της Ελλάδας (Tsachalidis and Hadjisterkotis, 2009;Ntolka et al., 2016), αλλά οι πυκνότητές τους παραμένουν χαμηλές συγκριτικά με άλλες ευρωπαϊκές χώρες (Meriggi et al., 2014;Imbert et al., 2016). Ο αγριόχοιρος είναι το μόνο θηρεύσιμο οπληφόρο είδος στην Ελλάδα. ...
Greek abstract of PhD thesis entitled: "Wolf ecology and predation behavior: a conflict minimization strategy towards sustainable livestock breeding" (in english).
Η παρούσα διατριβή εξετάζει τη σύγκρουση μεταξύ λύκου και ανθρώπου, η οποία προκύπτει κυρίως από τη θήρευση κτηνοτροφικών ζώων, αποτελώντας σημαντική απειλή τόσο για τη διατήρηση των λύκων όσο και για τον κλάδο της κτηνοτροφίας. Για να επιτευχθεί η βιώσιμη συνύπαρξη λύκου και ανθρώπου, απαιτείται πλήρης κατανόηση των οικολογικών και κοινωνικών παραγόντων που την επηρεάζουν. Η παρούσα διατριβή υιοθετεί μία διεπιστημονική προσέγγιση, διερευνώντας τις τροφικές συνήθειες των λύκων, τα χωροχρονικά του πρότυπα σε σχέση με τις ανθρώπινες δραστηριότητες, την επίδραση των κτηνοτροφικών πρακτικών στα επίπεδα θήρευσης από τον λύκο, καθώς και τις προκλήσεις που αντιμετωπίζουν οι κτηνοτρόφοι σε περιοχές με παρουσία λύκου. Στο παράπανω πλαίσιο, η διάρθωση της διατριβής είναι η ακόλουθη.
Το Kεφάλαιο 1 παρέχει μια γενική εισαγωγή στο θέμα και τις κύριες έννοιες της διδακτορικής διατριβής.
To Κεφάλαιο 2 εξετάζει τις τροφικές συνήθειες του λύκου σε μια περιοχή της Κεντρικής Ελλάδας, με έντονη ανθρωπογενή παρουσία. Ειδικότερα, αυτή η περιοχή χαρακτηρίζεται από υψηλή διαθεσιμότητα κτηνοτροφικών ζώων και περιορισμένους πληθυσμούς άγριων οπληφόρων.
Το Κεφάλαιο 3 διερευνά, με τη χρήση αυτόματων υπέρυθρων καμερών, τα χωροχρονικά πρότυπα δραστηριότητας του λύκου, των κυριότερων άγριων θηραμάτων του, και άλλων σαρκοφάγων θηλαστικών στη Βορειοδυτική Ελλάδα. Παράλληλα εξετάζει τη σχέση αυτών των χωροχρονικών προτύπων με τις ανθρώπινες δραστηριότητες.
Το Κεφάλαιο 4 εστιάζει στον ρόλο των κτηνοτροφικών πρακτικών στα επίπεδα θήρευσης από τον λύκο στη Βορειοδυτική Ελλάδα και παράλληλα παρουσιάζει μια επισκόπηση των τάσεων στις απώλειες που προκαλούνται από τον λύκο σε εθνικό επίπεδο.
Το Κεφάλαιο 5 εξετάζει τις προκλήσεις που αντιμετωπίζουν οι κτηνοτρόφοι που δραστηριοποιούνται σε περιοχές στη Βορειοδυτική Ελλάδα με παρουσία λύκου. Επίσης, αξιολογεί τις απόψεις τους ως προς τον αντίκτυπο που έχουν οι λύκοι στο επάγγελμα τους.
... In times of scarcity, wolves will eat carrion and also consume some fruits and vegetable. Episodes of cannibalism against wounded, weakened specimens, or carcasses are known (Canalis, 2012;Geptner et al., 1988;Mech, 1971;Mech et al., 2004;Meriggi et al., 2016;Nowak, 1983). ...
The analysis of animal hair is a useful and non-invasive investigation method for monitoring metal content, whose beginning dates to a few decades ago. This study addresses the issue of wildlife mortality in Piedmont and Aosta Valley by linking the evidence to the characteristics of the territories and to hair elemental profile. The considered animal species were badger, fox, marten, and wolf. The quantitative data for 11 trace metals and 16 lanthanides were evaluated considering animal location and death causes regarding impacts, whose relevant number was confined to the Turin province and to the badger and fox species. The evaluation of the analytical results was performed after the out of bounds sample identification by mean of statistics. For trace metals, some areas have thus been identified, mainly in Turin province, in which the large excess of As, Cd, and Pb was related to district pollution. Moreover, the lower contents of Fe, Mn, and Al in Aosta Valley badgers’ hair seem influenced by the different characteristics of the subsoil, in comparison to Piedmont, and a relationship with the living environment and the habits was suggested. Regarding lanthanides, Biella province represents an area in which environmental contamination and abundance of rare-earth elements was recorded.
... v) Environmental factors that have been previously reported may drive the consumption of different prey species [46,47], thus, season and year are predicted to potentially influence wolf diet in the study area. We predict that in spring the consumption of wild ungulates will be higher than domestic ones due to the greater availability of young prey of different species. ...
The Iberian wolf (Canis lupus signatus) is recolonizing historical distribution areas after decades of absence. As in other human-dominated landscapes, finding a balance to protect this species by favoring recolonization and mitigating human-wildlife conflicts is a challenge. Since wolves are often generalist opportunistic predators, we studied their diet composition in the Central Spain to evaluate the consumption of domestic ungulates and provide reliable data that help local authorities to deal with the current wolf-cattle ranchers conflict and coexistence. Diet composition (% prey occurrence, % prey ingested biomass) was analyzed through the identification of prey hairs present in 671 scats collected between 2017 and 2021. Wolves fed more on wild ungulates (82% occurrence) than domestic ones (18%). Wild boar (Sus scrofa, 44% occurrence) and roe deer (Capreolus capreolus, 35%) were the most consumed prey. Wolves positively selected these two species. The wolf's diet varied between seasons, years, and forest regions, but a diet based on wild ungulates predominated over domestic ones. Food niche breadth and showed variations depending on seasons and years. Preserving the availability and diversity of wild ungulates may favor reducing livestock attacks and would be an achievable goal that help to conserve this species and reduce conservation conflicts.
... The wolf (Canis lupus) preferably predates wild animals (Imbert et al., 2016;Miller et al., 2016), and domestic ungulates (Kaartinen et al., 2009), for this reason causing greater conflicts with humans and their economic interests regarding livestock (Murray, 2006). Furthermore, dogs (Canis familiaris) are widely distributed in southern European countries, coexisting with wolves, thus wolf predation can sometimes be confused with those caused by other carnivores (Cozza et al., 1996). ...
... The analyses have been frequently conducted to provide data for mitigating economic loss due to livestock depredation. For example, domestic animal predation by the wolf (Imbert et al. 2016), caracal (Jansen et al. 2019), and leopard have been reported (Ott et al. 2007). Besides those prominent studies on depredation, study on the indirect effect of carnivores on food production through trophic interaction received less attention (Baker et al. 2008). ...
Dietary study of carnivores has garnered a great deal of attention for a very long time since it provides information for comprehending ecological processes and tracking the economic effects of animals on food production. Few studies reported alternative prey that is detrimental to food production, such as rodents and insects, in addition to the numerous studies on livestock predation. Therefore, this study aims to provide information on the diet composition and overlap of two mesocarnivore species, Javan mongoose (Urva javanica) and leopard cat (Prionailurus bengalensis), in Java, Indonesia. The macroscopic observation technique was applied for identifying food remnants found in the 90 feces of the mongoose and 100 feces of the cat collected in 2018. Twenty three and 21 taxa of prey were found in the leopard and Javan mongoose feces. Rodentia was the most frequent taxa found in the feces of both species, followed by Galliformes. Insects comprised the majority of diet items but at low frequencies. Two mesocarnivores were specialists (Levin’s index of leopard cat: 0.08; Javan mongoose: 0.26) with fairly high diet overlap (Pianka’s index = 0.78). The results provided compelling evidence of the positive and negative impacts of mesocarnivores on food production in Java.
... Wolves require rich and diverse ungulate populations as a prey-base to reduce predation on livestock and subsequent wolf-human conflicts [79]. Increased roe deer availability for wolves can significantly decrease livestock consumption, as it acts as a second potential prey species in Mediterranean landscapes, in addition to wild boar [80]. We suggest that human disturbance could indirectly increase wolf-livestock conflict, since it was found to increase wolf nocturnality and, consequently, reduce opportunities for wolves to prey on roe deer during the day, while spatially restricting roe deer availability. ...
In an era of increasing human pressure on nature, understanding the spatiotemporal patterns of wildlife relative to human disturbance can inform conservation efforts, especially for large carnivores. We examined the temporal activity and spatial patterns of wolves and eight sympatric mammals at 71 camera trap stations in Greece. Grey wolves temporally overlapped the most with wild boars (∆ = 0.84) and medium-sized mammals (∆ > 0.75), moderately with brown bears (∆ = 0.70), and least with roe deer (∆ = 0.46). All wild mammals were mainly nocturnal and exhibited low temporal overlap with human disturbance (humans, vehicles, livestock, and dogs; ∆ = 0.18-0.36), apart from roe deer, which were more diurnal (∆ = 0.80). Six out of nine species increased their nocturnality at sites of high human disturbance, particularly roe deer and wolves. The detection of wolves was negatively associated with paved roads, the detection of roe deer was negatively associated with human disturbance, and the detection of wild boars was negatively associated with dogs. The detection of bears, boars, and foxes increased closer to settlements. Our study has applied implications for wolf conservation and human-wildlife coexistence.
... In fact, the northernmost GPS location recorded during this movement occurred along a hypothetical corridor (the Ticino River Natural Park), but in an area where an abrupt decline of ecological connectivity was predicted due to human settlements and roads [41]; moreover, during the last few years, two wolves were killed by car accidents exactly in this area. After the unidirectional dispersal southward, the NSD of W2357M showed two peaks and a turnaround, suggesting it had not found an available area to settle in the Northern Apennines, probably due to high pack density characterising the area [45,86,87]. ...
Simple Summary
In the last decades, a significant recovery and natural expansion of the wolf (Canis lupus) populations has occurred across Europe. This remarkable recolonisation was made possible by the high plasticity of the species, which was crucial in such a human-altered environment. Despite re-establishing within their former distribution range, understanding the behavioural responses adopted by this large carnivore to navigate in an increasingly anthropogenic world remains challenging. In this study, we investigated the movement ecology of three rehabilitated wolves in one of the most human-dominated landscapes of Europe, the Po Plain in Northern Italy, and obtained preliminary evidence of the ability of wolves to spatio-temporally segregate from human activities. We observed that (i) when wolves settled, they made considerably longer daily movements; (ii) when dispersing, wolves were more nocturnal in order to avoid encounters with humans; (iii) long-distance movements were aided by the availability of small-wooded patches used as resting areas. Our results provide important insight into the movement patterns of wolves in anthropogenic ecosystems, which may be used to inform future management actions that aim to facilitate wolf dispersal and settlement in human-dominated landscapes and to reduce human–wolf encounters, with the goal of promoting coexistence.
Abstract
Assessing the behavioural responses of floating wolves to human presence is crucial for investigating the chance of wolf populations expanding into urbanised landscapes. We studied the movement ecology of three rehabilitated wolves in a highly human-dominated landscape (Po Plain, Italy) to explore wolf’s plasticity amid widespread human pressure. To reach this aim, we estimated individual 95% utilisation distributions (UD) after the release and inspected both 95% UDs and net squared displacements to identify individual movement patterns; tested for differences in movement patterns during day and night; and analysed the selection of resting sites during dispersal movement in a highly human-altered environment. Both the 95% UDs and step lengths were smaller for wolves settling in suitable areas than for those settling in more urbanised areas. All wolves exhibited strong temporal segregation with humans during all movement phases, particularly while dispersing across highly urbanised areas. Main roads and proximity to built-up areas were shown to limit wolves’ dispersal, whereas small-wooded patches that provide shelter during rest facilitated long-distance movements. This study provides important insights into wolf movement and settling in urban and peri-urban areas, providing critical knowledge to promote human–carnivore coexistence.
... The wolf and the European wildcat experienced very similar demographic scenarios in Italy, with protracted isolation south of the Alps and recurrent bottlenecks that made them sharply genetically differentiated from any other wolf or wildcat population [13,14]. Nowadays, both species are geographically re-expanding and numerically increasing trough the Peninsula, thanks to legal protection and their ecological plasticity [15,16], but they are still threatened by habitat fragmentation [16,17], accidental or illegal killings [18,19] and by anthropogenic hybridisation [20,21]. ...
Simple Summary
Wolves and European wildcats are two iconic predator species that can live in overlapping ecological contexts and also share their habitats with their domestic free-ranging relatives, increasing the risk of anthropogenic hybridisation and its possible deleterious consequences. By applying a multidisciplinary approach, we morphologically and molecularly analysed the cat remains found in a canid faecal sample collected in a forested area of central Italy. Individual multilocus genotypes of both predator and prey were identified turning out to be, respectively, a wolf showing traces of dog ancestry at autosomal microsatellite loci and a domestic cat.
Abstract
Non-invasive genetic sampling is a practical tool to monitor pivotal ecological parameters and population dynamic patterns of endangered species. It can be particularly suitable when applied to elusive carnivores such as the Apennine wolf (Canis lupus italicus) and the European wildcat (Felis silvestris silvestris), which can live in overlapping ecological contexts and sometimes share their habitats with their domestic free-ranging relatives, increasing the risk of anthropogenic hybridisation. In this case study, we exploited all the ecological and genetic information contained in a single biological canid faecal sample, collected in a forested area of central Italy, to detect any sign of trophic interactions between wolves and European wildcats or their domestic counterparts. Firstly, the faecal finding was morphologically examined, showing the presence of felid hair and claw fragment remains. Subsequently, total genomic DNA contained in the hair and claw samples was extracted and genotyped, through a multiple-tube approach, at canid and felid diagnostic panels of microsatellite loci. Finally, the obtained individual multilocus genotypes were analysed with reference wild and domestic canid and felid populations to assess their correct taxonomic status using Bayesian clustering procedures. Assignment analyses classified the genotype obtained from the endothelial cells present on the hair sample as a wolf with slight signals of dog ancestry, showing a qi = 0.954 (C.I. 0.780–1.000) to the wolf cluster, and the genotype obtained from the claw as a domestic cat, showing a qi = 0.996 (95% C.I. = 0.982–1.000) to the domestic cat cluster. Our results clearly show how a non-invasive multidisciplinary approach allows the cost-effective identification of both prey and predator genetic profiles and their taxonomic status, contributing to the improvement of our knowledge about feeding habits, predatory dynamics, and anthropogenic hybridisation risk in threatened species.
... Kojola et al., 2004;Nowak et al., 2011;Ję drzejewski et al., 2000;Wagner, 2012) have revealed that wolves typically predate large wild ungulates such as elk Alces alces, wapiti Cervus canadensis, reindeer Rangifer tarandus, and red deer Cervus elaphus, alongside medium sized species such as wild boar Sus scrofa, white-tailed deer Odocoileus virginianus and roe deer Capreolus capreolus. Of note is that consumption of livestock is generally low but is directly dependent on the abundance of wild ungulates, which are preferred as a more risk-free resource (Meriggi and Lovari, 1996;Imbert et al., 2016;Janeiro-Otero et al., 2020). Nevertheless, in some regions, seasonal resource availability, the selection of some prey types over others, and the demands of provisioning for young have allowed wolves to add berries (Homkes et al., 2020), smaller mammalian prey such as hare Lepus spp. ...
The wolf (Canis lupus L., 1754) has been a major keystone predator in the Palaearctic since the late Middle Pleistocene. Today, wolves display considerable dietary plasticity over their range, characterised by their preferential consumption of large and medium-sized wild ungulates, supplemented by smaller prey, including small mammals, fish and plant foods. However, the origins of this dietary flexibility (arguably the key to the wolf's long persistence) are poorly understood in terms of responses to different drivers over the course of the Pleistocene, including changing climate, environment and competition from other large carnivores. Here, in the first study using direct palaeodietary measurements on British fossil wolves, carnivore competitors and potential prey species, we compare stable isotope (δ¹³C and δ¹⁵N) evidence from three sites representing a late Middle Pleistocene interglacial (Marine Oxygen Isotope Stage [MIS] 7c-a, c.220-190ka BP), the early Devensian (last cold stage, MIS 5a, c.90-80ka BP) and the middle Devensian (MIS 3, c. 60-25ka BP). The results reveal clear patterns of changing wolf prey choice through time. Notwithstanding issues of collagen preservation obscuring some dietary choices in the oldest samples, both small and large prey (hare, horse) were taken by wolves in the MIS 7c-a interglacial, large prey only (reindeer, bison) during MIS 5a and a broader range of large prey items (horse, woolly rhinoceros, bison) during MIS 3. The results also reveal two further important aspects: (1) that where wolves and spotted hyaenas co-existed, they occupied the same dietary niche and the former was not outcompeted by the latter, and (2) that the stable isotope evidence indicates prey choices during MIS 7c-a and MIS 3 that are not in synchrony with palaeodietary reconstructions from previous studies based on wolf cranio-dental morphology. This establishes for the first time a likely lag between changing predatory behaviour and morphological response but is interestingly not seen in the wolves from MIS 5a, where the prey choices are echoed by the cranio-dental morphology.
... This is a good perspective for Germany where sheep losses are still on the rise as the wolf recolonization is "young, " but they are expected to recede over time with the wolf population approaching its carrying capacity (Fechter and Storch, 2014) and farmers protecting their livestock and becoming more tolerant (Cretois et al., 2021). Imbert et al. (2016) also report that livestock protection and stabilization of wolf packs lead to the decline of livestock losses over time. ...
Recovery of predator populations triggers conflicts due to livestock depredation losses, particularly in Germany where the wolf (Canis lupus) population grows exponentially and livestock (especially sheep) losses raise public concerns and motivate the authorities to control wolf numbers. Yet, the effects of wolf numbers and alternative factors, such as abundance of prey and livestock, on livestock losses in this country are not investigated. In this study, we collected and analyzed data on the numbers of reproductive units of wolves (packs and pairs together) as a surrogate of adult wolf numbers, sheep killed by wolves, living sheep, red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild boar (Sus scrofa) in every German state and year from 2002 to 2019. We applied a negative binomial Generalized Linear Mixed Model (GLMM) to estimate the effects of these predictors on the numbers of sheep killed by wolves. We also examined the relationships between the percentages of killed/living sheep and the numbers of living sheep. Ranking of 63 models based on the Akaike information criterion revealed that sheep losses were determined by state, year, and number of living sheep, not by wolf numbers, at high precision and accuracy. The number of sheep killed by wolves increased consistently by 41% per year and by 30% for every additional 10,000 sheep, mainly in the north where most wolf territories are concentrated. This means that sheep are protected insufficiently and/or ineffectively. The percentages of killed/living sheep consistently increased by 0.02–0.05% per state and year, with the maximum percentage of 0.7%, on a backdrop of decreasing numbers of living sheep. In conclusion, we demonstrate that sheep losses in Germany have been driven by the expansion of the wolf population, not by wolf numbers, and by the number of sheep available. We suggest that Germany’s wolf conservation policy should focus on alternative non-lethal interventions, enforcement and standardization of intervention monitoring, and promotion of wolf tolerance rather than on lethal control of wolf population size.
... This supports the assertion that wolves prefer wild ungulates if they are sufficiently abundant (e.g. Imbert et al., 2016;Jedrzejewski et al., 2012;Meriggi et al., 2015). When wolves predate on horses in southern Europe, they usually target unprotected animals in open pastures (e.g. ...
We report on the first results of our efforts to generate validated data on horse-wildlife interactions in northern and eastern Germany. Data were gathered from pastures hosting horses within or nearby areas been frequented by territorial wolves. Including recommendations adressed to equestrians concerned.
... distance to forest; Treves et al., 2011), meteorological conditions (Towns et al., 2009), breed of domestic animals (Bassi et al., 2021), herd size (Dar et al., 2009), status of local carnivore populations (e.g. stability of wolf packs; Imbert et al., 2016;Santiago-Avila et al., 2018), historic presence of large carnivores in the area (Linnell, 2013), carnivores' space use (Melzheimer et al., 2020) and their habituation to human presence (Majić Skrbinšek and Krofel, 2015). Due to small sample sizes and limited availability of such information in LIFE project reports, we were not able to evaluate these factors on the effectiveness of prevention measures used in these projects. ...
Governments around the world invest considerable resources to reduce damages caused by large carnivores on human property. To use these investments more efficiently and effectively, we need to understand which interventions successfully prevent such damages and which do not. In the European Union, the LIFE program represents by far the largest financial instrument to help EU Member States with the implementation of conservation activities, including mitigation of damages caused by large carnivores. However, we currently lack information about the effectiveness of this funding program in reducing carnivore damages. We reviewed 135 LIFE projects dealing with large carnivores between 1992 and 2019 to provide an overview of the use of damage prevention methods and evaluate their functional and perceived effectiveness. Methods evaluated ranged from non-lethal and lethal interventions, to information dissemination and compensation schemes. The largest number of the projects was focused on grey wolf (Canis lupus) and brown bears (Ursus arctos) in the Mediterranean countries and in Romania. Electric fences were reported as the most successful method for reducing damages by large carnivores, and most of the non-lethal methods used showed at least moderate effectiveness. However, standards of measuring and reporting effectiveness were in general relatively low, which limits our ability to measure actual impact. We urge project managers and evaluators to improve these standards, as well as the dissemination of the project results. We provide a list of recommendations for improving measuring and reporting success of implemented interventions for the benefit of future projects aimed to reduce damages caused by wildlife.
... In Pakistan, wolves are endangered and persecution remains one of the biggest hurdles to recovery because of livestock depredation and its associated impact on livelihoods (Khan et al., 2019). Hence, public awareness, strong legal protection, supportive media coverage and furthered ecological research in Pakistan could aid in supporting the conservation of this keystone species (Imbert et al., 2016). In consideration of the lack of knowledge on wolves in Pakistan, we undertook this project to further research on these unique wolves' biology and ecology across Pakistan. ...
The gray wolf (Canis lupus) is a less studied wide-ranging endangered carnivore in
Pakistan. The current investigation is the first to report their body morphometrics
and chemical immobilization in Pakistan. Body morphometrics was examined for
12 wolves by measuring 15 variables. The majority of the 12 wolves had body
weights that were more similar or slightly higher than the weights of Indian wolves
from Central India. Principal Component Analysis (PCA) revealed that the wolves
from the southern lowland region have differing morphology, independent of body
size compared to the wolves from other regions of Pakistan. To record the body
morphometrics, wolves were immobilized using Zoletil™-50 (Z) (n=6) and
Xylazine-Ketamine hydrochloride (X-K) combination (n=3). The wolves were
immobilized by using drug doses 5-6 mg/kg for Z, and 1.25 mg/kg for X and 2-3
mg/kg for K. The first sign (minutes) of anesthesia was noted after 3.15±1.9 for Z
and 4.97±2.3 for X-K combination. The recumbency time was 7.7±2.5 for Z and
11.7±3 for X-K combination. The sign of recovery was recorded at 40.4±13.5 for Z
and 34.1±2.4 for X-K combination, while the sedation duration was recorded at
45.3±12.5 for Z and 39.6±3.5 for X-K combination. These results suggest that Z
induced quicker induction, more profound recumbency and swifter recovery than XK combination. Additionally, physiological parameters including rectal temperature,
respiration, heart rate and palpebral and capillary reflexes with both combinations
remained within the safe ranges.
... distance to forest; Treves et al., 2011), meteorological conditions (Towns et al., 2009), breed of domestic animals (Bassi et al., 2021), herd size (Dar et al., 2009), status of local carnivore populations (e.g. stability of wolf packs; Imbert et al., 2016;Santiago-Avila et al., 2018), historic presence of large carnivores in the area (Linnell, 2013), carnivores' space use (Melzheimer et al., 2020) and their habituation to human presence (Majić Skrbinšek and Krofel, 2015). Due to small sample sizes and limited availability of such information in LIFE project reports, we were not able to evaluate these factors on the effectiveness of prevention measures used in these projects. ...
Governments around the world invest considerable resources to reduce damages caused by large carnivores on human property. To use these investments more efficiently and effectively, we need to understand which interventions successfully prevent such damages and which do not. In the European Union, the LIFE program represents by far the largest financial instrument to help EU Member States with the implementation of conservation activities, including mitigation of damages caused by large carnivores. However, we currently lack information about the effectiveness of this funding program in reducing carnivore damages. We reviewed 135 LIFE projects dealing with large carnivores between 1992 and 2019 to provide an overview of the use of damage prevention methods and evaluate their functional and perceived effectiveness. Methods evaluated ranged from non-lethal and lethal interventions, to information dissemination and compensation schemes. The largest number of the projects was focused on grey wolf ( Canis lupus ) and brown bears ( Ursus arctos ) in the Mediterranean countries and in Romania. Electric fences were reported as the most successful method for reducing damages by large carnivores, and most of the non-lethal methods used showed at least moderate effectiveness. However, standards of measuring and reporting effectiveness were in general relatively low, which limits our ability to measure actual impact. Therefore we urge project managers and evaluators to improve these standards, as well as the dissemination of the project results. We provide a list of recommendations for improving measuring and reporting success of implemented interventions for the benefit of future projects aimed to reduce damages caused by wildlife.
Article impact statement
Electric fences were reported as the most effective method to prevent large-carnivore damages and are recommended for future use.
The primary food source of wolves is wild ungulates, mainly deer, but farm animals, including sheep, may also fall prey to these predators. The aim of this study was to characterize the scale of wolf attacks on flocks of sheep of native breeds in Poland in 2015–2020, with particular emphasis on mountainous regions, considered to be the grazing areas most at risk of attacks by wolves. The study included an analysis of documentation submitted by breeders reporting replacements of sheep (replacing one sheep with another as a result of death, selection, etc.) due to wolf attacks. As expected, the analysis showed that most attacks took place in the south of the country (76.9%), although some occurred in other parts of Poland as well, especially in the northeast. The most frequently attacked breed was the Podhale Zackel, kept in mountainous regions, areas which are also inhabited by wolf packs. The greatest number of attacks occurred in July and August. Ewes were most frequently attacked, followed by lambs, and finally rams. Over the years of the study, there was an increase in the number of cases of sheep attacked by wolves, which means that methods of protecting flocks should be improved.
Il s'agit d'un rapport d'expert indépendant (rapport d’étude) publié par OneVoice sur l'histoire et l'état actuel du pastoralisme en France, et sur la manière dont ses pratiques sont liées à l'environnement et aux populations de grands carnivores en France, en particulier les loups et les ours. Le rapport est basé sur une revue de la littérature narrative, ainsi que sur des analyses bibliométriques et médiatiques.
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This is an independent expert report published by OneVoice regarding the history and current status of pastoralism in France, and how its practices relate to the environment and large carnivore populations in France, particularly wolves and bears. The report is based on a narrative literature review, alongside bibliometric and news media analyses.
As wolves recolonize human-modified landscapes across the Old World, management programs aiming to mitigate livestock depredation while preserving wolf populations are falling short. The combination of human activity, habitat fragmentation, and mixed land use creates complex challenges for conservation management. Recolonized by wolves in the 1970s, the Golan Heights in northern Israel poses a dense mosaic landscape of risk, comprising grazing pastures, military zones, nature reserves, agriculture, and minefields. Today it contains one of the highest densities of wolves worldwide. While wolves are protected by law, the government maintains an active lethal management program by which about 25% of the population is culled annually. To evaluate this program’s effectiveness, we used 60 motion-triggered camera traps over 5,997 nights to monitor wolf activity across the Golan Heights. Using long-term culling and landscape data, we assessed the spatiotemporal responses of single wolves and wolf packs to culling pressure, land use and human activity. We found a positive relationship between culling pressure and single wolf activity, while pack responses to culling varied over a gradient of land uses. Single wolves utilized high-risk areas near cattle despite intense culling, while packs dominated protected, no-culling zones. Overall, culling did not deter wolves, singles or packs, from conflict zones; all zones were occupied by wolves. However, wolves shifted temporally to avoid daytime human activity and were predominantly nocturnal in high-culling areas. Understanding wolves’ responses to lethal management across diverse anthropogenic pressures offers lessons for other areas, particularly in Europe, currently undergoing wolf recolonization in similar landscapes.
Simple Summary
We conducted a study in central Spain to understand the dietary habits of the Iberian wolf. Our aim was to evaluate the extent to which they prey on domestic ungulates. We analyzed the composition of their diet by examining prey hairs found in 671 wolf scat samples collected between 2017 and 2021. The wolves predominantly consumed wild ungulates rather than domestic ones. Among their preferred prey were wild boar and roe deer. Although their diet varied with seasons, years, and forest regions, a preference for wild ungulates over domestic ones remained consistent.
Abstract
The Iberian wolf (Canis lupus signatus) is recolonizing historical distribution areas after decades of absence. As in other human-dominated landscapes, finding a balance to protect this species by favoring recolonization and mitigating human–wildlife conflicts is a challenge. Since wolves are often generalist opportunistic predators, we studied their diet composition in central Spain to evaluate the consumption of domestic ungulates and provide reliable data that could help local authorities to deal with the current wolf–cattle ranchers conflict and coexistence. Diet composition (% prey occurrence, % prey ingested biomass) was analyzed through the identification of prey hairs present in 671 scats collected between 2017 and 2021. The wolves fed more on wild ungulates (82% occurrence) than domestic ones (18%). Wild boar (Sus scrofa, 44% occurrence) and roe deer (Capreolus capreolus, 35%) were the most consumed prey. The wolves positively selected these two species. The wolves’ diets varied between seasons, years, and forest regions, but a diet based on wild ungulates predominated over domestic ones. Food niche breadth showed variations depending on seasons and years. Preserving the availability and diversity of wild ungulates may favor reducing livestock attacks and would be an achievable goal that would help to conserve this species and reduce conservation conflicts.
Wolves returned to Germany in 2000, leading to fear in German horse owners that their horses could be in danger of wolf attacks or panic-like escapes from pastures when sighting wolves. However, reports from southern European countries indicate that wolf predation on horses diminishes with increasing presence of wildlife. Therefore, we conducted a long-term, filed observation between January 2015 and July 2022 on 13 non breeding riding horses, mares and geldings, kept permanently on two pastures within the range of wildlife and a stable wolf pack with annual offspring. Wildlife cameras at the fences of the pastures made 984 times recordings of wolves and 3151 times recordings of wildlife in and around the pastures. Between 1 January 2022 and 23 March 2022 we observed two stable horse groups. Pasture 1 was grazed by five horses of mixed breed, four mares and one gelding, with the median age of 8 years (min. = 6y, max. = 29y). Pasture 2 was grazed by eight heavy warmbloods and draught horses, three mares and five geldings, with the median age of 16 years (min. = 13y, max. = 22y). During this period no wolf was recorded at pasture 2, but wild boar several times, whereas at pasture 1, wolves were recorded 89 times, and for the wildlife mostly hare. Wolves may have avoided pasture 2 because of the presence of wild boar or because the large group of older, heavy breed horses may have formed a stable, protective group. The latter needs to be confirmed in a follow-up field observation, which records anti-predator behavior and welfare indicators in horses. In conclusion, wolves did not attack the mature horses on pastures with plenty of wildlife and the horses did not respond to the presence of wolves with visible signs of reduced welfare or panic. This indicates that wolves may prefer to prey on easily accessible wildlife around and at horse pastures and that Central European horses become accustom to the presence of non-hunting wolves.
The gray wolf (Canis lupus) expanded its distribution in Europe over the last few decades. To better understand the extent to which wolves could re-occupy their historical range, it is important to test if anthropization can affect their fitness-related traits. After having accounted for ecologically relevant confounders, we assessed how anthropization influenced i) the growth of wolves during their first year of age (n = 53), ii) sexual dimorphism between male and female adult wolves (n = 121), in a sample of individuals that had been found dead in Italy between 1999 and 2021. Wolves in anthropized areas have a smaller overall variation in their body mass, during their first year of age. Because they already have slightly higher body weight at 3–5 months, possibly due to the availability of human-derived food sources. The difference in the body weight of adult females and males slightly increases with anthropization. However, this happens because of an increase in the body mass of males only, possibly due to sex-specific differences in dispersal and/or to “dispersal phenotypes”. Anthropization in Italy does not seem to have any clear, nor large, effect on the body mass of wolves. As body mass is in turn linked to important processes, like survival and reproduction, our findings indicates that wolves could potentially re-occupy most of their historical range in Europe, as anthropized landscapes do not seem to constrain such of an important life-history trait. Wolf management could therefore be needed across vast spatial scales and in anthropized areas prone to social conflicts.
Gray wolf Canis lupus predation on domestic dogs Canis familiaris is a considerable wolf–human conflict issue in several regions of Europe and North America but it has not been well documented in the scientific literature. Livestock depredations by wolves may be related to the abundance of wild prey. Regardless of the presumed motivations of wolves for attacking dogs (likely due to interference competition and predation), the abundance of wild prey populations may also influence the risk of wolf attacks on dogs. We examined whether the annual number of fatal attacks by wolves on dogs was related to the abundance of primary prey, including wild boar Sus scrofa and roe deer Capreolus capreolus in Estonia, as well as the abundance of moose Alces alces in Finland. Statistical models resulted in significant negative relationships, thus providing evidence that the risk of attacks in both house yards (Estonia) and hunting situations (Finland) was highest when the density of wild prey was low. Wild ungulates cause damage to agriculture and forestry, but they seem to mitigate conflicts between wolves and humans; therefore, it is necessary to develop a holistic, multispecies management approach in which the importance of wild ungulates for large carnivore conservation is addressed.
Applying research results to new locations and situations can be confounded by differences in the geographic context between the original and the applied study sites. Replication studies and meta‐analyses may be similarly hindered. We investigated how often canid management research reports (e.g., journal articles, conference proceedings) included information on historical/current lethal control, alternative prey availability, landscape features, and seasonal and settlement characteristics. We included experimental research published between 1970 and 2018, focusing on livestock depredations by wolves and coyotes in North America. Reporting on contextual factors was highly variable; seasonal context was included in 83% of research findings; human settlement characteristics were reported in only 8%. Contextual information was more common in journal versus grey literature, and in reports with academic versus government‐affiliated primary authors. Discussions of the effects of contextual factors on livestock depredation mitigation strategies were underdeveloped. Yet, geographic context of research is vital; it can alter animal behaviour and reduce the efficacy of applied mitigation. We suggest reporting guidelines to improve comparisons and meta‐analysis opportunities, which may enhance comparisons and management decision making.
Resource selection analyses based on detection data are widely used to parametrize resistance surfaces used to identify ecological corridors. To successfully parametrize resistance, it is crucial to decouple resident and disperser behaviours yet to date connectivity studies using detection data have not addressed this issue. Here, we decoupled data of resident and dispersing wolves by analysing detection data collected within a natural corridor crossing a human dominated plain in Italy. To decouple residents and dispersers, we ran a Kernel Density analysis to investigate whether clusters of wolf detection points characterized by sharply higher points’ density exist and checked whether the areas outlined by these clusters (core areas) hold specific characteristics. Habitat selection analysis was then performed to compare the intensity of habitat selection carried out by putative residents and dispersers. We identified a high-density cluster of 30 detection points outlining a small core area stably located in the central part of the park. The dramatic differences of the R² and the AUC of the habitat selection models performed inside (R² = 0.506; AUC = 0.952) and outside (R² = 0.037; AUC = 0.643) the core area corroborated the hypothesis that the core area effectively encloses detection points belonging to residents. Our results show that through simple space use analyses it is possible to roughly discriminate between detection points belonging to resident-behaving and disperser-behaving individuals and that habitat selection models separately performed on these data have extremely different results with strong possible effects on resistance surfaces parametrized from these models.
Human-wolf (Canis lupus) conservation conflicts in Europe have increased as wolf presence has expanded. Understanding how different stakeholders perceive coexistence, especially in locations identified as ecologically important for wolves is necessary to minimize conflict. We conducted a survey in an area of northwest Italy identified as a critical corridor linking separate, cross-boundary populations. The objective was to understand how stakeholder identities, social demographics, communication, and exposure influence the success of coexistence strategies. The study found conservationists and, significantly, hunters were most positive about wolves, while farmers were least tolerant, irrespective of exposure. Tolerance also correlated positively with higher levels of formal education and engagement with science-based knowledge. In contrast, less tolerant attitudes were influenced more by informal knowledge discourses and age. The article concludes that coexistence strategies need to be inclusive, reflexive and adapted to the specific circumstances of different stakeholder types.
The gray wolf (Canis lupus) is one of the most conflictual mammals in Europe. Wild boar (Sus scrofa) are an essential part of gray wolf diet in central Europe, but after the emergence of African swine fever (ASF) in Europe, a sharp decline of the wild boar occurred. We examined how the wild boar population decline, due to African swine fever outbreak and mitigation efforts, affected the number of livestock killed by wolves in Poland using long-term data on wild ungulate and livestock population sizes and wolf-induced mortality between 2013 and 2019. We examined the influence of multiple factors on livestock kill rate, and the influence of wild boar population declines on the number of Cervidae killed by wolves using linear mixed models. We also explored the possibility of predicting a dramatic decrease in the wild boar population based on livestock depredation patterns. The number of livestock killed by wolves decreased with wild boar and roe deer (Capreolus capreolus) population size, and increased with red deer (Cervus elaphus) population size. A decline in the wild boar population was significantly correlated with an increase in the number of both red and roe deer killed by wolves. A drastic decline of wild boar population (over 30%) could be predicted by the numbers of livestock killed by wolves. Our study confirms that large changes in the number of naturl prey can increase livestock depredation, although these changes may be difficult to detect when the fluctuations in the numbers of natural prey are smaller. In our opinion, this indicates that the assessment of factors influencing livestock depredation should consider historical changes in prey dynamics. We suggest managers and conservationists use the predator population as a 'first alert system' for indirect monitoring of prey species. In this system, a sudden increase in wolf attacks on livestock across a large area of should trigger an alarm and prompt verification of the number of natural prey in the environment.
Habitat use and diet of wolves Canis lupus were examined in a mountainous area in the northern Apennines (northern Italy) from December 1987 to March 1989. Wolf signs were looked for along 22 transects representative of the different habitat types of the study area in order to define seasonal differences in habitat use. Scats were collected and analysed to identify the main food items used by wolves in each season. Changes in range surface area were recorded in different seasons in relation to food availability and territoriality of the wolves.
A major challenge in carnivore conservation worldwide is identifying priority human–carnivore conflict sites where mitigation efforts would be most effective. Spatial predation risk modeling recently emerged as a tool for predicting and mapping hotspots of livestock depredation using locations where carnivores attacked livestock in the past. This literature review evaluates the approaches and applications of spatial risk modeling for reducing human–carnivore conflict and presents a workflow to help conservation practitioners use this tool. Over the past decade 18 studies were published, most which examined canid and felid (10 and 8 studies on each group, respectively) depredation on cattle (14) and sheep (12). Studies employed correlation modeling, spatial association and/or spatial interpolation to identify high-risk landscape features, and many (but not all) validated models with independent data. The landscape features associated with carnivore attacks related to the species (carnivore and prey), environment, human infrastructure and management interventions. Risk maps from most studies (14) were used to help livestock owners and managers identify top-priority areas for implementing carnivore deterrents, with some efforts achieving >90 % reductions in attacks. Only one study affected policy, highlighting a gap where risk maps could be useful for more clearly communicating information to assist policymakers with large-scale decisions on conflict. Studies were used to develop a six-step workflow on integrating risk modeling into conservation. This review reveals a need for future predation risk modeling to focus more on validating models, accounting for feedbacks and impacting conflict-related policy in order to reliably improve the mitigation of human–carnivore conflict globally.
The Casentinesi Forests, in the northern Apennines, harbour a rich community of wild ungulates, with the wolf representing the largest predator in the area. Between 1993 and 2000, wolf pack distribution in the area was monitored and estimates of pack size were obtained by wolf-howling surveys, snow-tracking, and occasional observations. Three to five packs were detected yearly, with sizes averaging 4.2 � 0.9 wolves (maximum 7). The overall density in the area was 4.7 wolves per 100 km 2 with an average distance between adjacent packs of 11.1 km. The high wolf density in the Casentinesi Forests is mostly related to abundance and size of wild prey. In this, like in other areas at low latitudes, wolf density depends mainly on the number of packs, as pack size is rather small and recruitment limited by early dispersal and high mortality. Three homesites used in several years by resident packs were discovered. Homesite fidelity and pack reproductive success were higher in fully protected rather than harvested areas. Establishing a network of protected areas with high ungulate diversity and abundance is proposed as the main factor for allowing a full recovery of the wolf population in Italy. 2004 Elsevier Ltd. All rights reserved.
Information on population parameters is rarely collected from carcasses. This method can be particularly useful – with limitations – when protected species are involved (e.g. the grey wolf Canis lupus in Italy). Local data on population structure, reproduction, survivorship and causes of mortality are necessary to build reliable conservation models to assess the state of a population and to predict its evolution. On the other hand, ‘best guesses’ or data from ecologically different areas have often been used to build population viability analysis and other conservation-oriented models. A sample of 154 wolf carcasses was found, collected and analysed from 1991 to 2001 in central-eastern Italy, the historic core of the wolf distribution range. Collision with a vehicle was the main cause of death in both sexes; however, road kills may be biased with a greater detectability, and we treated our data accordingly. Road kills were concentrated on the younger (≤4 years old) age classes, whereas fully adult wolves died mainly because of poaching, intraspecific strife and pathologies. Cubs and subadults (≤2 years old) showed a mortality peak in November/December, at the beginning of the dispersal period, whereas adults died mainly in January/February (mating season). The population structure of our sample of wolf carcasses appeared to be well balanced, although perinatal and cub mortality was underestimated. The sex ratio was 1:1 in the younger age classes and 1:0.7 in the older age classes. Only 20.7% of females, 2–6 years old, showed signs of reproduction; placental scar and embryo number varied from one to seven (mean, 4.4) per individual. Survivorship theoretical curves indicated a fair survival of cubs and subadults, but a steep decline as wolves approached maximum life span (9 years old). Our data and other published data on food habits and genetic features of the wolf in central-eastern Italy suggest that, despite ongoing heavy human-induced losses, this predator has fully recovered in the last 30 years from the brink of extinction.
Predation by wolves Canis lupus Linnaeus, 1758 in ungulate communities in Europe, with special reference to the multi-species system of Bialowieza Primeval Forest (Poland/Belarus), was assessed on the basis results of original research and literature. In hisstorical times (post-glacial period), the geographical range of the wolf and most ungulate species in Europe decreased considerably. Community richness of ungulates and potential prey for wolves, decreased over most of the continent from 5-6 species to 2-3 species. The wolf is typically an opportunistic predator with a highly diverse diet; however, cervids are its preffered prey.
Long-term changes of wolf feeding habits have been recorded in several parts of
its European range and were related to the increase of wild ungulate populations. An
increased use of wild prey often is linked to a decrease of predation on livestock, and
consequently it can reduce wolf–human conflict and make for easier predator conservation.
We analysed wolf diet from scats collected from 2007 to 2012 in a 927.3-km2
area of the Northern Apennines (Northern Italy) to compare the present diet to that
recorded in the late 1980s when the area was colonised by wolves, and to examine
short-term changes of wolf feeding habits together with annual and seasonal variations
of wild and domestic ungulate selection, possibly related to the increase of wild
ungulate abundance. We calculated the availability proportions of wild ungulate
species from the presence signs recorded on itineraries and those of livestock species
from the data on farms of veterinary services. We identified 17 kinds of prey, pooled
into six categories, of which wild ungulates (70%) and livestock (20%) were the most
important. During the study period, we observed a reduction in the diet breadth of
wolves, to a greater use of wild ungulates, depending on the increase in the availability
of wild prey species. Significant seasonal variations were observed in the trophic habits
of the wolf, but generally there was a wide use of wild ungulates, especially wild boars.
Livestock was mainly used in summer, which corresponds to the period of greatest
presence of domestic ungulates on the pastures. Considering the study period, a fast
response of wolf feeding habits to the increase of wild ungulate species resulted,
together with changes in selection patterns. The change in diet towards a greater use
of wild ungulates is an important step in the recovery and conservation of the wolf.
Mortality due to illegal killing is still today one of the main threats to wolf
conservation, and an effective management of the conflict between wolf presence and
husbandry is a key element for species conservation. The research was aimed at
identifying the farm characteristics and the environmental factors that influence
predation, and at formulating predictive models of predation risk. We collected and
analysed the data on official predation events that occurred during the period
2005–2012 in an area of the northern Apennines, and on the characteristics of livestock
farms recorded at the veterinary services. Furthermore, we mapped the grazing
areas used by livestock farms and measured 23 variables of the pastures. Our results
showed that the majority of predation events were upon cattle, and that grazing
management significantly influences the number of predation events. In particular,
the pastures that suffered predation were those in which births occur directly on the
pasture, those that had at least one period of free grazing during the year, and cattle
farms that were lacking in any preventive methods. The number of killed animals per
event was higher for sheep than for cattle and goats and increased progressively
during the study period. Predation risk increases if the farms practice free grazing
at least for a period during the year, if they are not protected by any preventive
method and if the degree of surveillance is regular or constant. The risk of predation
also increases with the increasing complexity of pasture shape, the decrease in the
percentage of coniferous forest and if the pasture is exposed to the north. The model
of predation risk showed that 56.6% of the pastures in the study area are potentially
exposed to wolf predation, and it allowed us to identify the grazing areas where it is
worthwhile intervening with preventive methods.
The diet strategy of the wolf in Europe is reviewed on the basis of 74 basic and 14 additional literature
sources. The comparative analysis reveals clear dependence on the latitude (and, therefore, on the changing
environmental conditions) correlated with the wild ungulate abundance and diversity. Following a
geographic pattern, the wolf is specialised on different species of ungulates: moose and reindeer in Scandinavia,
red deer in Central and Eastern Europe and wild boar in Southern Europe. Where this large prey
is taken, the roe deer is hunted with almost the same frequency in every region. The wolf diet in Europe
shows two ecological adaptations formed by a complex of variables: 1. Wolves living in natural habitats
with abundance of wild ungulates feed mainly on wild prey. 2. In highly anthropogenic habitats, with low
abundance of wild prey, wolves feed on livestock (where husbandry of domestic animals is available) and
take also a lot of plant food, smaller prey (hares and rodents) and garbage food. The frequency of occurrence
of wild ungulates in the diet of wolves in North Europe varies from 54.0% in Belarus to 132.7% in
Poland, while that of livestock is in the range from 0.4% in Norway to 74.9% in Belarus. In South Europe,
the frequency of occurrence of wild prey varies from 0% in Italy and Spain to 136.0% in Italy, while of domestic
ungulates ranges between 0% and 100% in Spain. The low density or lack of wild prey triggers the
switch of the wolf diet to livestock, plant food (32.2-85% in Italy) or even garbage (up to 41.5% in Italy).
The conservation of large carnivores is a formidable challenge for biodiversity conservation. Using a data set on the past
and current status of brown bears (Ursus arctos), Eurasian lynx (Lynx lynx), gray wolves (Canis lupus), and wolverines (Gulo gulo) in European countries, we show that roughly one-third of mainland Europe hosts at least one large carnivore species, with
stable or increasing abundance in most cases in 21st-century records. The reasons for this overall conservation success include
protective legislation, supportive public opinion, and a variety of practices making coexistence between large carnivores
and people possible. The European situation reveals that large carnivores and people can share the same landscape.
Predator control and sport hunting are often used to reduce predator populations and livestock depredations, – but the efficacy of lethal control has rarely been tested. We assessed the effects of wolf mortality on reducing livestock depredations in Idaho, Montana and Wyoming from 1987–2012 using a 25 year time series. The number of livestock depredated, livestock populations, wolf population estimates, number of breeding pairs, and wolves killed were calculated for the wolf-occupied area of each state for each year. The data were then analyzed using a negative binomial generalized linear model to test for the expected negative relationship between the number of livestock depredated in the current year and the number of wolves controlled the previous year. We found that the number of livestock depredated was positively associated with the number of livestock and the number of breeding pairs. However, we also found that the number of livestock depredated the following year was positively, not negatively, associated with the number of wolves killed the previous year. The odds of livestock depredations increased 4% for sheep and 5–6% for cattle with increased wolf control -up until wolf mortality exceeded the mean intrinsic growth rate of wolves at 25%. Possible reasons for the increased livestock depredations at #25% mortality may be compensatory increased breeding pairs and numbers of wolves following increased mortality. After mortality exceeded 25%, the total number of breeding pairs, wolves, and livestock depredations declined. However, mortality rates exceeding 25% are unsustainable over the long term. Lethal control of individual depredating wolves may sometimes necessary to stop depredations in the near-term, but we recommend that non-lethal alternatives also be considered. OPEN ACCESS
1994. Status and diet o f wolves in Greece. Acta theriol. 39: 411—416. The stomach contents of 32 wolves Canis lupus Linnaeus, 1758 collected through out Greece during autumn and winter 1991 and 1992 were examined for prey remains. O f these 28 stomachs contained measurable quantities of food. Of the 19 species that were found in the w olf diet 12 species were mammals. The main food items were goats, sheep, pigs and cows. Large wild herbivores did not play a significant role in the w olf diet. The observed consumption of grasses and fruits are important when they are available. The maximum food content in the examined stomachs was 1860 g.
After centuries of population decline and range contraction, gray wolves (Canis lupus) are now expanding in Europe. Understanding wolf social structure and population dynamics and predicting their future range expansion is mandatory to design sound conservation strategies, but field monitoring methods are difficult or exceedingly expensive. Non-invasive genetic sampling offers unique opportunities for the reliable monitoring of wolf populations. We conducted a 9-year long monitoring program in a large area (approximately 19,171 km2) in northern Italy, aiming to identify individuals, estimate kinship, reconstruct packs, and describe their dynamics. Of 5,065 biological samples (99% scats), we genotyped and sexed 44% reliably using 12 unlinked autosomal microsatellites, 4 Y-linked microsatellites and a diagnostic mtDNA control-region sequence. We identified 414 wolves, 88 dogs, and 16 wolf x dog hybrids. Wolves in the study area belonged to at least 42 packs. We reconstructed the genealogy of 26 packs. The mean pack size was 5.6 ± 2.4 (± SD), including adoptees, with a mean minimum pack home range of 74 ± 52 km2. We detected turnovers of breeding pairs in 19% of the packs. Reproductive wolves were unrelated and unrelated dispersers founded new packs, except for 1 pack founded by a brother-sister pair. We did not detect multiple breeding females in any packs. Overall, the population was not inbred. We found significant isolation-by-distance and spatial autocorrelation, with non-random genetic structure up to a distance of approximately 17 km. We detected 37 dispersers, 14 of which became breeders in new or already existing packs. Our results can be used to model habitat uses by wolves, to estimate survival rates, to predict future expansion of the wolf population, and to build risk maps of wolf-human conflicts.
After centuries of range contraction and demographic declines wolves are now expanding in Europe, colonizing regions from where they have been absent for centuries. Wolf colonizing the western Alps originate by the expansion of the Italian population. Vagrant wolves of Italian and Dinaric-Balkan origins have been recently observed in the Eastern Alps. In this study we compared the genetic structure of wolf populations in Italy and Croatia, aiming to identify the sources of the ongoing recolonization of the Eastern Alps. DNA samples, extracted from 282 Italian and 152 Croatian wolves, were genotyped at 12 autosomal microsatellites (STR), four Y-linked STR and at the hypervariable part of the mitochondrial DNA control-region (mtDNA CR1). Wolves in Croatia and Italy underwent recent demographic bottlenecks, but they differ in genetic diversity and population structure. Wolves in Croatia were more variable at STR loci (NA = 7.4, HO = 0.66, HE = 0.72; n = 152) than wolves in Italy (NA = 5.3, HO = 0.57, HE = 0.58; n = 282). We found
four mitochondrial DNA (mtDNA CR1) and 11 Y-STR haplotypes in Croatian wolves, but only one mtDNA CR1 and three Y-STR haplotypes in Italy. Wolves in Croatia were subdivided into three genetically distinct subpopulations (in Dalmatia, Gorski kotar and Lika regions), while Italian wolves were not sub-structured. Assignment testing shows that the eastern and central Alps are recolonized by wolves dispersing from both the Italian and Dinaric populations. The recolonization of the Alps will predictably continue in the future and the new population will be genetically admixed and very variable with greater opportunities for local adaptations and survival.
A composite approach mixing design-based and model-based inference is considered for analyzing line-transect or point-transect data. In this setting, the properties of the animal abundance estimator stem from the sampling scheme adopted to locate transects or points on the study region, as well as from the modeled detection probabilities. Moreover, the abundance estimation can be viewed as a “generalized” version of Monte Carlo integration. This approach permits to prove the superiority of the stratified placement of transects or points (based on a regular tessellation of the study region) over the uniform random placement. Even if the result was already established for the fixed-area sampling, i.e., when a perfect detection takes place, it was lacking in distance sampling. Comparisons with other widely-applied schemes pursuing an even placement of transects or points are also considered.
We reviewed 20 studies on wolf diet in Italy, to relate the changes in diet composition to the increase of wild ungulate population in Italy. Researches covered the period from 1976 to 2004 and the whole range of wolves from southern Apennines to western Alps. We used the frequency of occurrence of seven food categories and of the wild ungulate species occurring in the diet. Estimates of wild ungulate populations were obtained from the literature and we extrapolated their trend in the period considered. Differences among geographic areas (south-central Apennines, northern Apennines, and western Alps) were tested by nonparametric multivariate analysis of variance, while the trends of the wild ungulate and livestock use and of diet breadth were analysed by regression and curve-fit analyses. We used the same method to support the relationships between the use and availability of wild ungulates. Wolves preyed on wild herbivores more in the northern Apennines and in the western Alps than in the southern Apennines; the contrary was the case for livestock. Among wild ungulate species, wild boar, roe deer and red deer were the main prey of the wolf. The occurrence of wild ungulates in the wolf diet increased from 1976 to 2004 together with a decrease of livestock; the increase was mainly due to roe deer, red deer and chamois. The results of scat analysis in the province of Genoa showed an increase of the occurrence of wild ungulates from 1987 to 2005, in particular roe deer and fallow deer. Wolves in Italy seem to select wild ungulates over domestic ones where the former are available with rich and diversified guilds and abundant populations.
We investigated the patterns of wild ungulate selection by wolves in an 860 km study area of the Northern Apennines (North Italy), in order to detect seasonal variation in wolf diet and changes of feeding habits in relation to the increased abundance of wild ungulates. From June 2007 to May 2008 we collected wolf scats and recorded the signs of presence of wild ungulate species along 25 transects, representative of the different habitat types within the study area. We analysed the scats to identify the main food items used by wolves in each season and we compared the proportions of biomass of wild ungulate species in the diet (use) to those calculated from the signs of presence (availability). We recorded significant seasonal changes in wolf feeding habits, but the main food items were wild ungulates, mainly wild boars, and medium-sized mammals. Livestock (sheep, goats, and calves) were used only in summer and early autumn, during the grazing period, in small quantities. Comparing the current diet of wolves to that recorded in the past in the same study area highlighted significant differences for wild ungulates and livestock, the former being more used nowadays and the latter less. Our study showed that, in the Northern Apennines, the feeding behaviour of wolves has evolved towards a greater use of wild prey, so reducing the impact on livestock and consequently wolf-human conflicts. Moreover, predation by wolves was directed towards the most abundant species of wild ungulates.
The expansion of the wolf population all over Europe has posed problems on wolf-man coexistence in those areas where the wolf was not present for a long time. Breeding activities are now threatened by high predation on livestock. We investigated the relationship between wolf and its habitat, in order to evaluate wolf habitat suitability and to predict its presence. The study areas covered 3289 km2. A 23 km2 grid was used to identify 143 sample squares. For each sample square 58 habitat variables were measured from land-use (1:25 000), and from topographical maps. Wolf presence (4 classes) was assessed by scat collection, direct observations, snow tracking, wolf-howling, and predation records. Univariate and multivariate statistical analyses were carried out to investigate the influence of habitat features on wolf presence. The habitat suitability model was based on two different equations: a dichotomous logistic regression model, and a polytomous one. The first one discriminated between suitable and unsuitable habitats, and its predictions were confirmed in 93% of grouped cases; the second provided predicted values of wolf presence that were concordant with the observed response levels in 82.5% of contrasts. Both models underlined the importance of three factors in determining wolf habitat suitability: wild prey abundance, human presence and forest cover.
We describe extensions to the method of Pritchard et al. for inferring population structure from multilocus genotype data. Most importantly, we develop methods that allow for linkage between loci. The new model accounts for the correlations between linked loci that arise in admixed populations (“admixture linkage disequilibium”). This modification has several advantages, allowing (1) detection of admixture events farther back into the past, (2) inference of the population of origin of chromosomal regions, and (3) more accurate estimates of statistical uncertainty when linked loci are used. It is also of potential use for admixture mapping. In addition, we describe a new prior model for the allele frequencies within each population, which allows identification of subtle population subdivisions that were not detectable using the existing method. We present results applying the new methods to study admixture in African-Americans, recombination in Helicobacter pylori, and drift in populations of Drosophila melanogaster. The methods are implemented in a program, structure, version 2.0, which is available at http://pritch.bsd.uchicago.edu.
THE FIRST REAL BEGINNING to our understanding of wolf social ecology came from wolf 2204 on 23 May 1972. State depredation control trapper Lawrence Waino, of Duluth, Minnesota, had caught this female wolf 112 km ( 67 mi) south of where L. D. Mech had radio-collared her in the Superior National Forest 2 years earlier. A young lone wolf, nomadic over 100 km2 (40 mi2) during the 9 months Mech had been able to keep track of her, she had then disappeared until Waino caught her. From her nipples it was apparent that she had just been nursing pups. "This was the puzzle piece I needed," stated Mech. "I had already radio-tracked lone wolves long distances, and I had observed pack members splitting off and dispersing. My hunch was that the next step was for loners to find a new area and a mate, settle down, produce pups, and start their own pack. Wolf 2204 had done just that."
Closed form representations of the gradients and an approximation to the Hessian are given for an asymptotic approximation to the log likelihood function of a multidimensional autoregressive moving average Gaussian process. Their use for the numerical maximization of the likelihood function is discussed. It is shown that the procedure described by Hannan for the estimation of the parameters of one dimensional autoregressive moving average processes is equivalent to a three stage realization of one step of the Newton Raphson procedure for the numerical maximization of the likelihood function, using the gradient and the approximate Hessian. This makes it straightforward to extend the procedure to the multidimensional case. The use of the block Toeplitz type characteristic of the approximate Hessian is pointed out.
Within the 2700km² Beltrami Island State Forest, near the W edge of the primary range of Canis lupus in Minnesota, wolf population density was low at the start of the study in 1972 but increased substantially up to 1977 (end of study). At least 8 of 13 social units present in mid-1976 had formed since 1972. Size of litters of established packs averaged 4.6 pups, and those of newly-formed pairs averaged 4.1. Mortality decreased over the study period, and recruitment of young wolves exceeded mortality following legal protection. A high rate of dispersal of young from packs was documented. Dispersal peaked in autumn. Most wolves paired within a few days of leaving their packs. Average territory size decreased as both population and pack numbers increased. Behaviour of alpha males, alpha females and subordinate members of the packs is discussed. Deer and moose comprised 94% of animal biomass eaten by wolves, with deer along accounting for 67%. Seasonal differences in food taken and energy requirements are noted.-P.J.Jarvis
In an area of the Abruzzo National Park, Italy, wolf preyed mainly upon wild and domestic ungulates. Occurrence of red deer Cervus elaphus in the diet exceeded that of the roe deer Capreolus capreolus, parallel to the relative local abundance of these species. Although consumption of reintroduced deer species increased during the study period, no corresponding decrease of predation on livestock could be assessed. Diet of the wolf did not show seasonal variations in the study area, whereas that of the fox did. Use of different trophic resources by these two carnivores was likely to generate such a difference, the former taking prey steadily available throughout the year, while the latter makes use of seasonally fluctuating food resources (eg rodents and fruits). The dependence on anthropogenic food resources was noticeable for the wolf, but mainly restricted to domestic ungulates, and was negligible for the fox. Food competition is unlikely in well preserved habitats, where they can fill different trophic niches. -from Authors
In the early 1990s the wolf (Canis lupus) settled in the Mercantour's mountains, southeastern France. Its predation on domestic and game species prompted a strong opposition from part of the local residents. In this context, it appeared of prime importance to collect data on the occurrence of ungulates in the wolf's diet. The diet was determined through the analysis of 236 scats collected from April 1994 to March 1995 in a 280 km2 mountainous study area where a pack of 8 wolves was settled. The chamois (Rupicapra rupicapra) is abundant (about 1,000/100 km2) in this area and a population of 300 moufflons (Ovis gmelini) was present. There are also small populations of roe deer (Capreolus capreolus), red deer (Cervus elaphus), ibex (Capra ibex) and wild boar (Sus scrofa). Concerning livestock, 10,000 sheep (Ovis aries), some cows (Bos taurus), goats (Capra hircus) and horses (Equus caballus) graze from June to October in the study area. Ungulates (wild and domestic pooled) made up the bulk of the wolf's diet, occurring in 97% of the scats analysed. Their relative occurrence varied from 94% in winter and spring to 87% in summer and autumn. Wild ungulates remains, mainly those of moufflons and chamois, occurred in 80% of the scats analysed. In summer, the chamois occurrence relative to wild ungulates was more than twice that of the moufflon (71% vs 29%). Conversely, the relative occurrence of the moufflon was almost twice that of the chamois in spring (68% vs 32%), autumn (64% vs 33%) and winter (59% vs 26%) despite the fact that moufflons were ten times less abundant than chamois. The seasonal selectivity of moufflons by wolves was probably related to the fact that the moufflon i) cannot escape easily when the snow depth is important, ii) occurs in herds that are predictably associated with good habitat patches during winter, iii) tends to stay in lower altitude than the chamois from autumn to spring and iv) is the first ungulate species to breed in the Mercantour. Concerning domestic ungulates, free ranging sheep were the main target of wolf predation on livestock because of their local abundance and accessibility. The sheep's occurrence relative to the total ungulate's one was 50% in summer (main grazing period) and 22% in autumn. Wild ungulates were preferred to livestock. However, our data show that, when prevention methods are lacking, the predation on domestic ungulates is not necessarily rare in areas characterized by a rich and diverse wild ungulate guild.
A decline in the gray wolf (Canis lupus) population in Isle Royale National Park prompted an intensive study of radiocollared individuals in 1988-1991, complementing an ongoing study begun in 1958. During winter, 1959-1991, the proportion of lone wolves was higher when the population was at low levels, whereas average size of packs declined with density of wolves. Sizes of territories (1971-1991) were not related to size of packs. In winter, food availability (kg wolf-1 day-1) and kill interval (days/kill) varied inversely with size of packs. Social relationships of radiocollared lone wolves were flexible; they often formed temporary groups of two or three wolves. Small groups (less than four) and lone wolves roamed large areas of the island (300 km2 and 540 km2, respectively), frequently moving through territories of established packs. Solitary wolves and pairs readily killed adult moose, in contrast to a common belief that larger packs benefit from cooperative hunting.
We give suggestions for the presentation of research results from frequentist, information-theoretic, and Bayesian analysis paradigms, followed by several general suggestions. The information-theoretic and Bayesian methods offer alternative approaches to data analysis and inference compared to traditionally used methods. Guidance is lacking on the presentation of results under these alternative procedures and on nontesting aspects of classical frequentist methods of statistical analysis. Null hypothesis testing has come under intense criticism. We recommend less reporting of the results of statistical tests of null hypotheses in cases where the null is surely false anyway, or where the null hypothesis is of little interest to science or management.
Knowledge about recruitment in a population can be critical when making con-servation decisions, particularly for harvested species. Harvest can affect popula-tion demography in complex ways and this may be particularly true for cooperatively breeding species whose successful reproduction is often linked with complex social dynamics. We currently have a poor understanding of how harvest affects recruitment in cooperatively breeding species. We used non-invasive genetic sampling and a natural experiment to estimate recruitment in a population of gray wolves Canis lupus before and after harvest in the northern Rocky Moun-tains, US (2008–2013). We hypothesized that recruitment would decline after hunting and trapping began and that the decline in recruitment would be attrib-utable to the harvest of pups and not to the subtler mechanisms associated with group dynamics and reduced reproductive success. We collected fecal samples from wolves in 10 packs for 6 consecutive years, extracted DNA and genotyped 154 individual pups across 18 microsatellite loci. Population harvest rates aver-aged 23.8% (sd = 9.2). Our hypothesis that recruitment would decline was sup-ported; survival from 3 to 15 months of age decreased from 0.60 [95% confidence interval (CI): 0.48–0.72] without harvest to 0.38 (95% CI: 0.28–0.48) with harvest and recruitment declined from 3.2 (95% CI: 2.1–4.3) to 1.6 (95% CI: 1.1–2.1) pups per pack after harvest was initiated. We cannot unequivocally dismiss other factors that could have reduced recruitment, however, an increase in recruitment when harvest temporarily ceased lends support to our conclusion that harvest reduced recruitment. We attributed just 18–38% of pup mortality directly to harvest and suggest that there are indirect effects of harvest on recruitment that may be associated with changes in group size and structure. Models that do not include both direct and indirect effects of harvest on recruitment may underesti-mate the potential impact of harvest on population growth in social species.
(1) The food of European badgers Meles meles L. is described from six areas in Scotland, using faecal analysis. The estimated relative volume of various foods in the diet is compared with the frequency of their occurrence. Variation between areas and differences between seasons are related to food availability, and the food diversity and variance of different foods with time are presented. (2) The dominant food everywhere was earthworms, Lumbricus terrestris L. and L. rubellus Hoffmeister but their importance in the diet varied little in time and between areas, and there was no correlation with availability. Other less important foods included rabbits, cereals, insects and tubers, and several minor food categories; they were consumed relative to availability in those cases where this availability could be measured. (3) It is suggested that badgers change their foraging effort to compensate for fluctuations in earthworm availability, consuming a range of secondary foods opportunistically.
IN SOME APPLICATIONS of the general linear model, the usual assumptions of homoscedastic disturbances and fixed coefficients may be questioned. When these requirements are not met, the loss in efficiency in using ordinary least squares (OLS) may be substantial and, more importantly, the biases in estimated standard errors may lead to invalid inferences. This has caused a number of writers to propose models which relax these conditions and to devise estimators for their more general specifications, e.g., Goldfeld and Quandt (8) for heteroscedasticity and Hildreth and Houck (11) for random coefficients. However, because the effect of introducing random coefficient variation is to give the dependent variable a different variance at each observation, models with this feature can be considered as particular heteroscedastic formulations for the purpose of detecting departure from the standard linear model. A test for heteroscedasticity with the same asymptotic properties as the likelihood ratio test in standard situations, but which can be computed by two least squares regressions, thereby avoiding the iterative calculations necessary to obtain maximum likelihood estimates of the parameters in the full model, is considered in this paper. The approach is based on the Lagrangian multiplier
The 400–500 wolves currently living in the Apennine range of peninsular Italy are slowly recolonizing the Alps and are expected to move northward. A nationwide management plan for the Italian wolf population is being prepared, and a zoning system with connecting corridors has been suggested. We developed a large-scale probabilistic model of wolf distribution as a contribution to the planning process. Thirteen environmental variables related to wolf needs and human presence were analyzed in 12 well-studied wolf territories and in 100 areas where the species has been absent for the past 25 years. These two areas were used as a training set in a discriminant analysis to evaluate potential wolf presence throughout the entire country. We used the Mahalanobis distance statistic as an index of environmental quality, calculated as the distance from the average environmental conditions of the wolf territories. Based on the Mahalanobis distance statistics, we constructed an actual and potential spatial distribution of the wolf for all of peninsular Italy. The jackknife procedure was used to assess the stability of the distance model and showed good confidence in our model (coefficient of variation ≤ 13%). Distance from the wolf territories’ centroid as an index of environmental quality for the wolf was tested using 287 locations where wolves have been found dead in the past 25 years as a consequence of human action (poison, shotgun, car accidents). A useful contribution to conservation planning resulted from comparing the frequency distribution of the Mahalanobis distance of the dead wolf locations with the percentage of study area within each distance class. This showed how the number of wolf casualties would greatly decrease with protection of only a minor part of the study area and indicated the usefulness of our approach for evaluation of other conservation options, such as core areas and corridor identification.
We consider testing for an exponential distribution with unspecified rate parameter when it is only possible to observe the counts in groups with boundaries specified before sighting the data. On the basis of a size and power study we recommend that tests of fit for the exponential distribution be based on the Anderson–Darling statistic and the SW2SW2 statistic recommended by Gulati and Neus [(2001). Goodness-of-fit statistics for the exponential distribution when the data are grouped. In: Huber-Carol, C., Balakrishnan, N., Nikulin, M.S., Mesbah, M., (Eds.), Goodness-of-Fit Tests and Validity of Models. Birkhauser, Boston, pp. 113–123 (Chapter 9)]. We also suggest that inference based on one of these be complemented by examination of the components of the X2X2 statistic. We illustrate how to use these components to give improved models.
Following several years of occasional occurrence, several wolvesCanis lupus Linnaeus, 1758 have established a resident population in northeastern Saxony (Eastern Germany). From 2001 to 2003, we collected
and analysed 192 scats ofC. lupus. Results of our study are expressed as the frequency of occurrence of prey species and the percentage of biomass consumed
using coefficients of digestibility as well as two variants of an equation for prey mass per collectable scat. Diet composition
of the wolves was restricted to a few food items, mostly wild ungulates. These remains were found in 97% of the scats, representing
99% of the biomass consumed by the wolves. Roe deerCapreolus capreolus was the most frequent and most important prey, constituting nearly of one half the biomass. Red deerCervus elaphus was recorded in one-third of the samples, followed by wild boarSus scrofa, mouflonOvis am mon musimon and brown hareLepus europaeus. Compared with game occurrence, roe deer was clearly preferred over the other species. A difference between winter and summer
diets was mainly due to the high occurrence of young wild boar in summer. The general diet pattern of the wolf in Saxony corresponds
with that found in the naturally occurring populations in Europe.
In West Galicia, the diet is centered on scraps from industrial poultry and pig farms, and domestic ungulates. In the Cantabrian and Demanda Mountains, the diet is more diversified, ranging from an almost exclusive dependence on wild ungulates with domestic ungulates as a complementary prey, to the opposite situation. In the Douro Meseta lagomorphs were added to the conventional prey groups, according to season. In Extremadura, wolves prey on both wild and domestic ungulates, while in Sierra Morena they prey almost exclusively on big game. Young wolves consumed more chickens, arthropods and fruit, while adults consumed more wild ungulates. -from Authors
The brown bear Ursus arctos, wolf Canis lupus, and Eurasian lynx Lynx lynx vanished during the 18th and 19th centuries from all regions of high human activity in Europe because of direct persecution and environmental changes. Bear, wolf, and lynx were vulnerable in different ways to deforestation and the destruction of wild ungulate populations. Analysing the ecological factors responsible for the fall of the large carnivores can help to prepare their recovery. The return of large predators into semi-natural areas such as the Alps is possible, as the forests have expanded, and the wild ungulate populations increased. Lynx reintroduction in the Alps started in the 1970s. Wolves returned to the south-western Alps from the central Italian population in the early 1990s. The brown bear is recolonising the Austrian Alps from Slovenia. However, the modern protective legislation is not backed by a cooperative attitude among the affected people. In rural areas, large carnivores are still regarded as unrestrained killers of wildlife and livestock. Ecological conditions and husbandry in the Alps have been altered substantially since the large carnivores were eradicated, and the potential for conflicts has diminished. But stockmen have lost any remaining tradition of coexistence with large predators, and sheep are again very abundant in the Swiss Alps. The return of the large predators will not be possible without changing the system of sheep-husbandry. The rural people are not yet willing to do so. They generally object to any change in their lifestyle induced from outside, and the large predators become a negative symbol for restrictive conservation measures considered to hinder economic development. Nature conservation, including the reintegration of large predators, must be integrated into rural development; local people must be much more involved in this process.
I examined the components generating selectivity in wolves preying on five ungulate species in Banff National Park, Alberta. Overall selectivity for elk and deer species, and apparent avoidance of bighorn sheep and mountain goats, were due primarily to lower habitat overlap of wolves with the latter two species, and therefore lower encounter rates. For social ungulates, I argue that the herd should be considered the unit of encounter, with encounter rates proportional to the number of herds rather than the number of individuals. However, large herds predictably associated with certain areas may be visited intentionally by wolves, increasing effective encounter rates. Foraging theory suggests that all ungulate prey should be equally profitable to wolves upon encounter and therefore the factors affecting encounter rates are critical in determining prey selectivity. A simple model incorporating different habitat overlap, herd sizes, and predictable herds predicts qualitatively different functional responses of wolves to changes in density of the different prey types. The model also demonstrates how apparent selectivity for a prey type can result from the different ways in which prey are encountered.