RECENT RESEARCH | November – December 2020

The grey wolf (Canis lupus) is a widespread but locally endangered species across Pakistan. The current study investigates the occupancy of grey wolf and conflicts with humans in Khanbari valley in Gilgit Baltistan. The study was conducted from the 5th of December 2014 to the 11th of January 2015. The study area was delineated into blocks following natural watersheds, and overall 47 motion-triggered cameras were installed in various locations encompassing an area of 810 km2 with an effort of 1428 trap nights. A human-wolf conflict survey was carried out through questionnaires, where 57 respondents were randomly chosen from 8 villages in the valley. Grey wolf was photo-captured at 11 different camera stations, occupancy estimated at 0.37±0.22 S.E., and detection probability of 0.29±0.19 S.E. A total of 166 livestock were killed which incurred an economic loss of USD 17,046 (USD 299 per household) in five years. Predation on goat was highest, though consumed as per availability. Sheep predation indicates selection for this animal because predation was much higher than availability. Cattle was predated as per availability and accounts for the least part of the livestock loss. Predation of livestock was greatly influenced by four factors: habitat, prey type, prey age, and time of predation. We recommend conservation initiatives like compensation for economic losses, predator-proof corrals, and awareness campaigns to promote human-wolf co-existence in the area.

Knowledge on interference competition between species, particularly for scarce crucial resources, such as water, is a topic of increasing relevance for wildlife management given climate change scenarios. This study focuses on two sympatric canids, the African wolf and the side-striped jackal, to evaluate their group size and spatiotemporal activity patterns in the use of a limited resource by monitoring artificial waterholes in a semi-arid environment located in Senegal (West Africa). Remote cameras were deployed at five artificial waterholes to evaluate the number of individuals, age and activity patterns of resource use. African wolves (n = 71; 31% of all carnivore detections) and side-striped jackals (n = 104; 45%) were the most detected carnivore species. While both canids tended to occur alone at waterholes, they showed an evident monthly variation in group size. Both species showed a high activity overlap, with a bimodal activity pattern in waterhole use. However, we found evidence of unidirectional spatiotemporal avoidance, suggesting African wolves might be dominant over side-striped jackals. Our findings provide useful insights to investigate niche partitioning on the use of limited resources and have conservation implications for regions with a prolonged dry season.

Livestock guardian dogs (LGDs) in the Romanian Carpathians are as old as the pastoral presence and activity in the region. The main role of these dogs is to protect livestock from predation by large carnivores. The Carpathian Mountains, as opposed to other European mountain ranges, have always had considerable populations of wolf, brown bear, and lynx; conflict with the herders is inevitable. Here, the shepherds rely only on themselves and their dogs to keep their animals safe from predation during pastoral movements. We investigated 12 sites from the historical regions of Banat and Transylvania, where we have collected traditional ecological knowledge (TEK) on the use of native LGDs as an ancient non-lethal method for the prevention of livestock depredation. By monitoring the behavior of their dogs, the shepherds establish a complex ethno-ethological relationship with them, which helps them foretell the movements and presence of large carnivores in their vicinity. We have also investigated the recent positive change of attitude of some of the Romanian nature conservationists towards the Romanian Carpathian Shepherd Dog breed, which is also currently promoted by important international nature conservation NGOs as an ecologically friendly method to mitigate the conflict with large carnivores. The uninterrupted use of endemic LGD breeds by pastoralists in Romania might be one of the main reasons for the survival and conservation of large carnivores here in the past and in the future.

Coyotes (Canis latrans) have adapted and learned to live alongside humans but not without cost. Seen as a pest, nuisance or biosecurity threat, coyotes often receive indirect or direct violence from the communities they live alongside with.It is our position that the values and behaviours justifying violence towards coyotes energize the same systems of oppression responsible for the marginalization of other groups (such as Indigenous peoples, non-male genders, differently-abled, people of colour, etc.). Herein, we liken the systemic violence that coyotes experience to that enacted against marginalized human groups. We argue that speciesism is foundational to the treatment of coyotes and that this structural oppression must be interrogated as we would racism, classism and sexism.Next, we suggest that both human and wildlife welfare need to be considered jointly in future conservation efforts, as the oppression of marginalized human and nonhuman animal groups are often linked. We conclude that structural changes in academia, wildlife management policies,and grassroots education are essential to dismantling traditions of violence towards coyotes.

In 1966 the gray wolf (Canis lupus) was regarded as functionally extinct in Norway and Sweden (the Scandinavian peninsula). In 1978 the first confirmed reproduction on the peninsula in 14 years was recorded. During 20 successive winters, from 1978–1979 to 1997–1998, the status, distribution, and dynamics of the wolf population were monitored by snow-tracking as a cooperative Swedish–Norwegian project. After the 1978 reproduction in northern Sweden, all new pairs and packs were located in south-central parts of the Scandinavian peninsula. Between 1983 and 1990 wolves reproduced each year except 1986, but in only one territory. There was no population growth during this period and the population never exceeded 10 animals. In 1991 reproduction was recorded in two territories. After that there were multiple reproductions each year and the population started growing. In 1998 there were 50–72 wolves and six reproducing packs on the peninsula. Between 1991 and 1998 the annual growth rate was 1.29 0.035 (mean SD). A minimum of 25 litters were born during the study period. The early-winter size of packs reproducing for the first time was 6.2 1.4 wolves (n = 9), and this decreased with time during the study. The size of packs that had reproduced more than once was 6.4 1.8 wolves (n = 12), and this increased with time over the study period. All but 1 of 30 reported wolf deaths were human-caused. The annual mortality rate was 0.13 0.11, and this decreased with time during the study period. The minimum dispersal distance was 323 212 km for males and 123 67 km for females. Of 10 new wolf territories where breeding occurred, only 1 bordered other, existing territories. The distance from newly established wolf pairs to the nearest existing packs was 119 73 km. Simulation of population growth based on known reproductions and mortalities showed a close similarity to the results from population censuses up to the mid-1990s. To what extent this population is genetically isolated is at present unclear.

Damage to livestock is the main cause of conflict between human activities and the grey wolf (Canis lu-pus) throughout most of the species’ range. In the past, people responded by persecuting wolves, eradicating them from many areas (Boitani, 2003). Later, wolves received legal protection in many countries and damage compensation schemes were implemented as part of a strategy to alleviate conflicts.In recent decades, the wolf has been naturally re-covering in many regions of Europe (Chapron et al., 2014), returning to areas with high densities of live-stock but where traditional methods to protect them from predators are no longer used (Linnell and Cre-tois, 2018). Various methods to protect livestock from wolves and other large carnivores have been tested around the world (Linnell et al., 1996; Shivik, 2006). These differ in terms of effort and cost to install and maintain, user-friendliness, longevity, flexibility and, of course, effectiveness (Gehring et al., 2010). Not all techniques are suitable in every situation: methods should be chosen and adapted to the predation risk and specific conditions in each holding (Linnell and Cretois, 2018). Among the most widely used and rec-ommended measures to prevent damage and hence promote coexistence are livestock guarding dogs (LGDs) and electric fences (e. g. Boitani, 2000; Breit-enmoser et al., 2005; Rigg, 2001; Wade, 1982).In Europe, many projects and initiatives have aimed to reduce damage caused by large carnivores, some of them funded by the EU LIFE Programme1 (Sal-vatori, 2013). One such project, LIFE Coex (LIFE04 NAT/IT/00144), was implemented from 2004 to 2008 in Portugal, Spain, France, Italy and Croatia. One of its main activities was to implement and promote damage prevention measures for livestock, beehives and crops. During the project, 290 electric fences, 22 conventional wire-netting fences and 245 LGDs were implemented, monitored and assessed (see: LIFE Coex, 2008; Salvatori and Mertens, 2012). In Spain and Portugal, measures were focused on reducing losses of livestock to wolves at a total of 144 holdings..
… In this article, we present results from an assessment of the use of three types of damage prevention measures in Portugal and Spain a decade after they were implemented during the LIFE Coex project in 2004 – 2008. Specifically, we wanted to know the level of satisfaction of the beneficiaries, their perceptions of the efficacy and maintenance costs of the measures or, if applicable, their main reasons for no longer using them, as well as their suggestions to encourage other farmers to implement them. Whenever appropriate, a comparison was made with assessments made at the end of the project.

We modeled resource selection by wolves (Canisspp.), eastern coyotes (C. latrans), and admixed canids during the pup-rearing season at den and rendezvous sites (collectively, homesites) within a largely unprotected landscape proposed as the recovery zone for federally and provincially threatened eastern wolves (C. lycaon) in Ontario, Canada. Overall, canids selected wetlands, while avoiding secondary roads and open-structure rock-grass habitat patches. Packs with greater wolf ancestry selected wetlands and tertiary roads more strongly, while avoiding mixed conifer-hardwood forests. Contrary to our prediction, canids with greater coyote ancestry did not establish homesites closer to roads, which likely mitigated their risk of human-caused mortality during pup-rearing. Packs exhibited increased selection of wetlands within territories as a function of increasing availability of wetlands. Packs with abundant access to wet-lands may prioritize this habitat type to exploit beavers, a valuable prey species during pup-rearing. Packs with higher pup survival selected hardwood forests and avoided conifer forests more than packs with lower pup survival. This is consistent with our understanding of habitat relations of the main prey species for canids in central Ontario and suggests that selecting prey-rich habitat types at homesites increases fitness. A proposed goal of eastern wolf recovery is numerical and geographical expansion outside of the population core in Algonquin Provincial Park. Thus, our results provide valuable information for conserva-tion by quantifying resource selection of wolves, coyotes, and hybrids during pup-rearing and identifying links between fitness and homesite selection.

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat‐ or disturbance‐mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous‐browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193‐km2 section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire‐return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km2), likely because of the scarcity of deciduous or mixed‐wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km2). Wolf density (3.1 wolves/1,000 km2) and pack sizes (urn:x-wiley:0022541X:media:jwmg21982:jwmg21982-math-0001 = 4.0 wolves/pack) were low and resident (established) territories were large (urn:x-wiley:0022541X:media:jwmg21982:jwmg21982-math-0002 = 4,360 km2; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km2). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous‐browse and alternate‐prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.

In Finland, all four large carnivore species, brown bear, grey wolf, Eurasian lynx and wolverine, have established populations, though their presence is not easily accepted by some. Large carnivores pose a threat to livestock and cause fear in the locals living in their territories. Wolf – hunting dog conflict is especially prominent in Finland south of reindeer husbandry area and the poaching of wolves hinders the population’s management.

Attitudes towards large carnivores are often influenced by personal background, such as education level, ecological knowledge and respondent’s position in possible human – wildlife conflict. Residence can have an effect, as well, since conditions between living in urban or rural areas often differ. Hypothesis for this study are 1) attitudes towards large carnivores get worse while getting closer to protected areas, 2) attitudes towards large carnivores differ between eastern and western study areas, and 3) a higher education level increases positive attitudes towards large carnivores. The effect of ecological knowledge, prior experiences with large carnivores, age, sex and position in conflict was also explored.

Study was conducted as a questionnaire, with face-to-face interviews and web survey distribution targeting two areas in Finland with large carnivore occupancy, one in the West and one in the East. A link between negatively perceived personal experiences and negative opinions towards large carnivores and their management was found. Living in the western area, where large carnivores have resided for a shorter time, predicted attitudes towards stricter management of the species. Third level education influenced attitudes positively.

By understanding local attitudes towards large carnivores, it is possible to better understand the conflict between humans and predators, and so, find more likely solutions. Conservation actions where locals have been included, have been documented as successes. Regional differences in attitudes should be further studied and included in future decision making.

Even though the Persian leopard Panthera pardus saxicolor is an endangered subspecies with the main population inhabiting in Iran (Khorozyan and Abramov, Zool Middle East 41:11–24, 2007; Kiabi et al., Zool Middle East 26(1):41–47, 2002), earlier studies (Sanei et al., Assessment of the Persian leopard mortality rate in Iran. In: Proceedings from UMT 11th International Annual Symposium on Sustainability Science and Management (pp. 1458–1462, 2012). Terengganu, Malaysia: Universiti Malaysia Terengganu) demonstrated that the majority of leopard mortalities are recorded to be as a result of intentional hunting, revenge killing, and poisoning of the specimens. To mitigate livestock–carnivore conflicts and reduce the subsequent revenge killings, an innovative model including a medium and a long-term insurance schemes together with awareness raising, trust building, and participatory conservation strategies is designed. Accordingly, the medium term insurance scheme addresses three main subjects of (1) improving conservation practices in the areas of leopard mortality hot spots, (2) medical payments and wergild for possible human injuries/maim/death because of human–leopard conflicts and (3) recompensing livestock depredation. Also, since the wolf Canis lupus distribution is comparable with the leopard range in the country, because of conservation concerns, damages caused by wolf depredation are also planned to be recompensed partially in the first type (i.e. medium term) insurance scheme and fully recompensed in the long term (i.e. second type) insurance program. Introducing sessions about the relative regulations and instructions were conducted for provincial wildlife wardens and DoE staff who are well familiar with wildlife sign surveys and have a quick access to the habitats in each region. Subsequently, they took the responsibility for identification of wildlife species in livestock–carnivore conflicts. Improvements in husbandry practices, linking the payments to the acceptable husbandry enhancements and participation in reducing risk of damages by local people are some of the instructions considered in the model to improve the efficacy and outcomes. So far, Department of Environment of Iran together with a private insurance company has partially launched the short term insurance scheme since 2016 and launching other sections of this model is in progress.

nimal social strategies are of importance when avoiding predation. Since horses are the least hunted among all farm animal species, we suppose that the alert reaction to a predator’s vocalisation, followed by anti-predator social behaviour, still exists in domestic horses. Recorded vocalisations of three different predators (grey wolf, Arabian leopard and golden jackal) were played to 20 horses of two horse breeds—namely, Konik polski and Arabian. Social responses and tactics in antipredator behaviour differed between the breeds and between predators. Koniks exposed to vocalisations of a howling wolf resulted in tight groupings, while Arabians exposed to the growling of a leopard responded with linear group formation. The behaviour of studied horses, expressed by alertness and defensive formations, indicates existence of the social anti-predator behavior, which in turn may explain the low rates of horses falling prey to predators as compared with other farm animal species.
Abstract
We tested the hypothesis that social defensive responses to the vocalisation of a predator still exist in horses. The recordings of a grey wolf, an Arabian leopard and a golden jackal were played to 20 Konik polski and Arabian mares. Durations of grazing, standing still, standing alert and the number of steps in walk and trot/canter were measured. In one-minute scans, the distances of the focal horse from the reference horse (DIST-RH) and from the nearest loudspeaker (DIST-LS) were approximated. The vocalisation of a leopard aroused the Arabians more than the Koniks (less grazing, stand-still and walk, more stand-alert and trotting/cantering). Koniks showed more relaxed behaviours to the leopard vocalisation (more grazing, stand-still and walk), but high alertness to the wolf playback (stand-alert, trotting/cantering). Spatial formation of the herd of Koniks showed tight grouping (lower DIST-RH) and maintaining distance from the potential threat (DIST-LS) in response to the wolf howling, while the Arabians approached the loudspeakers in linear herd formation when the leopard growls were played. Adult horses responded to potential predation by changing spatial group formations. This ability to apply a social strategy may be one of the explanations for the least number of horses among all hunted farm animal species.

Although there are recent claims of a lack of evidence of self-consciousness in many tested species, the ability to recognize oneself in a mirror, which seems an exceedingly rare capacity in the animal kingdom, may not be the only way to check for animal self-awareness (i.e. the capacity to become the object of your own attention). A new testing approach, based on a different sensory modality (such as the sniff-test for self-recognition, STSR), recently proved to be effective with dogs. We applied this sniff test to a group of four captive grey wolves, living in male-female couples in two different enclosures at the Wolf Park in Indiana, USA. In this preliminary study, wolves showed some signs of the ability to recognize themselves through the “olfactory mirror” and exhibited some clues of mark-directed responses, particularly scent-rolling, which may shed more light on this still unclear behavior and represent a sort of olfactory equivalent to passing the original mirror test.

The occurrence of carnivore species in wolf diet has been overlooked and poorly studied despite the potential implications for wolf ecology and wildlife management. We conducted an extensive literature review, focusing on 120 wolf diet studies worldwide to assess global patterns of carnivore consumption by wolves and their ecological and human-related determinants. We used a total of 143 sampling sites with data on the consumption of carnivores by wolves. In total, 35 carnivore species were reported to be consumed by wolves, comprising members of all taxonomic carnivore families represented within the gray wolf range. The carnivores were mostly limited to occasional consumption (<5% of wolf diet) but could account for as much as 25% in some study areas. The most frequently consumed carnivore species were those with reported scavenging behavior, belonging to medium-sized generalist canids. Generalized linear model (GLM) analysis revealed that higher magnitudes of carnivore consumption were related to nonprotected areas as well as lower occurrences of wild ungulates, domestic ungulates, and small mammals in wolf diet, while higher numbers of consumed carnivore species were related to nonprotected areas with low vegetation productivity and lower occurrences of domestic ungulates and small mammals in wolf diet. Our results suggest that carnivore consumption by wolves is driven by altered ecosystems and human-dominated landscapes, where mesopredator densities are often increased and prey densities decreased, which intensify competition and the need for alternative food sources.

Generally, Plitvice Lakes National Park’s values have so far been preserved, however a number of concerns exist, especially in relation to its water ecosystem. All lakes forming the central part of the park, which include 16 large lakes and some smaller ones, are subject to continuous eutrophication – a natural process whereby a water body gets enriched with nutrients, often excessively. However, due to potential higher levels of organic pollution, the eutrophication process might become considerably faster. Human activities are influencing this acceleration, including water use and waste water discharge from tourism infrastructure and households, agricultural practices with the use of chemicals in the upper watershed of the site and outside its boundary, and livestock farming. Such threats require measures to maintain the water flow and reduce sources of pollution, namely stopping urbanisation and building an efficient waste water management system. However, no concrete measures have so far been developed and pressures have only been increasing. Recently, concerns have been raised over the rapid expansion of tourism facilities within the property, including with regards to spatial planning regulations and their implementation. In addition to posing a threat to the site’s sensitive hydrogeological system, growing tourism infrastructure and uncontrolled visitor numbers are affecting the visual integrity of the site. The recent UNESCO/IUCN reactive monitoring mission (2017) concluded that, while the site’s ecological integrity has so far been preserved, the current and potential serious threats to the Outstanding Universal Value (OUV) arising from these developments – together with related threats of excessive visitor numbers, water usage, water pollution, road infrastructure and traffic – are of significant concern. While the relevant authorities show commitment to address the issue, urgent measures are required in order to reduce pressures and prevent any irreversible damage to its values.

An emerging conflict with Trans-Himalayan pastoral communities in Ladakh’s Changthang Plateau threatens the conservation prospects of the kiang (Equus kiang) in India. It is locally believed that Changthang’s rangelands are overstocked with kiang, resulting in forage competition with livestock. Here, we provide a review and preliminary data on the causes of this conflict. Erosion of people’s tolerance of the kiang can be attributed to factors such as the loss of traditional pastures during an Indo-Chinese war fought in 1962, immigration of refugees from Tibet, doubling of the livestock population in about 20 years, and increasing commercialization of cashmere (pashmina) production. The perception of kiang overstocking appears misplaced, because our range-wide density estimate of 0.24 kiang km−2 (± 0.44, 95% CL) is comparable to kiang densities reported from Tibet. A catastrophic decline during the war and subsequent recovery of the kiang population apparently led to the overstocking perception in Ladakh. In the Hanle Valley, an important area for the kiang, its density was higher (0.56 km−2) although even here, we estimated the total forage consumed by kiang to be only 3–4% compared to 96–97% consumed by the large livestock population (78 km−2). Our analysis nevertheless suggests that at a localized scale, some herders do face serious forage competition from kiang in key areas such as moist sedge meadows, and thus management strategies also need to be devised at this scale. In-depth socioeconomic surveys are needed to understand the full extent of the conflicts, and herder-centered participatory resolution needs to be facilitated to ensure that a sustainable solution for livelihoods and kiang conservation is achieved.

One of the seven equid species in the world, the kiang Equus kiang, occurs in parts of China and India, with small populations also reported from Pakistan and Nepal. Unlike the Asian wild ass E. hemionus, of which the population has declined drastically over the last century, kiang continues to have a wide distribution with fairly large populations (Schaller 1998). Within India, Ladakh (approximately 75° 50′ to 75° 80′ E; 32° 30′ N to 32° 37′ N) remains a stronghold for the kiang (Fox and others 1991; Shah 1996), where local Buddhist communities have been fairly tolerant of the species, and large herds of kiang can be relatively easily seen in eastern Ladakh. Kiang is classed as a Least Concern species under the IUCN Red List category; however the Western Kiang, E. k. kiang, which occurs in Ladakh, is classed as “data deficient” (Shah 2002).

Pacific pink salmon (Oncorhynchus gorbuscha) invasions, thought to originate from populations introduced and established in Russia, occurred along the Norwegian coast in 2017 and 2019. Despite several thousand pink salmon entering and establishing in northern Norwegian rivers, current understanding of the ecological effect of the species in northern Europe is limited. Scavengers feeding on pacific salmon carcasses are important vectors for the transport of marine derived energy and nutrients to terrestrial ecosystems in the Pacific Northwest, North America, where the salmon naturally occur. However the role of terrestrial and aquatic scavengers in the consumption and removal of pink salmon beyond the salmon’s native range is unknown. This study has identified terrestrial and sub-aquatic vertebrate scavengers on pink salmon carcasses in a sub-arctic river in northern Norway. Avian scavengers filmed by a camera placed near sites baited with pink salmon carcasses included the Eurasian magpie (Pica pica), hooded crow (Corvus cornix), common raven (Corvus corax), the European herring gull (Larus argentatus), redwing (Turdus iliacus) and goosander (Mergus merganser). However, the largest carcass weight was removed by red foxes (Vulpes vulpes). Carcasses entering Vesterelv River in 2019 were estimated to provide energy and nutrients to the river ecosystem an order of magnitude lower than in the Pacific Northwest. This study provides some of the first information in northern Europe on the mechanisms and quantification of energy and nutrient transfer from the ocean to riparian environments via introduced Pacific pink salmon. Results help to begin to determine the ecological effect of pink salmon and the development of appropriate management strategies.

The mechanisms of interactions among carnivore species range from facilitation (mainly through increased availability of prey carcasses) to competition. We assessed the potential for facilitative/competitive interactions between the two most widespread carnivores in the world, the wolf and the red fox, in a prey-rich area recently recolonized by the apex predator. One could expect that the superior competitor would ecologically suppress the inferior one, leading to avoidance of the former by the latter. In a Mediterranean coastal area (2017–2018), we assessed spatiotemporal and dietary interspecific overlap and investigated whether the recovery of wolves affected food habits of foxes. Spatiotemporal overlap was extensive (0.84–0.89). Wild ungulates were the staple of the wolf diet (~88–90%); foxes used mainly invertebrates and fruits (~78%), with ungulates being a substantial food category (13% of diet; 66% of occurrences among vertebrate prey). Interspecific dietary overlap was low (0.23), but extensive (0.89) for vertebrate prey. In comparison to a preceding wolf-free period, the volume and occurrence of large mammals in the diet of foxes showed a 2.8- to 3.5-fold increase. Apparently, foxes did not avoid wolves, which provided additional food to the foxes as prey leftovers. In a rich community, the presence of wolves may increase the food spectrum of foxes. Temporal variation of facilitation vs. competition should be assessed in relationship to spatiotemporal changes of predator–prey numbers.

In facing our greatest challenges, researchers have questioned where the ‘wild things’ will reside in the future, and large carnivores have been a primary focal area.
The built environment plays a critical role in the propagation of countless species including carnivores; however, contemporary conceptualizations of human–nature relations do not satisfactorily attend to where the built environment should be placed within existing human–nature relation frameworks or how it impacts our ability to find space for carnivores.
This paper fills this information gap by investigating the role of the built environment in social–ecological systems (SES), specifically wildlife and carnivore conservation.
The paper unfolds in four stages: The first reviews empirical efforts to capture the relationship between human–natural–wildlife systems and the built environment. Second, using insights from the built environment literature, I argue that moving away from a common pool resource focus, decoupling wildlife and natural systems, investigating all infrastructure types and their interactions across systems, and considering the notion of hybrid systems offer pathways forward. Third, an explanation of the built environment's linkages to human and carnivore systems is undertaken to illustrate how the built environment facilitates the material and symbolic interactions through a blending of properties from human, wildlife and natural systems. Lastly, the argument is made that attending to the role of the built environment in human–wildlife relations can stimulate new research that reveals unhelpful habitual behaviour, feedbacks and barriers, and may also help explain unintended or unexplained consequences impacting human–carnivore relations not fully considered under existing frameworks.

The rapid extinction of species over the past few decades has created a biodiversity crisis. Factors contributing to recent extirpations are linked to increased human population growth, habitat loss and fragmentation, and over-exploitation of wildlife. Only decisive, effective action to combat biodiversity loss can reverse these trends. The use of indicator species as surrogates for biodiversity provides a way to identify areas with high biodiversity so that conservation efforts can be accelerated and supported in those areas. Predators are considered important indicators of healthy, biodiverse ecosystems due to their high trophic level and their direct and indirect interaction with other species. Using camera trap data from 221 cameras set across five vegetation types and five land use zones in South Africa, we evaluated carnivores as potential surrogates for biodiversity. We used the leopard (Panthera pardus), and three meso-predators: caracal (Caracal caracal), honey badger (Mellivora capensis), and black-backed jackal (Canis mesomelas), as candidate indicator species. We used mammals captured at the camera traps as a measure of biodiversity referred to as mammalian species richness. The mammalian species richness was highest in the Orange River Nama Karoo vegetation type and in privately owned game reserves. We found that predator sightings were associated with significantly higher mammalian species richness which increased with increasing number of predator species. These findings suggest that the surrogate species concept can be applied to leopard and meso-predators.

Deer pressure on forest regeneration constitutes a serious problem in commercial forests in the northern hemisphere due to the increase in deer populations. However, other drivers, such as climate, landscape structure and the level of human activity, have a strong influence on deer pressure. The direct, density‐related impacts of ungulates on forest regeneration have been well studied, but there is limited empirical evidence related to the indirect factors mentioned above. We conducted a field experiment in three forest divisions in Poland to evaluate the role of a common element of human infrastructure, i.e. small, unpaved forestry roads. Additionally, we assessed the modifying effect of visibility driven by vegetation cover and forage availability.

RESULTS

The proximity of unpaved roads affected deer habitat use and foraging behaviour, and limited browsing pressure on regenerating forests. Low visibility and higher winter forage availability increased the probability of tree browsing. We observed different responses to roads in two deer species: red deer avoided roads, while roe deer browsed in the vicinity of roads.

CONCLUSIONS

A typical forest network of unpaved roads creates a landscape of fear for red and roe deer, and limits browsing pressure on regenerating forests due to the changes in deer habitat use, activity patterns and foraging behaviour. Knowledge of the factors influencing browsing pressure can help to spatially optimise the application of protective measures for tree seedlings.

Oil sands development in Alberta, Canada, results in a uniquely fragmented landscape characterized by a high-density of linear features and fewer polygonal disturbance features. Uncertainty remains concerning the effects of this type of industrial footprint on boreal mammal species and communities. We examined anthropogenic and natural factors that might exert the strongest positive and negative effects on the density of 11 species of boreal mammals and grouse species. We collected wildlife track density and anthropogenic feature data along 9-km snow tracking triangles. Eighty-six different triangle locations were sampled with 56 of these replicated from 2 to 13 winters between 2005 and 2018. Twenty-five explanatory variables were tested for their influence on track density by species. Variables were organized into anthropogenic, natural habitat, weather/season, small prey density and survey year. Industrial linear features did not significantly reduce trail density for most focal species. The exception were negative effects of low impact seismic lines on fisher (Pekania pennanti) and red squirrel (Tamiasciurus hudsonicus). Wider linear features including roads, pipelines and powerlines had positive effects on Canada lynx (Lynx canadensis), coyote (Canis latrans), white-tailed deer (Odocoileus virginianus) and red squirrel track density. Time of the year, snow depth and temperature were highly influential in terms of their composite across-species impact on density of small-bodied focal species. Upland deciduous forest had a strong positive effect on density of large-bodied mammals namely moose (Alces americanus), white-tailed deer and wolves (Canis lupus). Small prey density had a strong positive effect on Canada lynx, coyote, fisher, and ermine (Mustela erminea) density. Snowshoe hare (Lepus americanus) density followed a well-defined 10-yr cycle. Canada lynx, coyote and fisher followed this cycle with temporal density lags. The occurrence of well-defined snowshoe hare and predator cycles of typical amplitude, in spite of a >8%-per-year increase of human development footprint may indicate a system that demonstrated some resilience to the exploration and early-to mid-development stages of in situ oil sands exploration and production.

Dorsal guard hairs from three canid species that included red fox (Vulpes vulpes), golden jackal (Canis aureus), and gray wolf (Canis lupus) found in Turkey were characterized using light microscopy to study hair features, including cuticle and medullary patterns, medullary index, and hair length and root. The morphological features of the medulla and cuticle structures were quite similar for the three species, but a distinct difference was observed in the medulla structure. While the medullary structures of both C. aureus and C. lupus had a vacuolated structure, a multiseriate structure was observed in the V. vulpes. The average longest hairs (means ± standard errors [SE]) were observed to be 6.66 ± 1.22, 6.02 ± 0.75, and 5.56±5.57 cm in C. aureus, V. vulpes, and C. lupus, respectively. The highest medullary index (0.72 ± 0.52 μm) was recorded in C. aureus followed by V. vulpes (0.70 ± 0.56 μm) with the lowest in C. lupus (0.42 ± 0.64 μm). Hair root diameter values were determined as 136.27±29.25, 102.30±17.19, and 62.23±10.37 μm in C. lupus, C. aureus, and V. vulpes. Based on the one-way analysis of variance, significant differences were identified in both hair length and diameter, medullary diameter and index, and hair root diameter values among the three species (P <0.05).

The effects of anthropogenic climate change are apparent in the Greater Yellowstone Ecosystem (GYE), USA, with forest die-off, insect outbreaks, and wildfires impacting forest ecosystems. A long-term perspective would enable assessment of the historical range of variability in forest ecosystems and better determination of recent forest dynamics and historical thresholds. The objectives of this study were to (1) develop tree-ring chronologies for Engelmann spruce and Douglas fir growing at the study location, (2) correlate the annual ring widths of each species to monthly climate variables, (3) examine the instrumental climate data for regimes shifts in the mean state of variables, and (4) determine when ecological disturbances occurred through a quantification of growth releases. Finally, we discuss both climate-growth relationships and growth releases in the context of climate regime shifts and known forest disturbances. Engelmann spruce and Douglas fir showed some similar climate responses using moving correlation analysis including negative correlations between ring width and June – August current year temperature and previous growing season temperature. Regime shift analysis indicated significant (p < 0.05) shifts in minimum and maximum GYE temperature in the latter half of the 20th century. Disturbance analysis indicated that both tree species responded to wildfire and insect outbreak events with growth releases in up to 25% of the trees. Disentangling the influence of climate regime shifts and forest disturbances on the climate-growth relationships can be difficult because climate and forest disturbances are intricately linked. Our evidence indicates that regime shifts in monthly climate variables and forest disturbances as recorded by growth releases can influence the ring width response to climate over time. Trees are key to providing a long-term perspective on climate and ecological health across the GYE because they integrate both climate and ecology in their annual ring widths.

Caribou (Rangifer tarandus) populations are sympatric with Cougars (Puma concolor) in only a few areas, primarily in western Canada. Records of Cougar–Caribou interactions are limited and no published accounts describe Cougar predation on the shallow-snow, terrestrial-lichen-eating Northern Mountain Caribou (Rangifer tarandus caribou), referred to as Designatable Unit (DU) 7 by the Committee on the Status of Endangered Wildlife in Canada. In 2018 and 2019, two incidents of confirmed Cougar predation on radio-collared Caribou were documented in the declining Itcha-Ilgachuz subpopulation in west-central British Columbia. To the best of our knowledge, this is the first published record of Cougar predation on DU7 Northern Mountain Caribou. Increased landscape disturbance and climate change may be increasing apparent competition between deer (Odocoileus spp.), feral Horses (Equus ferus caballus), and Caribou, leading to Cougar predation in areas of Caribou range where it previously has not been documented. Cougar predation may become a conservation concern, as declining Caribou herds are susceptible to any increased predation pressure.

Understanding the causes behind population cycles is a fundamental issue in ecology, and has been the focus of research for decades. One of the most prominent examples of cyclical species is the snowshoe hare (Lepus americanus), a primary prey species in the North American boreal forest that exhibits fluctuations in abundance over 8-10 year periods. The potential for food limitation to cause changes in hare abundance has been examined, through manipulative experiments where individuals are provided supplemental food. However, in previous studies food addition was administered on a large scale, which can be problematic due to increased immigration to the area by other hares causing a predator-pantry effect. Individual-based food addition is therefore an optimal method for investigating the role of food in the snowshoe hare cycle. My primary objective was to evaluate the effect of food limitation on snowshoe hare demography, and determine the potential mechanisms of how food acts by examining behaviour, body condition, and physiology. To accomplish this, I radio-collared hares in the Kluane Lake area of the Yukon, Canada during the increase, peak, and decline phases of the cycle and fed a subset of individuals throughout the winter from selective feeders. These feeders allow controlled access to only the specific pit-tagged individuals. Over the study, I found that food supplementation significantly improved survival, with the largest effect size occurring during the population increase phase. Fed individuals also gave birth to larger offspring and produced larger litters. The demographic effects between treatments were likely driven by differences in behaviour or body condition. Fed individuals foraged less per day and spent a larger proportion of the foraging time vigilant, while maintaining a larger body mass. They further had reduced movement rates, and selected for conifer habitat relative to controls. Surprisingly, food supplementation increased hare stress responses to a hormone challenge, but simultaneously increased their ability to buffer against this heightened response, through increased corticosteroid binding capacity. Food supplemented individuals had improved body condition, in terms of blood indices and lower over-winter mass loss. I also found that changes in the indicators of stress in snowshoe hares during this study did not echo those observed in previous cycles. In combination, this thesis provides evidence that food limitation contributes to the demographic changes observed across the cycle, particularly in the increase phase, which may be caused by changes in behaviour and body condition.

Canis lupus (Gray Wolf or Wolf) can move great distances with important con-sequences to the persistence of isolated populations. We used global position system (GPS) tracking technology to summarize the movements of 4 Wolves translocated to Isle Royale National Park, MI, in 2018–2019. We quantified aspects of movement behavior of individual Wolves following translocation to, and an emigration event from the Isle Royale ecosystem. Introduction Immigration and emigration are essential to wildlife population growth and genetic diversity. Natural and anthropic barriers can limit these movements, affecting colonization of new areas and genetic exchange among populations (Adams et al. 2011, Mills 2013). Isle Royale National Park (IRNP) is an isolated island system in Michigan where reduced immigration and emigration has strongly influenced the is-land’s Canis lupus L. (Gray Wolf or Wolf) population (Hedrick et al. 2017). Wolves naturally colonized IRNP in the late 1940s, increased to 50 individuals by 1980, then declined to 10–20 Wolves by 2000 (Hedrick et al. 2017). After a 1997 immigra-tion event, the Wolf population increased to 30 Wolves in 2005, before declining to 2 Wolves by 2016 (Adams et al. 2011, Hedrick et al. 2017). Limited immigration and emigration were ascribed as main factors that lead to declining Wolf abundance on Isle Royale (Mech 2013, Vucetich et al. 2012). Unlike for mainland Wolf populations, immigration or emigration in the Isle Royale system is tied to ice-bridge formations across Lake Superior connecting the island to mainland (Fig. 1). To restore this apex predator and related ecosystem processes to IRNP, the US National Park Service (NPS) initiated Wolf translocations in 2018 (US Department of Interior 2018). Here, we describe first-year movements of initial Wolf transloca-tions to Isle Royale.

nimal domestication induces not only biological changes of the targeted populations but also cultural shifts in the way the domesticated populations are perceived. The grey wolf (Canis lupus) was certainly the first animal to be involved in this bio-cultural transformation, leading to the appearance of the dog (Canis familiaris), although the precise timing of this change is still intensely debated. Regardless, during the Late Upper Palaeolithic, alterations in the animal’s morphology and modifications in the behaviour of humans and “proto-domesticated dogs” alike consensually characterise the early steps of dog domestication. Once domesticated, the dog accompanied human groups in their migrations across the world until regions where the wolf has never been present, such as Australia and South America.

However, this strong relationship never became exclusive and human societies maintained strong economic and symbolic interactions with the dog’s wild relatives. In Europe, the red fox (Vulpes vulpes) has been used, by turns, as a companion8 and a fur provider. Archaeological evidence also points toward the symbolic consideration of the grey wolf in many cultures across the Northern hemisphere, as a participant in ritual paraphernalia. Nonetheless, all the studies aiming at precisely reporting the distinct role of dogs and non-dog canids in the archaeological record have been hampered by the difficulty in identifying the different canid species from their osteological remains.

With 18 to 20 identified species, the American continent hosts the largest diversity of canid species in the world. The multiplicity of biologically closely related species increases both the potential of interaction with human societies and the ambiguity in their skeletal identification. In Mesoamerica, four different species are present since the beginning of the Holocene: the grey fox (Urocyon cinereoargenteus), the coyote (Canis latrans), the Mexican wolf (Canis lupus baileiy) and the dog. Because of its very small size (on average 3-7 kg for a body length of 1 m including the tail), the grey fox can often be disentangled from the other species, even from fragmented bones. As a matter of fact, this species is often identified in the archaeozoological record, albeit never in large proportions. It is particularly scarce in the iconography and, unlike the other canids, the fox does not stand as a particular symbol in the Mesoamerican cosmogony (see Cultural background). Therefore, we focused our study on the identification of the three other species, namely the dog, the wolf and the coyote.

In this paper, we aim to deepen our understanding of the role of large canids (Canis spp.) in Mesoamerica by providing a more accurate identification of the canid remains to the species level. After introducing the ecological and cultural background of each animal, we first present a geometric morphometric (GMM) analysis of the first lower molar and test its efficiency to discriminate specimens of known species using a reference collection of 42 modern and archaeological specimens. This approach is then applied to 22 archaeological teeth, each from a unique individual, from four archaeological sites in central Mexico. The subsequent identification is used to contrast our perception of canid diversity and their interaction with human societies in the region.

Understanding how predators affect prey populations is a fundamental goal for ecologists and wildlife managers. A well‐known example of regulation by predators is the predator pit, where two alternative stable states exist and prey can be held at a low density equilibrium by predation if they are unable to pass the threshold needed to attain a high density equilibrium. While empirical evidence for predator pits exists, deterministic models of predator–prey dynamics with realistic parameters suggest they should not occur in these systems. Because stochasticity can fundamentally change the dynamics of deterministic models, we investigated if incorporating stochasticity in predation rates would change the dynamics of deterministic models and allow predator pits to emerge. Based on realistic parameters from an elk–wolf system, we found predator pits were predicted only when stochasticity was included in the model. Predator pits emerged in systems with highly stochastic predation and high carrying capacities, but as carrying capacity decreased, low density equilibria with a high likelihood of extinction became more prevalent. We found that incorporating stochasticity is essential to fully understand alternative stable states in ecological systems, and due to the interaction between top–down and bottom–up effects on prey populations, habitat management and predator control could help prey to be resilient to predation stochasticity.

The taxonomic status of extinct Japanese or Honshu wolves (Canis lupus hodophilax) has been disputed since the name hodophilax was first proposed by Temminck in 1839 on the basis of specimens stored in Leiden, the Netherlands. Points of controversy include whether the type specimen of hodophilax (Jentink c: RMNH.MAM.39181) and the other two specimens from Leiden (Jentink a: RMNH.MAM.39182 and Jentink b: RMNH.MAM.39183) represent different varieties or subspecies of Japanese wolves or not. Two Japanese names, ookami and jamainu, used to describe wild Canis species, further complicate the issue. In this study, the taxonomic status of Japanese wolves was clarified using mitochondrial DNA of the three specimens stored at the Naturalis Biodiversity Center in Leiden, in addition to three Japanese wolf specimens stored at the Museum für Naturkunde in Berlin and five new samples from Japan. The mitochondrial genomes of the type specimen of hodophilax (Jentink c) and another sample from Leiden (Jentink b) as well as Berlin specimens were included in the cluster of Japanese wolves distinct from other grey wolves. However, the other sample from Leiden (Jentink a) was identified as a domestic dog. A mitochondrial genome analysis suggested that Japanese wolves could be categorized into two distinct clusters. Studies of nuclear genomes are needed to further clarify the taxonomic status, divergence time, and population genetic structure of Japanese wolves.

Context: Ungulate prey can use increased vigilance to reduce their risk of predation, but little is known of the combined and interactive risk effects from humans and wolves in determining ungulate behaviour across time and space. Understanding the interplay between these risk effects is increasingly important, considering the recolonisation of several large carnivores to more human-dominated landscapes in Europe.

Aim: The aim of the present study was to assess the vigilance behaviour expressed by red deer (Cervus elaphus) in response to both humans and wolves in the Polish Białowieża Forest.

Methods: Using a camera-trap transect, the effect of distance to human settlements, hunting season, patterns of space use by wolves (Canis lupus), canopy openness, canopy height, time of day, as well as sex/age of individuals, on the vigilance behaviour observed in red deer was studied using a model-selection approach.

Key results: We did not find a clear effect of patterns of space use by wolves or distance to human settlements on red deer vigilance behaviour at the landscape scale. However, red deer showed increased vigilance during the hunting season and during the day outside of protected areas and reserves, because disturbance from human hunters is highest. Conversely, we also found that red deer were more vigilant at night within more protected areas, which is likely to be explained by the increased activity of wolves because human activity is strictly limited.

Conclusions: Our study showed that vigilance behaviour of red deer in Białowieża Primeval Forest is more driven by human hunting than by the frequency of wolf presence at a landscape scale. This could be explained by the higher temporal and spatial predictability of human hunting activities than wolf risk. We found that patterns of wolf space use, as opposed to the omnipresent fear effects from humans, had only localised effects by increasing vigilance levels during night hours in non-hunting areas of the forest. The reverse was observed outside of protected reserves. Understanding how prey species respond to this new combination of risk from natural predators and humans, is increasingly important in a landscape where human risk is becoming ever more potent and carnivores recolonise.

The depopulation of rural areas by humans (or rural exodus) in southern Europe, and the associated abandon-ment of cropland, had marked ecological consequences on wildlife, which became evident approximately fifteen years ago. Shrub and tree encroachment, and the expansion of forest birds and the formerly persecuted mammalian ungulates and carnivores, were highlighted as the major consequences of the rural exodus in Italy. In this report, we provide a more integrative view, and show that a rural exodus also explains other ecological phenomena that are usually treated independently. After reviewing the ecological consequences of the rural exodus that has been affecting a large part of Spain during the last six decades, we suggest that this set of ecological consequences also includes the movement of shy-selected predators and large, big game species out of their former ecological refuges, as well as increased frequencies of individuals with bolder-behaviours in recovering populations. We develop a tentative conceptual model linking the increasing approach of wildlife to anthropogenic habitats and human depopulation of rural areas. These links are created by the increasing diffi-culty to survive and reproduce in recovering, high-predation wild areas, due to mesopredator release and the loss of fear to humans, among other factors. We acknowledge that the recovery of formerly persecuted wildlife in depopulated landscapes has been helped by conservation policies, but we suggest that policies alone cannot explain the observed changes. Finally, we propose that the processes we analyse on a national scale could be taking place in Europe on a continental scale as well, and will most likely occur in the future in other regions of the world, with the current growth in economies.

The foraging movement behaviour of grey wolves (Canis lupus) is unique in summer, when the focus of breeding wolves is on taking care of new-born offspring. In this study, we analysed the movement tracks of nine radio-collared wolves during their pup-rearing season. The wolves lived in the boreal zone in Finland. Our interest was in analysing the foraging trips of wolves from the den site, which serves as the cen-tral place of the pack in summer. Based on the information on spatial relocations and time of the day, the movement tracks of the collared wolves were split into seg-ments using hidden Markov models (HMM). Those segments were considered to be produced by different movement behaviour modes that were not observed. We first split the movement tracks of the wolves into separate foraging trips using a two-state HMM and further extracted the different movement modes with a four-state HMM. The modes were interpreted as rest, moderate activity, homing to the den site and fast movement for other purposes, such as leaving the den. Our analysis showed that, for most of the individuals, the movement during homing was fastest and most persistent. This research highlights the foraging behaviour of wolves during the pup-rearing season, which has gained less attention than the nomadic behaviour outside of this season. Our study showed how cyclic foraging trips can be divided into be-havioural phases using HMMs, and how these behaviours appear in different times of the day. These results have potential uses, for instance, when studying the habitat requirements and usage or assessing the risk of human–wildlife conflicts.