Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

Beaver activity increases aquatic subsidies to terrestrial consumers

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Ancient DNA evidence for the loss of a highly divergent brown bear clade during historical times

The genetic diversity of present-day brown bears (Ursus arctos) has been extensively studied over the years and appears to be geographically structured into five main clades. The question of the past diversity of the species has been recently addressed by ancient DNA studies that concluded to a relative genetic stability over the last 35,000 years. However, the post-last glacial maximum genetic diversity of the species still remains poorly documented, notably in the Old World. Here, we analyse Atlas brown bears, which became extinct during the Holocene period. A divergent brown bear mitochondrial DNA lineage not present in any of the previously studied modern or ancient bear samples was uncovered, suggesting that the diversity of U. arctos was larger in the past than it is now. Specifically, a significant portion (with respect to sequence divergence) of the intraspecific diversity of the brown bear was lost with the extinction of the Atlas brown bear after the Pleistocene/Holocene transition.     

The impacts of beavers Castor spp. on biodiversity and the ecological basis for their reintroduction to Scotland,UK

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Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore

The increased abundance of large carnivores in Europe is a conservation success, but the impact on the behavior and population dynamics of prey species is generally unknown. In Europe, the recolonization of large carnivores often occurs in areas where humans have greatly modified the landscape through forestry or agriculture. Currently, we poorly understand the effects of recolonizing large carnivores on extant prey species in anthropogenic landscapes. Here, we investigated if ungulate prey species showed innate responses to the scent of a regionally exterminated but native large carnivore, and whether the responses were affected by human‐induced habitat openness. We experimentally introduced brown bear Ursus arctos scent to artificial feeding sites and used camera traps to document the responses of three sympatric ungulate species. In addition to controls without scent, reindeer scent Rangifer tarandus was used as a noncarnivore, novel control scent. Fallow deer Dama dama strongly avoided areas with bear scent. In the presence of bear scent, all ungulate species generally used open sites more than closed sites, whereas the opposite was observed at sites with reindeer scent or without scent. The opening of forest habitat by human practices, such as forestry and agriculture, creates a larger gradient in habitat openness than available in relatively unaffected closed forest systems, which may create opportunities for prey to alter their habitat selection and reduce predation risk in human‐modified systems that do not exist in more natural forest systems. Increased knowledge about antipredator responses in areas subjected to anthropogenic change is important because these responses may affect prey population dynamics, lower trophic levels, and attitudes toward large carnivores. These aspects may be of particular relevance in the light of the increasing wildlife populations across much of Europe.     

Changes in wolf spider (Araneae) assemblages across woodland–pasture boundaries in the central wheat‐belt of New South Wales,Australia

The abundance of wolf spiders (Lycosidae) was measured across woodland–pasture boundaries in the wheat‐belt of New South Wales, Australia, to determine the nature and magnitude of any edge effect. Spiders were collected by spotlighting along sample plots in woodlands located at distances of 5, 20, 35 and 200 m from the edge, and along sample plots in paddocks located at distances of 5 and 20 m from the edge. The wolf spider assemblage changed significantly across the edge, but the difference could be accounted for only by a change between the woodland and the paddock and not by any changes within the woodland at different distances from the edge. Ground cover (wolf spider microhabitat) changed significantly between the paddock and the woodland, but there were no consistent differences in microhabitat with distance from edge within either paddocks or woodlands. There was a significant correlation between an ordination of sites based on spider species abundance and an ordination based on microhabitat variables, suggesting that the wolf spider assemblage was responding to differences in microhabitat. Fine‐scale selection of microhabitat by most wolf spider species was non‐random, with most species preferring locations with grass cover, rather than more open locations. The present study indicates that wolf spiders are mostly unaffected by edge conditions at the woodland–paddock boundary. Accordingly, small and/or linear remnants with high edge‐to‐area ratios may constitute suitable faunal habitat for wolf spiders and perhaps other terrestrial arthropod species, despite the fact that this configuration is unsuitable for many vertebrate species.     

Black bear (<Emphasis Type="Italic">Ursus americanus</Emphasis>) and wolf (<Emphasis Type="Italic">Canis</Emphasis> spp.) summer diet composition and ungulate prey selectivity in Ontario,Canada

Understanding predator-prey dynamics is an important component of management strategy development for wildlife populations that are directly affected by predation. Ungulates often serve as a significant source of prey for many large mammal predators, and patterns of predation are known to influence population dynamics. Although black bear and wolf diets have been investigated extensively, prey preference has been less commonly examined, especially in analyses that take into account age class (i.e., juvenile and adult) of the ungulate prey. We examined black bear (Ursus americanus), wolf (Canis lupus), and hybrids (Canis spp.) prey preference in Ontario based on the availability of three ungulate species—elk (Cervus elaphus), moose (Alces alces), and white-tailed deer (Odocoileus virginianus). We analyzed the presence of prey items in black bear and wolf scats collected over 3 years by examining prey hair cuticular scale patterns. We applied correction factors to frequencies of occurrence of prey items found in predator scat and related diet composition to the availability of ungulates, determined by fecal pellet transect surveys. In addition, non-ungulate diet items were identified to obtain full diet composition profiles. We found that black bear diet consisted of more than 87% vegetation, and they were opportunistic, not selecting for any particular ungulate species in either adult or juvenile age class. Wolf diet was comprised mainly of ungulates (~?73.2%), muskrat (Ondatra zibethica; ~?8.5%), and beaver (Castor canadensis; ~?14.6%), and although moose were at least 1.5 times more abundant then each of the other ungulate prey species in the study area, wolves preferred elk, using moose less than expected. Although we found black bear diet to be opportunistic during the summer, wolves in our study heavily utilized both juvenile and adult ungulates, however, among ungulate species, displayed preference for elk. The preference displayed by wolves provides insight that wildlife managers can use to guide further investigation and assist with the development of strategies to ensure continued elk reintroduction success, and moose and white-tailed deer population sustainability.     

Restoring ecological balance to the British mammal fauna

• 1 The mammal fauna of the British Isles has seen major perturbations since the end of the last ice age, some natural and some anthropogenic. Today, 61 species of terrestrial mammals breed in the British Isles, but only 39 of them are native species, the rest have been introduced. Furthermore, 19 native species have disappeared from the fauna including all the large predators, lynx Lynx lynx, wolf Canis lupus and brown bear Ursus arctos.
• 2 Inevitably, these changes in species composition have been accompanied by major changes in community function including changes in patterns of energy flow through the mammalian community. For example, a high percentage of all the energy now flows through the introduced rabbit Oryctolagus cuniculus population and red deer Cervus elaphus, which, in the absence of natural predators, are living at extremely high densities.
• 3 Could the reintroduction of species help to reverse such changes in community structure and function? The successful return to the wild of species such as the Arabian oryx Oryx leucoryx and beaver Castor fiber has shown that reintroductions are certainly possible. However, the impact on community function of returned species is more difficult to evaluate.
• 4 This question is addressed in relation to the consequences for deer populations of any possible reintroduction of the wolf to Scotland. Based on what we know of wolves elsewhere, predictions are made about the likely demography and patterns of killing behaviour of introduced wolves. These values are then used to parameterize a Leslie matrix simulation of the impact of wolves on contemporary populations of red deer in Scotland. The simulations suggest very strongly that wolves are very unlikely to have any significant impact on the high‐density populations of deer now living in the Scottish landscape.

     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Evaluating the Mechanisms of Landscape Change on White-Tailed Deer Populations

Understanding how landscape change influences the distribution and densities of species, and the consequences of these changes, is a central question in modern ecology. The distribution of white-tailed deer (Odocoileus virginianus) is expanding across North America, and in some areas, this pattern has led to an increase in predators and consequently higher predation rates on woodland caribou (Rangifer tarandus caribou)—an alternate prey species that is declining across western Canada. Understanding the factors influencing deer distribution has therefore become important for effective conservation of caribou in Canada. Changing climate and anthropogenic landscape alteration are hypothesized to facilitate white-tailed deer expansion. Yet, climate and habitat alteration are spatiotemporally correlated, making these factors difficult to isolate. Our study evaluates the relative effects of snow conditions and human-modified habitat (habitat alteration) across space on white-tailed deer presence and relative density. We modeled deer response to snow depth and anthropogenic habitat alteration across a large latitudinal gradient (49° to 60°) in Alberta, Canada, using motion-sensitive camera data collected in winter and spring from 2015 to 2019. Deer distribution in winter and spring were best explained by models including both snow depth and habitat alteration. Sites with shallower snow had higher deer presence regardless of latitude. Increased habitat alteration increased deer presence in the northern portion of the study area only. Winter deer density was best explained by snow depth only, whereas spring density was best explained by both habitat alteration and the previous winter's snow depth. Our results suggest that limiting future habitat alteration or restoring habitat can alter deer distribution, thereby potentially slowing or reversing expansion, but that climate plays a significant role beyond what managers can influence. © 2020 The Wildlife Society.     

Spatial variability in wolf diet and prey selection in Galicia (NW Spain)

We studied wolf (Canis lupus) diet for three different landscapes in the north-western Iberian Peninsula, differing in land uses and availability of food for wolves. We examined 2740 scats, collected over a period of 4 years, in order to describe wolf diet, its geographic variation, and trophic preferences. The most consumed species were wild pony, roe deer and cattle. We observed differences in wolf diet among the three study sites, related to the availability and accessibility of food resources in each habitat. For the two study sites in northern and central Galicia, wolves showed similar diet, with high occurrence of wild pony (37 vs. 34%) and cattle (20 vs. 23%), but differing in the consumption of wild ungulates (16 vs. 8%) and carrion (7 vs. 14%). Wolf diet in eastern Galicia’s mountain ranges was entirely different, due to the higher consumption of wild ungulates (70%). Wolves showed clear prey selection patterns. Between wild ponies and livestock, wolves positively selected ponies. Among wild ungulates, wolves positively selected roe deer. Wild pony and roe deer are key species for wolf feeding in Galicia. In the Galician wild pony range, ponies are the main food for wolves. Given that the availability of wild ponies may contribute to the decrease in wolf predation on cattle, it is essential to develop innovative administrative decisions in such areas to preserve this traditional equid population. In the same way, the population of roe deer should be strengthened in the livestock areas outside the range of wild pony.     

Serum concentrations of vitamin D metabolites, vitamins A and E, and carotenoids in six canid and four ursid species at four zoos

Nutritional status for six captive canid species (n=34) and four captive ursid species (n=18) were analyzed. The species analyzed included: African wild dog (Lycaon pictus), arctic fox (Alopex lagopus), gray wolf (Canis lupus), maned wolf (Chrysocyon brachyurus), Mexican wolf (Canis lupus baleiyi), red wolf (Canis rufus), brown bear (Ursus arctos), polar bear (Ursus maritimus), spectacled bear (Tremarctos ornatus), and sun bear (Ursus malayanus). Diet information was collected for these animals from each participating zoo (Brookfield Zoo, Fort Worth Zoo, Lincoln Park Zoological Gardens, and North Carolina Zoological Park). The nutritional composition of the diet for each species at each institution met probable dietary requirements. Blood samples were collected from each animal and analyzed for vitamin D metabolites 25(OH)D and 1,25(OH)(2)D, vitamin A (retinol, retinyl stearate, retinyl palmitate), vitamin E (alpha-tocopherol and gamma-tocopherol) and selected carotenoids. Family differences were found for 25(OH)D, retinol, retinyl stearate, retinyl palmitate and gamma-tocopherol. Species differences were found for all detectable measurements. Carotenoids were not detected in any species. The large number of animals contributing to these data, provides a substantial base for comparing the nutritional status of healthy animals and the differences among them.     

Conservation genetics of carnivores in Italy

Pleistocene climatic changes shaped the patterns of biodiversity in Europe and around the Mediterranean. Describing the phylogeographic structure of animal populations and inferring past population dynamics is essential to develop a framework for conservation biology in Europe. Direct persecution, habitat loss, population fragmentation and hybridization with domesticated conspecifics, are the main threats to the survival of large mammalian species. In this paper I will summarize the available information on phylogeography and population genetics of brown bear, wolf, wildcat and otters in Italy and in Europe.     

National‐scale analyses of habitat associations of Marsh Tits Poecile palustris and Blue Tits Cyanistes caeruleus: two species with opposing population trends in Britain

Capsule Marsh Tits were strongly associated with both the amount and species diversity of woodland understorey; Blue Tits were associated with large trees and deadwood.

Aims To gather quantitative information on the habitat requirements of Marsh Tits, in comparison with those of Blue Tits, across a large number of sites in England and Wales, and secondly to evaluate the range of habitat conditions likely to encourage the presence, and increase the abundance of, each species.

Methods Counts of birds were made at each of 181 woods across England and Wales, and habitat data were collected from the same locations in each woodland. Marsh Tit and Blue Tit presence and abundance were related to habitat characteristics, interspecific competition and deer impact.

Results Shrub cover and species diversity were important for the presence and abundance of Marsh Tits, across their geographical range in Britain. Blue Tits were associated with large trees and deadwood.

Conclusion Our results support the hypothesis that changes in woodland management, leading to canopy closure and a decline in the understorey available, could have had an impact on Marsh Tits, and may have led to the observed population decline. These same changes were also consistent with population increase in Blue Tits.     

Factors affecting habitat occupancy by wolves in northern Apennines (northern Italy): a model of habitat suitability

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.     

19th century woodland structure controls stand-scale epiphyte diversity in present-day Scotland

Axioms developed from island biogeography theory (i.e. species–area relationships, effects of fragmentation and isolation) are central to the development of conservation strategy. Within this context, the 'extinction debt' hypothesis brings into question an often assumed relationship between species richness and present-day spatial habitat structure (i.e. extent, fragmentation), suggesting instead that the richness and composition of biological communities may lag behind spatial changes in habitat. We examined evidence for an extinction debt among epiphytic lichens, a highly diverse biological group of significant conservation concern. Using sites in Scotland, we compared epiphyte species richness in smaller-scale habitat units (aspen stands) to larger-scale woodland structure (extent and fragmentation) measured at two spatial scales (1 km² and 4 km²) and for two timeframes, modern (1990s to 2000s) and historic (1860s to 1880s). Species richness was positively related to woodland extent and negatively related to woodland fragmentation; however, richness was explained better by historic woodland structure at a 1-km² scale, than by modern woodland structure. The results indicate: (1) a coupling of stand-scale epiphyte assembly and dynamics of the wider woodland ecosystem, and (2) a significant lag in the response of epiphyte species richness to habitat spatial structure. However, the effect of spatial habitat structure is different between species groups with contrasting traits. The effect of decreasing woodland extent on epiphyte richness is generally more severe for microlichens (comprising a greater number of rare and specialist species) than the more generalist macrolichens.     

Responses of wolf feeding habits after adverse climatic events in central-western Belarus

Many studies have investigated the ecology of wolf populations of Eurasia, showing that although wolves are mostly opportunistic in seeking meso-large enough mammalian prey, they can also be selective, depending on local availability of prey and their population biomass. Yet prey preferences of the wolf have been poorly evaluated in situations of complex predator/prey systems because such ecological situations are extremely rare in Europe. In particular, the role of beaver is poorly known due to the extreme decline in its range and population over the last few centuries.We conducted a 15-year study (1999–2014) of wolf Canis lupus diet in the Naliboki forest of central-western Belarus to determine the dietary responses of the wolf population in a context of a rich prey supply (beaver 650 inds/100 km², elk 47 inds/100 km², red deer 98 inds/100 km, roe deer 398 inds/100 km², wild boar 234 inds/100 km²). The bison, although present, is not preyed on. We compared the seasonal and annual diet variations of both wolf adults and pups, by scat analysis and hair identification. In winter 2012–2013, the winter was quite harsh with a long period of snow, which severely affected the roe deer and wild boar populations. Five severe summer droughts also occurred (1999, 2001, 2002, 2004 and 2013), greatly decreasing the water level in rivers and canals. We took advantage of these stressful events to evaluate the diet responses of the wolves.In “normal” years, we identified 11 food categories, essentially beaver and medium-sized ungulates (66%), and large ungulates to a lesser extent (9% in summer, 20% in winter). The adults were found to selectively supply pups with beaver, probably because of its easy transportability. Beaver consumption also increased during summer droughts when water levels were very low. After the harsh winter of 2012–2013, which was followed by a sharp decline in medium-sized prey, we observed a shift in the winter diet breadth of the wolves towards greater consumption of both large wild ungulates and small carnivores. We concluded that:1. Beaver is a functional element in wolf ecology, as a primary food for adults and pups;2. A large range of available prey species is important to maintaining a viable wolf population in cases of extreme climatic events.     

The importance of spatial scale in habitat selection by European beaver

We evaluated habitat selection by European beaver Castor fiber L. across a spatial gradient from local (within the family territory) to a broad, ecoregional scale. Based on aerial photography, we assessed the habitat composition of 150 beaver territories along the main water bodies of the Vistula River delta (northern Poland) and compared these data with 183 randomly selected sites not occupied by the species. The beavers preferred habitats with high availability of woody plants, including shrubs, and avoided anthropogenically modified habitats, such as arable lands. Within a single family territory, we observed decreasing woody plant cover with increasing distance from a colony centre, which suggests that beaver habitat preferences depend on the assessment of both the abundance and spatial distribution of preferred habitat elements. We tested the importance of spatial scale in beaver habitat selection with principal coordinates of neighbour matrices analysis, which showed that the geographical scale explained 46.7% of the variation in habitat composition, while the local beaver density explained only 10.3% of this variability. We found two main spatial gradients that were related to the broad spatial scale: first, the most important gradient was related to the largest distances between beaver sites and was independent of woody plant cover and the local beaver site density. The second most important gradient appeared more locally and was associated with these variables. Our results indicate that European beaver habitat selection was affected by different scale‐related phenomena related 1) to central place foraging behaviour, which resulted in the clumped distribution of woody plants within the territory, and 2) local population density and woody plant cover. Finally, 3) habitat selection occurs independently across the largest spatial scale studied (e.g. between watersheds), which was probably due to the limited natal dispersal range of the animals.