Resource separation analysis with moose indicates threats to caribou in human altered landscapes

Species recovery is often impeded by inadequate knowledge on mechanisms of community interactions that cause and exacerbate species endangerment. Caribou and wild reindeer Rangifer tarandus are declining in many regions of their circumpolar range likely because of human‐induced landscape changes. In general, their niche specialization enables Rangifer to survive in nutrient‐poor habitats spatially separated from other ungulates and their shared predators. Research has indicated that shifts in primary prey distribution following human landscape alteration may result in spatial overlap with Rangifer. We studied overlap relationships of woodland caribou R. t. caribou and moose Alces alces, quantified by their differential use of environmental resources, and evaluated the role of human landscape alteration in spatial separation in south‐western Canada. Anthropogenic conversion of old‐growth forests to early seral stands is hypothesized to decrease the spatial separation between caribou and moose, the dominant prey for wolves Canis lupus, contributing to increased caribou mortality. Redundancy analysis (RDA) was first used to examine coarse scale resource separation across our study area. Second, at a finer spatial scale, we used logistic regression to compare resource‐ and spatial separation of sympatric pairs of 17 moose and 17 caribou. Finally, we tested if the frequency of predator‐caused caribou mortalities was higher in regions with higher moose resource use. Although environmental resource separation was strong at the coarser scale, we observed substantial spatial overlap (>50%) at the finer scale. In summer we reported a significant positive relationship between spatial overlap of moose and caribou and the degree of human landscape alteration. Most importantly, locations of caribou mortalities corresponded with areas of high resource use by moose in summer. Thus, consistent with the spatial separation hypothesis, our research suggests that early successional forest stages may decrease spatial separation between caribou and moose, resulting in increased mortality risk for threatened caribou.     

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     

Trophic niche partitioning between two prey and their incidental predators revealed various threats for an endangered species

Documenting trophic niche partitioning and resource use within a community is critical to evaluate underlying mechanisms of coexistence, competition, or predation. Detailed knowledge about foraging is essential as it may influence the vital rates, which, in turn, can affect trophic relationships between species, and population dynamics. The aims of this study were to evaluate resource and trophic niche partitioning in summer/autumn between the endangered Atlantic‐Gaspésie caribou (Rangifer tarandus caribou) population, moose (Alces americanus) and their incidental predators, the black bear (Ursus americanus) and coyote (Canis latrans), and to quantify the extent to which these predators consumed caribou. Bayesian isotopic analysis showed a small overlap in trophic niche for the two sympatric ungulates suggesting a low potential for resource competition. Our results also revealed that caribou occupied a larger isotopic niche area than moose, suggesting a greater diversity of resources used by caribou. Not surprisingly, coyotes consumed mainly deer (Odocoileus virginianus), moose, snowshoe hare (Lepus americanus), and occasionally caribou, while bears consumed mainly vegetation and, to a lesser extent, moose and caribou. As coyotes and bears also feed on plant species, we documented trophic niche overlap between caribou and their predators, as searching for similar resources can force them to use the same habitats and thus increase the encounter rate and, ultimately, mortality risk for caribou. Although the decline in the Gaspésie caribou population is mostly driven by habitat‐mediated predation, we found evidence that the low level of resource competition with moose, added to the shared resources with incidental predators, mainly bears, may contribute to jeopardize the recovery of this endangered caribou population. Highlighting the trophic interaction between species is needed to establish efficient conservation and management strategies to insure the persistence of endangered populations. The comparison of trophic niches of species sharing the same habitat or resources is fundamental to evaluate the mechanisms of coexistence or competition and eventually predict the consequences of ecosystem changes in the community.     

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.     

Moose movement rates are altered by wolf presence in two ecosystems

Predators directly impact prey populations through lethal encounters, but understanding nonlethal, indirect effects is also critical because foraging animals often face trade‐offs between predator avoidance and energy intake. Quantifying these indirect effects can be difficult even when it is possible to monitor individuals that regularly interact. Our goal was to understand how movement and resource selection of a predator (wolves; Canis lupus) influence the movement behavior of a prey species (moose; Alces alces). We tested whether moose avoided areas with high predicted wolf resource use in two study areas with differing prey compositions, whether avoidance patterns varied seasonally, and whether daily activity budgets of moose and wolves aligned temporally. We deployed GPS collars on both species at two sites in northern Minnesota. We created seasonal resource selection functions (RSF) for wolves and modeled the relationship between moose first‐passage time (FPT), a method that discerns alterations in movement rates, and wolf RSF values. Larger FPT values suggest rest/foraging, whereas shorter FPT values indicate travel/fleeing. We found that the movements of moose and wolves peaked at similar times of day in both study areas. Moose FPTs were 45% lower in areas most selected for by wolves relative to those avoided. The relationship between wolf RSF and moose FPT was nonlinear and varied seasonally. Differences in FPT between low and high RSF values were greatest in winter (?82.1%) and spring (?57.6%) in northeastern Minnesota and similar for all seasons in the Voyageurs National Park ecosystem. In northeastern Minnesota, where moose comprise a larger percentage of wolf diet, the relationship between moose FPT and wolf RSF was more pronounced (ave. across seasons: ?60.1%) than the Voyageurs National Park ecosystem (?30.4%). These findings highlight the role wolves can play in determining moose behavior, whereby moose spend less time in areas with higher predicted likelihood of wolf resource selection.     

Food habits of three carnivores in a mosaic landscape of São Paulo state,Brazil

São Paulo is the most developed state in Brazil, but despite intense land use changes, some mammal species still inhabit remnant fragments of native vegetation of the state. We used fecal analyses to investigate prey diversity, trophic niche breadth, and trophic niche overlap of pumas, ocelots, and maned wolves in protected and non-protected areas in São Paulo state. During 18 months, we inspected more than 10,000 km of roads and trails and collected 233 fecal samples. Maned wolves showed generalist food habit, ocelots were predominantly carnivores, and pumas were strict carnivores. The European hare, an exotic species which had not yet been reported in the study area, was identified in the feces of a puma. Trophic niche breadth was the lowest for maned wolves (BA?=?0.339) and the highest for pumas (BA?=?0.723). Trophic niche overlap was high between maned wolves and ocelots (O_jk?=?0.765) and between pumas and ocelots (O_jk?=?0.639). Significantly different diets of maned wolves and ocelots were observed between dry and rainy seasons. Our findings indicate that these three species may adapt their diets to the disturbed landscapes. This is the second study characterizing food habits of ocelots in Cerrado areas but the first considering a large number of samples.     

Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Feeding ecology and seasonal diet overlap between Stellifer brasiliensis and Stellifer stellifer in a tropical estuarine ecocline

Seasonal freshwater discharge was important for defining habitat utilization by different ontogenetic phases of Stellifer brasiliensis and Stellifer stellifer along the estuarine ecocline. The middle estuary was important as a nursery and feeding ground for young‐of‐the‐year, and a feeding ground for sub‐adults and adults of both species. These species are zoobenthivorous, but during their life cycle and between different habitats and seasons, their trophic guild can change to opportunist and zooplanktivore. During the late rainy season in the lower estuary, all phases of both species, except juveniles of S. brasiliensis and adults of S. stellifer, showed a niche overlap indicating similarity in prey utilization. The diet composition was qualitatively similar, showing an evident niche overlap of intra and interspecific competition among the Stellifer spp. Although the niches of these species appeared to significantly overlap, some resource partitioning patterns were apparent. The niche overlap was significantly reduced due to the seasonal difference in habitat use and prey consumption along the ecocline of the estuary by different ontogenetic phases. The ingestion of blue nylon fragments by both species was observed and quantified.     

Natural regulation of cervidae along a 1000 km latitudinal gradient: Change in trophic dominance

Summary The biomass of forage, herbivores (caribou and moose) and predators (wolf) were estimated for four assemblages of large mammals along a latitudinal gradient in the Québec-Labrador peninsula and related to predictions made by two types of multitrophic level models. Wolves were present in three study areas, but they had been extirpated in the last one. Annual production of preferred forage exhibited a clear north-south increase for moose, but not for caribou. Neither the herbivore nor predator biomass increased along the latitudinal gradient: the highest herbivore biomass occurred in the wolf-free area and in the northernmost site, while the greatest predator density was observed in the southernmost site. Consequently, the ratio of the herbivore to forage biomass was the highest in the area devoid of wolves and in the northernmost site occupied by migratory caribou. Availability of forage per herbivore was the greatest in the moose-wolf and the caribou-moose-wolf assemblages. The observed data supported the multitrophic level model incorporating classical predator-prey relationships and producing stepwise accrual of trophic level biomass with increasing food chain length. In the northernmost site, the system was limited to two functional trophic levels and caribou were regulated by summer forage. Three functional trophic levels appeared to exist in the central study area where caribou and moose were preyed upon by wolves. Both herbivores were at very low density, the first one due probably to its poor adaptation to predation and the second because of an unproductive range. In the southernmost site, moose were clearly regulated by predation and kept much below the carrying capacity. With the extirpation of wolves in the last study area, moose were regulated by forage and the density exceeded that in the moose-wolf system by seven times even in a less productive range. Caribou, having primarily evolved under resource limitation, is replaced by a cervid better adapted to predation, the moose, in more productive three-link ecosystems.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

Niche overlap and diet composition of three sympatric coastal dolphin species in the southwest Atlantic Ocean

Sympatric species are expected to differ in ecological requirements to minimize niche overlap and avoid competition. Here we assess the trophic interactions among three coexisting dolphin species from southern Brazil: the franciscana dolphin (Pontoporia blainvillei), the Guiana dolphin (Sotalia guianensis), and the Lahille's bottlenose dolphin (Tursiops truncatus gephyreus). We evaluated temporal variation in carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen to examine potential dietary shifts resulting from increased fishing activity over the past three decades. We estimated the degree of niche overlap among these species and the contribution of potential prey sources to their diet. δ¹⁵N values were consistent among species and across years, while δ¹³C values increased for Guiana dolphins and decreased for bottlenose dolphins, suggesting changes in diet and/or foraging habitats through time. The similar δ¹³C and δ¹⁵N values and the high niche overlap between Guiana and bottlenose dolphins indicate that these species are primarily feeding on demersal prey. The franciscana diet is primarily composed of pelagic prey, resulting in a lower niche overlap in comparison with the other dolphin species. Our study provides further information about the foraging ecology of this unique dolphin community in southern Brazil with implications for its management and conservation.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

Unveiling trade‐offs in resource selection of migratory caribou using a mechanistic movement model of availability

Habitat selection is a multi‐level, hierarchical process that should be a key component in the balance between food acquisition and predation risk avoidance (food–predation trade‐off). However, to date, studies have not fully elucidated how fine‐ and broad‐scale habitat decisions by individual prey can help balance food versus risk. We studied broad‐scale habitat selection by Newfoundland caribou Rangifer tarandus, focusing on trade‐offs between predation risk versus access to forage during the calving and post‐calving period. We improved traditional measures of habitat availability by incorporating fine‐scale movement patterns of caribou into the availability kernel, thus enabling separation of broad and fine scales of selection. Remote sensing and field surveys served to create a spatio‐temporal model of forage availability, whereas GPS telemetry locations from 66 black bears Ursus americanus and 59 coyotes Canis latrans provided models of predation risk. We then used GPS telemetry locations from 114 female caribou to assess food–predation trade‐offs through the prism of our refined model of caribou habitat availability. We noted that migratory movements of caribou were oriented mainly towards habitats with abundant forage and lower risk of bear and (to a lesser extent) coyote encounter. These findings were generally consistent across caribou herds and would not have been evident had we used traditional methods instead of our refined model when estimating habitat availability. We interpret these findings in the context of stereotypical migratory behaviour observed in Newfoundland caribou, which occurs despite the extirpation of wolves Canis lupus nearly a century ago. We submit that caribou are able to balance food acquisition against predation risk using a complex set of factors involving both finer and broader scale selection. Accordingly, our study provides a strong argument for using refined habitat availability estimates when assessing food–predation trade‐offs.     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.     

End of the road: Short-term responses of a large mammal community to forest road decommissioning

Anthropogenic disturbances are increasing worldwide, causing wildlife habitat loss, alteration, and fragmentation. In Canada, the decommissioning of linear anthropogenic structures is identified as a promising tool to restore the habitat of threatened populations of boreal caribou (Rangifer tarandus caribou) by reducing food availability for alternate prey and decreasing encounter probabilities with predators. In this study, we monitored the use of 40 km of decommissioned forest roads by caribou, gray wolves (Canis lupus), black bears (Ursus americanus), and moose (Alces americanus) 1–3 years after reclamation, using 232 motion-activated camera traps. We compared four additive treatments (meaning that each successive treatment included the treatment prior): closing the road to human access, decompacting its soil, planting black spruce (Picea mariana) trees, and adding enriched soil. We assessed the influence of treatments, use by other large mammals, and characteristics of the surrounding environment on road use by the four species. Caribou used the planted treatment (which also included closing and decompacting) more than the closed-only (reference) treatment, but treatments did not influence the use of decommissioned roads by bears and moose. We could not assess the use of treated roads by wolves because of low sample size. Road use by caribou declined with local moose density, but increased with local bear density. Caribou were observed more frequently on roads surrounded by regenerating and mature coniferous stands; caribou also preferentially used roads surrounded by wetlands. Our results suggest that the treatment combining road closure, soil decompaction, and tree planting could be beneficial to caribou, highlighting the relevance of including active restoration efforts in caribou conservation programs. We recommend that such a treatment be added to road decommissioning protocols for the conservation of caribou, alongside broad-scale habitat protection.     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

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 ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 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.     

Feeding ecology and trophic segregation of two sympatric mesopredatory sharks in the heavily exploited coastal ecosystem of the Adriatic Sea

The ecological roles and trophic interactions of two commercially important mesopredatory shark species, Squalus acanthias and Mustelus punctulatus that co‐occur on the continental shelf of the north‐central Adriatic Sea were investigated. Both shark species are dietary specialists, with a significant dietary overlap recorded only during the spring season. They showed different patterns of feeding as they grew: S. acanthias extended its trophic niche with an increase in size, while M. punctulatus developed a more specialized diet. These two sharks partition food resources and reduce niche overlap by foraging at different trophic levels. Mustelus punctulatus is a crustacean feeder, specialized in foraging on scavenging malacostracans frequently found along trawl tracks or on discards in the Adriatic fishing zone. Conversely, S. acanthias prefers small pelagic fishes, which are commercially exploited and in decline. The different foraging strategies adopted by these two species suggest that they should be managed separately. Dietary specialization, direct competition with humans for prey and their higher intrinsic vulnerability make S. acanthias particularly susceptible to the effects of anthropogenic perturbations.     

Potential conflict between future development of natural resources and high-value wildlife habitats in boreal landscapes

We used the Muskwa-Kechika Management Area in northeast British Columbia, Canada as a case study to determine potential conflicts between future resource development and high-value habitats of large mammals in an undeveloped boreal landscape. More than 50 % of high-value habitats for caribou, moose, elk, wolves and grizzly bears were located in Special Resource Management Zones, where natural resource developments could occur. We developed geographic information system (GIS) layers of potential forest resources, oil and gas, minerals, wind power, all resources combined, and roads; and quantified the proportions of high-value habitats overlapping these potentials. Greater proportions of high-value habitats across seasons for moose, elk, and wolves overlapped areas with high cumulative resource potential (winter, 49–70 %, growing season, 35–63 %) more than for three other species (grizzly bears, Stone’s sheep, mountain goats). This pattern was similar for forest resources, oil and gas, wind power, and roads. Caribou were more seasonally influenced. The proportions of their high-value habitat in areas with high cumulative resource potential (winter, 53 %, growing season, 16 %), as well as high forest and oil and gas potentials, were greatest in winter; in contrast, overlap with high mineral potential was greatest during the growing season. We recommend a quantitative and visual GIS approach to scenario planning in the Muskwa-Kechika to maintain the abundance and diversity of wildlife populations there. Resource development would likely increase early seral habitats, presumably benefiting moose, elk, and wolves, but could adversely affect caribou and grizzly bears through habitat loss and increased access.     

Molecular identification of Taenia spp. in wolves (Canis lupus), brown bears (Ursus arctos) and cervids from North Europe and Alaska

Taenia tapeworms of Finnish and Swedish wolves (Canis lupus) and Finnish brown bears (Ursus arctos), and muscle cysticerci of Svalbard reindeer (Rangifer tarandus platyrhynchus), Alaskan Grant's caribou (Rangifer tarandus granti) and Alaskan moose (Alces americanus) were identified on the basis of the nucleotide sequence of a 396 bp region of the mitochondrial cytochrome c oxidase subunit 1 gene. Two species were found from wolves: Taenia hydatigena and Taenia krabbei. The cysticerci of reindeer, caribou and one moose also represented T. krabbei. Most of the cysticercal specimens from Alaskan moose, however, belonged to an unknown T. krabbei-like species, which had been reported previously from Eurasian elks (Alces alces) from Finland. Strobilate stages from two bears belonged to this species as well. The present results suggest that this novel Taenia sp. has a Holarctic distribution and uses Alces spp. as intermediate and ursids as final hosts.