Logging, linear features, and human infrastructure shape the spatial dynamics of wolf predation on an ungulate neonate.

Humans are increasingly recognized as important players in predator-prey dynamics by modifying landscapes. This trend has been well-documented for large mammal communities in North American boreal forests: logging creates early seral forests that benefit ungulates such as white-tailed deer (Odocoileus virginianus), while the combination of infrastructure development and resource extraction practices generate linear features that allow predators such as wolves (Canis lupus) to travel and forage more efficiently throughout the landscape. Disturbances from recreational activities and residential development are other major sources of human activity in boreal ecosystems that may further alter wolf-ungulate dynamics. Here, we evaluate the influence that several major types of anthropogenic landscape modifications (timber harvest, linear features, and residential infrastructure) have on where and how wolves hunt ungulate neonates in a southern boreal forest ecosystem in Minnesota, USA. We demonstrate that each major anthropogenic disturbance significantly influences wolf predation of white-tailed deer fawns (n = 427 kill sites). In contrast with the "human shield hypothesis" that posits prey use human-modified areas as refuge, wolves killed fawns closer to residential buildings than expected based on spatial availability. Fawns were also killed within recently-logged areas more than expected. Concealment cover was higher at kill sites than random sites, suggesting wolves use senses other than vision, probably olfaction, to detect hidden fawns. Wolves showed strong selection for hunting along linear features, and kill sites were also closer to linear features than expected. We hypothesize that linear features facilitated wolf predation on fawns by allowing wolves to travel efficiently among high-quality prey patches (recently logged areas, near buildings), and also increase encounter rates with olfactory cues that allow them to detect hidden fawns. These findings provide novel insight into the strategies predators use to hunt ungulate neonates and the many ways human activity alters wolf-ungulate neonate predator-prey dynamics, which have remained elusive due to the challenges of locating sites where predators kill small prey. Our research has important management and conservation implications for wolf-ungulate systems subjected to anthropogenic pressures, particularly as the range of overlap between wolves and deer expands and appears to be altering food web dynamics in boreal ecosystems.

The ethology of wolves foraging on freshwater fish in a boreal ecosystem.

Through global positioning system (GPS) collar locations, remote cameras, field observations and the first wild wolf to be GPS-collared with a camera collar, we describe when, where and how wolves fish in a freshwater ecosystem. From 2017 to 2021, we recorded more than 10 wolves (Canis lupus) hunting fish during the spring spawning season in northern Minnesota, USA. Wolves ambushed fish in creeks at night when spawning fish were abundant, available and vulnerable in shallow waters. We observed wolves specifically targeting sections of rivers below beaver (Castor canadensis) dams, suggesting that beavers may indirectly facilitate wolf fishing behaviour. Wolves also cached fish on shorelines. We documented these findings across five different social groups at four distinct waterways, suggesting that wolf fishing behaviour may be widespread in similar ecosystems but has probably remained difficult to study given its annual brevity. Spawning fish may serve as a valuable pulsed resource for packs because the spring spawning season coincides with low primary prey (deer Odocoileus virginianus) availability and abundance, and when packs have higher energetic demands owing to newly born pups. We demonstrate the flexibility and adaptability of wolf hunting and foraging behaviour, and provide insight into how wolves can survive in a myriad of ecosystems.

A simple, low-cost method to age mammals? An alternative to cementum annuli analysis.

One of the most common and ubiquitous methods to age mammals is by counting the cementum annuli in molars, premolars, incisors, or canines. Despite the ubiquity and perceived simplicity of the method, cementum annuli analysis can be time-consuming, expensive, inaccurate, and imprecise, and require specialized equipment. Using beavers (Castor canadensis) as a test species, we developed a straightforward method to age mammals that requires little specialized equipment. The method consists of: (1) digitizing longitudinally sectioned teeth and measuring the proportion of tooth surface area comprised of cementum ("proportion cementum"), (2) evaluating the relationship between proportion cementum and specimen age (determined from either known-age samples or cementum annuli analysis), and (3) using the modeled relationship to estimate the age of other individuals based solely on proportion cementum. The relationship between proportion cementum and age was strongly correlated (R 2 = .97-.98 depending on observer), similar between observers, and similar between known-age specimens and those aged via cementum annuli analysis. Using this proportion cementum method, two independent observers accurately predicted the age of 80%-84% of specimens within 0.5 year and 96%-98% within 1 year. We suggest this aging method will likely work with most mammal species given the relatively consistent deposition of cementum throughout mammals' lives and has promise to be a simple and quick alternative to cementum annuli analysis regardless of whether one develops proportion cementum models using known-age specimens or those aged via alternative methods.

Statistical population reconstruction of moose (Alces alces) in northeastern Minnesota using integrated population models.

Given recent and abrupt declines in the abundance of moose (Alces alces) throughout parts of Minnesota and elsewhere in North America, accurately estimating statewide population trends and demographic parameters is a high priority for their continued management and conservation. Statistical population reconstruction using integrated population models provides a flexible framework for combining information from multiple studies to produce robust estimates of population abundance, recruitment, and survival. We used this framework to combine aerial survey data and survival data from telemetry studies to recreate trends and demographics of moose in northeastern Minnesota, USA, from 2005 to 2020. Statistical population reconstruction confirmed the sharp decline in abundance from an estimated 7,841 (90% CI = 6,702-8,933) in 2009 to 3,386 (90% CI = 2,681-4,243) animals in 2013, but also indicated that abundance has remained relatively stable since then, except for a slight decline to 3,163 (90% CI = 2,403-3,718) in 2020. Subsequent stochastic projection of the population from 2021 to 2030 suggests that this modest decline will continue for the next 10 years. Both annual adult survival and per-capita recruitment (number of calves that survived to 1 year per adult female alive during the previous year) decreased substantially in years 2005 and 2019, from 0.902 (SE = 0.043) to 0.689 (SE = 0.061) and from 0.386 (SE = 0.030) to 0.303 (SE = 0.051), respectively. Sensitivity analysis revealed that moose abundance was more sensitive to fluctuations in adult survival than recruitment; thus, we conclude that the steep decline in 2013 was driven primarily by decreasing adult survival. Our analysis demonstrates the potential utility of using statistical population reconstruction to monitor moose population trends and to identify population declines more quickly. Future studies should focus on providing better estimates of per-capita recruitment, using pregnancy rates and calf survival, which can then be incorporated into reconstruction models to help improve estimates of population change through time.

BASELINE PHYSIOLOGIC AND HEMATOLOGIC HEALTH IN WILD-CAUGHT MUSKRATS (ONDATRA ZIBETHICUS) FROM A NEAR-PRISTINE ECOSYSTEM IN NORTHERN MINNESOTA.

Muskrat (Ondatra zibethicus) populations show long-term and widespread declines across North America, necessitating research into potential mechanistic explanations, including population health. Previous research established reference hematology values, a proxy of individual health, of muskrats occurring in highly modified ecosystems. However, our knowledge of hematology metrics in muskrat populations occurring in more natural ecosystems is limited. We measured several hematological parameters of wild-caught muskrats (n = 73) in the Greater Voyageurs Ecosystem in northern Minnesota in 2018-2019 to establish baseline muskrat health in a relatively intact, near-pristine ecosystem. Additionally, we measured rectal temperature and heart and respiratory rates and collected whole blood for complete blood cell count assessment. We established baseline physiologic and hematologic reference ranges for the population and describe variations between total white blood cells, nucleated cell differentials, and basic erythron and platelet estimates and demonstrate methods of estimation to be poor proxies for more standardized counting methods. Our results establish a baseline to compare muskrat health assessments for populations affected by landscape change or in decline.

Ecological forecasts reveal limitations of common model selection methods: predicting changes in beaver colony densities.

Over the past two decades, there have been numerous calls to make ecology a more predictive science through direct empirical assessments of ecological models and predictions. While the widespread use of model selection using information criteria has pushed ecology toward placing a higher emphasis on prediction, few attempts have been made to validate the ability of information criteria to correctly identify the most parsimonious model with the greatest predictive accuracy. Here, we used an ecological forecasting framework to test the ability of information criteria to accurately predict the relative contribution of density dependence and density-independent factors (forage availability, harvest, weather, wolf [Canis lupus] density) on inter-annual fluctuations in beaver (Castor canadensis) colony densities. We modeled changes in colony densities using a discrete-time Gompertz model, and assessed the performance of four models using information criteria values: density-independent models with (1) and without (2) environmental covariates; and density-dependent models with (3) and without (4) environmental covariates. We then evaluated the forecasting accuracy of each model by withholding the final one-third of observations from each population and compared observed vs. predicted densities. Information criteria and our forecasting accuracy metrics both provided strong evidence of compensatory density dependence in the annual dynamics of beaver colony densities. However, despite strong within-sample performance by the most complex model (density-dependent with covariates) as determined using information criteria, hindcasts of colony densities revealed that the much simpler density-dependent model without covariates performed nearly as well predicting out-of-sample colony densities. The hindcast results indicated that the complex model over-fit our data, suggesting that parameters identified by information criteria as important predictor variables are only marginally valuable for predicting landscape-scale beaver colony dynamics. Our study demonstrates the importance of evaluating ecological models and predictions with long-term data and revealed how a known limitation of information criteria (over-fitting of complex models) can affect our interpretation of ecological dynamics. While incorporating knowledge of the factors that influence animal population dynamics can improve population forecasts, we suggest that comparing forecast performance metrics can likewise improve our knowledge of the factors driving population dynamics.

Outsized effect of predation: Wolves alter wetland creation and recolonization by killing ecosystem engineers.

Gray wolves are a premier example of how predators can transform ecosystems through trophic cascades. However, whether wolves change ecosystems as drastically as previously suggested has been increasingly questioned. We demonstrate how wolves alter wetland creation and recolonization by killing dispersing beavers. Beavers are ecosystem engineers that generate most wetland creation throughout boreal ecosystems. By studying beaver pond creation and recolonization patterns coupled with wolf predation on beavers, we determined that 84% of newly created and recolonized beaver ponds remained occupied until the fall, whereas 0% of newly created and recolonized ponds remained active after a wolf killed the dispersing beaver that colonized that pond. By affecting where and when beavers engineer ecosystems, wolves alter all of the ecological processes (e.g., water storage, nutrient cycling, and forest succession) that occur due to beaver-created impoundments. Our study demonstrates how predators have an outsized effect on ecosystems when they kill ecosystem engineers.

Landscape structure and population density affect intraspecific aggression in beavers.

Intraspecific competition plays an important role for territory acquisition and occupancy, in turn affecting individual fitness. Thus, understanding the drivers of intraspecific aggression can increase our understanding of population dynamics. Here, we investigated intraspecific aggression in Eurasian (Castor fiber) and North American (Castor canadensis) beavers that are both monogamous, territorial mammals. Combined, we examined tail scars from >1,000 beavers (>2,000 capture events) as part of two long-term studies in Norway and the USA. We investigated the influence of landscape structure, population density, sex, age, and (for Eurasian beavers only) social status and group size on the number of tail scars caused by conspecifics. The number of tail scars was affected by population density in well-connected landscape types (large lakes and rivers), but not in more isolated areas (ponds), where individuals generally had fewer tail scars. Further, the relationship of population density was not linear. In the North American beaver population occurring in large lakes, intraspecific aggression increased with population density. Conversely, in the saturated Eurasian beaver population, intraspecific aggression was in a negative relationship with population density (except at the highest densities), likely due to inverse density-dependent intruder pressure via dispersers. Our findings emphasize that population density can affect intraspecific aggression depending on landscape structure, which might have important consequences for local patterns of dispersal, mate change, and territory occupancy, all of which can affect population dynamics.

Top-down effects of repatriating bald eagles hinder jointly recovering competitors.

The recovery of piscivorous birds around the world is touted as one of the great conservation successes of the 21st century, but for some species, this success was short-lived. Bald eagles, ospreys and great blue herons began repatriating Voyageurs National Park, USA, in the mid-20th century. However, after 1990, only eagles continued their recovery, while osprey and heron recovery failed for unknown reasons. We aimed to evaluate whether top-down effects of bald eagles and bottom-up effects of inclement weather, habitat quality and fish resources contributed to the failed recovery of ospreys and herons in a protected area. We quantified the relative influence of top-down and bottom-up factors on nest colonization, persistence (i.e., nest reuse) and success for ospreys, and occurrence and size of heronries using 26 years (1986-2012) of spatially explicit monitoring data coupled with multi-response hierarchical models and Bayesian variable selection approaches. Bald eagles were previously shown to recover faster due to intensive nest protection and management. Increased numbers of eagles were associated with a reduction in the numbers of osprey nests, their nesting success and heronry size, while higher local densities of nesting eagles deterred heronries nearby. We found little evidence of bottom-up limitations on the failed recovery of herons and ospreys. We present a conservation conundrum: bald eagles are top predators and a flagship species of conservation that have benefited from intensive protection, but this likely hindered the recovery of ospreys and herons. Returning top predators, or rewilding, is widely promoted as a conservation strategy for top-down ecosystem recovery, but managing top predators in isolation of jointly recovering species can halt or reverse ecosystem recovery. Previous studies warn of the potential consequences of ignoring biotic interactions amongst recovering species, but we go further by quantifying how these interactions contributed to failed recoveries via impacts on the nesting demography of jointly recovering species. Multi-species management is paramount to realizing the ecosystem benefits of top predator recovery.

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.

Finding wolf homesites: improving the efficacy of howl surveys to study wolves.

Locating wolf (Canis lupus) homesites is valuable for understanding the foraging behavior, population dynamics, and reproductive ecology of wolves during summer. During this period wolf pack members (adults and pups) readily respond to simulated wolf howls (i.e., howl surveys), which allows researchers to estimate the location of the homesite via triangulation. Confirming the actual locations of homesites via ground truthing is labor intensive because of the error surrounding estimated locations. Our objectives were (1) to quantify observer error during howl surveys and compare amongst experience levels, (2) provide a simple method for locating homesites in the field by incorporating observer error, and (3) further document the value of this method for monitoring wolf packs throughout the summer. We located 17 homesites by howl surveys during 2015-2017 in the Greater Voyageurs Ecosystem, Minnesota, USA. Of 62 bearings taken by observers during howl surveys, bearings erred by an average of 7.6° ± 6.3° (SD). There was no difference in observer error between novice and experienced observers. A simple way to increase efficiency when searching for homesites is to search concentric areas (bands) based on estimated observer error, specifically by: (1) adding ±10° error bands around howl survey bearings when ≥3 bearings can be obtained, (2) ±10° and ±20° error bands when 2 bearings are obtained, and (3) ±10° and ±26° error bands when 1 bearing is obtained. By incorporating observer error and understanding how frequently and how far wolves move homesites, it is possible to monitor wolf packs and confirm most, if not all, homesites used by a pack from at least June until August without having a collared individual in a pack.

Moose at their bioclimatic edge alter their behavior based on weather, landscape, and predators.

Populations inhabiting the bioclimatic edges of a species' geographic range face an increasing amount of stress from alterations to their environment associated with climate change. Moose Alces alces are large-bodied ungulates that are sensitive to heat stress and have exhibited population declines and range contractions along their southern geographic extent. Using a hidden Markov model to analyze movement and accelerometer data, we assigned behaviors (rest, forage, or travel) to all locations of global positioning system-collared moose (n = 13, moose-years = 19) living near the southern edge of the species' range in and around Voyageurs National Park, MN, USA. We assessed how moose behavior changed relative to weather, landscape, and the presence of predators. Moose significantly reduced travel and increased resting behaviors at ambient temperatures as low as 15 °C and 24 °C during the spring and summer, respectively. In general, moose behavior changed seasonally in association with distance to lakes and ponds. Moose used wetlands for travel throughout the year, rested in conifer forests, and foraged in shrublands. The influence of wolves Canis lupus varied among individual moose and season, but the largest influence was a reduction in travel during spring when near a wolf home range core, primarily by pregnant females. Our analysis goes beyond habitat selection to capture how moose alter their activities based on their environment. Our findings, along with climate change forecasts, suggest that moose in this area will be required to further alter their activity patterns and space use in order to find sufficient forage and avoid heat stress.

Erratum: Moose at their bioclimatic edge alter their behavior based on weather, landscape, and predators.

[This corrects the article DOI: 10.1093/cz/zox047.]

Lambdapapillomavirus 2 in a Gray Wolf ( Canis lupus ) from Minnesota, USA with Oral Papillomatosis and Sarcoptic Mange.

Oral papillomatosis was diagnosed in a gray wolf ( Canis lupus ) with sarcoptic mange from Minnesota, US found dead in February 2015. Intranuclear inclusion bodies were evident histologically, and papillomaviral antigens were confirmed using immunohistochemistry. Sequencing of the L1 papillomavirus gene showed closest similarity to Lambdapapillomavirus 2.

A SEROSURVEY OF DISEASES OF FREE-RANGING GRAY WOLVES (CANIS LUPUS) IN MINNESOTA, USA.

We tested serum samples from 387 free-ranging wolves ( Canis lupus ) from 2007 to 2013 for exposure to eight canid pathogens to establish baseline data on disease prevalence and spatial distribution in Minnesota's wolf population. We found high exposure to canine adenoviruses 1 and 2 (88% adults, 45% pups), canine parvovirus (82% adults, 24% pups), and Lyme disease (76% adults, 39% pups). Sixty-six percent of adults and 36% of pups exhibited exposure to the protozoan parasite Neospora caninum . Exposure to arboviruses was confirmed, including West Nile virus (37% adults, 18% pups) and eastern equine encephalitis (3% adults). Exposure rates were lower for canine distemper (19% adults, 5% pups) and heartworm (7% adults, 3% pups). Significant spatial trends were observed in wolves exposed to canine parvovirus and Lyme disease. Serologic data do not confirm clinical disease, but better understanding of disease ecology of wolves can provide valuable insight into wildlife population dynamics and improve management of these species.

Where and How Wolves (Canis lupus) Kill Beavers (Castor canadensis).

Beavers (Castor canadensis) can be a significant prey item for wolves (Canis lupus) in boreal ecosystems due to their abundance and vulnerability on land. How wolves hunt beavers in these systems is largely unknown, however, because observing predation is challenging. We inferred how wolves hunt beavers by identifying kill sites using clusters of locations from GPS-collared wolves in Voyageurs National Park, Minnesota. We identified 22 sites where wolves from 4 different packs killed beavers. We classified these kill sites into 8 categories based on the beaver-habitat type near which each kill occurred. Seasonal variation existed in types of kill sites as 7 of 12 (58%) kills in the spring occurred at sites below dams and on shorelines, and 8 of 10 (80%) kills in the fall occurred near feeding trails and canals. From these kill sites we deduced that the typical hunting strategy has 3 components: 1) waiting near areas of high beaver use (e.g., feeding trails) until a beaver comes near shore or ashore, 2) using vegetation, the dam, or other habitat features for concealment, and 3) immediately attacking the beaver, or ambushing the beaver by cutting off access to water. By identifying kill sites and inferring hunting behavior we have provided the most complete description available of how and where wolves hunt and kill beavers.

Mercury Concentrations in Eggs of Red-Winged Blackbirds and Tree Swallows Breeding in Voyageurs National Park, Minnesota.

Most investigations of the environmental effects of mercury (Hg) have focused on aquatic food webs that include piscivorous fish or wildlife. However, recent investigations have shown that other species, including passerine songbirds, may also be at risk from exposure to methylmercury (MeHg). We quantified Hg concentrations in eggs of two species of songbirds, red-winged blackbirds (Agelaius phoeniceus) and tree swallows (Tachycineta bicolor), nesting in Voyageurs National Park, Minnesota, USA. Geometric mean concentrations of total Hg (THg) were lower in red-winged blackbird eggs [218 and 107 ng/g dry weight (dw) for 2012 and 2013, respectively] than in tree swallow eggs (228 and 300 ng/g dw for 2012 and 2013, respectively), presumably reflecting differences in the trophic positions of these two species. Concentrations of MeHg averaged 98.4 % of THg in red-winged blackbird eggs. Levels of THg observed in this study were well below critical toxicological benchmarks commonly applied to eggs of avian species, suggesting these breeding populations were not adversely affected by exposure to MeHg. In red-winged blackbirds, concentrations of THg in eggs collected in 2012 were twice those in eggs collected in 2013. Hg levels in eggs of both species increased with date of clutch initiation. In red-winged blackbirds, for example, temporal patterns showed that a 3-week delay in clutch initiation increased egg THg by 60 %. These observations indicate that in ovo exposure of wetland birds to MeHg can vary significantly within nesting season as well as between years.

Landscape influence on spatial patterns of meningeal worm and liver fluke infection in white-tailed deer.

Parasites that primarily infect white-tailed deer (Odocoileus virginianus), such as liver flukes (Fascioloides magna) and meningeal worm (Parelaphostrongylus tenuis), can cause morbidity and mortality when incidentally infecting moose (Alces alces). Ecological factors are expected to influence spatial variation in infection risk by affecting the survival of free-living life stages outside the host and the abundance of intermediate gastropod hosts. Here, we investigate how ecology influenced the fine-scale distribution of these parasites in deer in Voyageurs National Park, Minnesota. Deer pellet groups (N = 295) were sampled for the presence of P. tenuis larvae and F. magna eggs. We found that deer were significantly more likely to be infected with P. tenuis in habitats with less upland deciduous forest and more upland mixed conifer forest and shrub, a pattern that mirrored microhabitat differences in gastropod abundances. Deer were also more likely to be infected with F. magna in areas with more marshland, specifically rooted-floating aquatic marshes (RFAMs). The environment played a larger role than deer density in determining spatial patterns of infection for both parasites, highlighting the importance of considering ecological factors on all stages of a parasite's life cycle in order to understand its occurrence within the definitive host.