Maintaining subsistence species on hunting lands is essential to the food security and cultural preservation/flourishing of Indigenous peoples that rely on traditional foods. In northern North America, moose play a central role in subsistence, cultural, and stewardship practices but are declining in many parts of their range. Moose (Alces alces) in Minnesota are a threatened population that is integral to the lifeways of the Lake Superior Chippewa. This study, led by the Grand Portage Band, examines the shifting causes of adult moose mortality between 2010 and 2022 on the Grand Portage Indian Reservation and Voyageurs National Park. These efforts, rooted in principles of Indigenous sovereignty and co-stewardship, seek to sustain this vital species on ancestral lands. We observed that the relative importance of mortality causes varied over time, with Parelaphostrongylus tenuis and other health-related factors driving mortality during the initial decline period (2010-2014), while predation became a leading cause of mortality and quadrupled in probability during the stabilization period (2015-2022). Using a Bayesian framework, we integrated multiple contributing factors to accurately estimate cause-specific mortality probabilities and survival rates. The findings underscore the necessity for adaptive management strategies that address both parasitism and predation pressures to recover moose populations to pre-decline levels. Moreover, this study exemplifies how a long-term, Indigenous-led wildlife collaring and monitoring program is critical to capturing these dynamics and supporting the Grand Portage Band's ongoing stewardship. This research advances our understanding of moose mortality in a vulnerable population and reinforces the importance of Indigenous leadership in wildlife management and scientific co-production. By integrating traditional ecological knowledge found in Tribal-governmental planning documents with contemporary science led by their Natural Resources Management department, the Grand Portage Band is ensuring that moose remain a resilient and enduring part of their cultural and subsistence practices, thus contributing to the broader framework of Indigenous co-stewardship.
Despite changes in legislation and public attitudes towards large predators, human-caused mortality continues to impact survival and conservation of carnivore species. Understanding the drivers and timing of mortality is critical for informing evidence-based management and policy decisions aimed at improving carnivore conservation outcomes, particularly in areas with varying management regimes. Using GPS collar and mortality data representing 1,002 wolf-years from 608 wolves in Minnesota, Wisconsin, and Michigan USA during 2010–2023, we assessed cause-specific mortality and survival in relation to federal protection status, whether wolf harvest was allowed, sex-age class, and wolf residency. Human-caused mortalities represented 65% of recorded mortalities. Illegal kill represented 38% of mortalities and peaked in mid-November, concurrent with white-tailed deer firearm seasons in the region. Whether legal harvest was allowed did not influence mortality risk for causes other than legal kill. Mean annual regional wolf survival (0.74) was similar across years and sex-age categories and did not vary with protection status, but survival probability for resident wolves was greater than for non-residents. Legal kill was the greatest source of mortality for gray wolves in the western Great Lakes region in years when harvest was allowed, emphasizing the importance of continued population monitoring and adaptive management. Illegal kill was the greatest source of recorded mortality overall and was not reduced by federal protection, highlighting the need to enhance enforcement and address socio-political factors influencing public tolerance of wolves.
The moose (Alces alces; mooz in Anishinaabemowin, Ojibwe language) population has recently declined in Minnesota, USA, and gray wolf (Canis lupus; ma'iingan) predation is likely a contributing factor. We analyzed diet composition of gray wolves in northeastern Minnesota during 2011-2013 to evaluate the importance of moose as prey and seasonal and regional variations in wolf diet. We identified frequency of occurrence of prey items and biomass consumed in 1,000 wolf scats collected on and adjacent to the Grand Portage Indian Reservation and Voyageurs National Park and within the 1854 Ceded Territory (greater northeastern Minnesota). White-tailed deer (Odocoileus virginianus; waawaashkeshiwag [plural]), moose, and beaver (Castor canadensis; amikwag [plural]) composed the majority of wolf diet, with moose as the primary prey in Grand Portage (35-54% of diet by biomass across seasons) and deer the primary prey in the 1854 Ceded Territory (46-62%) and Voyageurs (63-79%). Relative importance of prey species differed by study area and season. Moose calves were an important prey item in spring in the 1854 Ceded Territory (12% of diet by biomass) but not in Grand Portage or Voyageurs. Although calves were not a majority of wolf diet by biomass, many calves were preyed upon by wolves (30% of calves born each year in Grand Portage), thus affecting recruitment in a declining moose population. Deer fawns composed 12% of wolf diet in spring and 10% in summer in Grand Portage and 19% in summer in Voyageurs. Beaver composed 16% of wolf diet by biomass in spring and 14% in summer in Grand Portage and composed 22% of wolf diet in spring and 30% in summer in Voyageurs. At most prey densities, moose were preferred and deer avoided in Grand Portage and the 1854 Ceded Territory and beaver were preferred in Voyageurs. Our results can be used in conjunction with predation and prey studies to evaluate the effect of wolves on prey populations.
Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission-fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids (Canis spp.) varied widely in the proportion of time they spent together (5%-100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.
Cooperative hunting can confer fitness benefits by increasing foraging efficiency. We documented a breeding pair of wolves in the Greater Voyageurs Ecosystem of Minnesota, USA that appeared to periodically use cooperative ambushing to hunt beavers. The breeding pair primarily chose to wait-in-ambush close to one another (< 65 m) but on different beaver feeding trails, which appears advantageous because: (1) feeding trails are where beavers are most active and vulnerable on land, (2) the probability that the pair encounters a beaver is increased, and (3) either wolf can quickly assist the other in killing a beaver. The cooperative ambush strategy these wolves used appears rare for most social Carnivora but we hypothesize this behavior is widespread in areas of wolf-beaver sympatry. This observation demonstrates that novel insights into the natural history of even well-studied predators are possible when technological advancements are combined with intensive fieldwork.
Appropriately monitoring vulnerable species is essential for resource management decision-making. We used two different metrics (occupancy and acoustic activity) to evaluate population trends of North American bats susceptible to a fungal disease and compared the outcomes of these two analyses. Our dataset consisted of passive acoustic survey data collected at nine U.S. national parks in the Great Lakes region between 2016 and 2020. Our focal species were little brown bat (Myotis lucifugus), northern long-eared bat (Myotis septentrionalis), tricolored bat (Perimyotis subflavus), and big brown bat (Eptesicus fuscus). Combining acoustic detection data with sitespecific and environmental covariates, we developed park- and species-specific models of occupancy (use) and acoustic activity. For little brown, northern long-eared, and tricolored bats, probability of use declined 20 % and activity declined 66 %, on average. For big brown bat, probability of use decreased 9 % and activity increased 17 %. Our results showed that use of space and level of activity were not predicted by the same covariates but were positively related. We also found probability of use could remain high while activity decreased to relatively low levels. Although we observed precipitous declines in activity rates of three species, they were still using a high proportion of the landscape. We conclude that analyzing activity provides different but complementary information to analyzing species' use of space. The two metrics vary on different temporal and spatial scales and have different biases. Appropriate choice of which parameter to analyze is crucial, as different parameters may lead to different ecological conclusions, thus affecting how management decisions are made and how species conservation and recovery efforts are implemented.
Transboundary movement of wildlife results in some of the most complicated and unresolved wildlife management issues across the globe. Depending on the location and managing agency, gray wolf ( Canis lupus ) management in the US ranges from preservation to limited hunting to population reduction. Most wildlife studies focus on population size and growth rate to inform management, but relatively few examine species biological processes at scales aside from that of the population. This is especially important for group-living species such as the gray wolf, for which the breeding unit is the social group. We analyzed data for gray wolf packs living primarily within several US National Park Service units (years of data): Denali National Park and Preserve (33 years), Grand Teton National Park (23 years), Voyageurs National Park (12 years), Yellowstone National Park (27 years), and Yukon-Charley Rivers National Preserve (23 years). We identified two gray wolf biological processes that differed from population size – namely, pack persistence and reproduction – and determined that while human-caused mortality had negative effects on both, pack size had a moderating effect on the impacts of mortality.
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.
Recovering species are not returning to the same environments or communities from which they disappeared. Conservation researchers and practitioners are thus faced with additional challenges in ensuring species resilience in these rapidly changing ecosystems. Assessing the resilience of species in these novel systems can still be guided by species’ ecology, including knowledge of their population size, life history traits, and behavioral adaptations, as well as the type, strength, and number of ways that they interact with other species in the community. We summarized broad trends of Common Loons (Gavia immer) breeding at Voyageurs National Park from 1973 to 2009, and evaluated the effects of increased risk from recovering Bald Eagles (Haliaeetus leucocephalus) on chick survival from 2004 to 2006. Adult Common Loons appear to have increased over time. Using Bayesian survival models that accounted for imperfect detection of unmarked individuals, we determined that chick survival of Common Loons was high from year to year and was unrelated to predation risk from Bald Eagles because chicks in territories closer to active nests did not experience greater mortality than those farther away. We suggest that Common Loon chicks were unaffected by the recovery of this top predator during the three years of sampling. Previous research indicates that Bald Eagles and other predators are an important source of egg losses, but Common Loons can compensate by re-nesting. Despite current uncertainties from anthropogenic threats, knowledge of a species’ ecology remains instrumental in determining its resilience during recovery.
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.
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.
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.
Frontiers in Ecology and the EnvironmentVolume 20, Issue 5 p. 275-277 Write Back Predator personalities alter ecosystem services Joseph Bump, Corresponding Author Joseph Bump bump@umn.edu Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNEmail: bump@umn.eduSearch for more papers by this authorThomas Gable, Thomas Gable Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorSean Johnson-Bice, Sean Johnson-Bice University of Manitoba, Winnipeg, CanadaSearch for more papers by this authorAustin Homkes, Austin Homkes Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorDanielle Freund, Danielle Freund Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorSteve Windels, Steve Windels Voyageurs National Park, International Falls, MNSearch for more papers by this authorStotra Chakrabarti, Stotra Chakrabarti Macalester College, Saint Paul, MNSearch for more papers by this author Joseph Bump, Corresponding Author Joseph Bump bump@umn.edu Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNEmail: bump@umn.eduSearch for more papers by this authorThomas Gable, Thomas Gable Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorSean Johnson-Bice, Sean Johnson-Bice University of Manitoba, Winnipeg, CanadaSearch for more papers by this authorAustin Homkes, Austin Homkes Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorDanielle Freund, Danielle Freund Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Minneapolis, MNSearch for more papers by this authorSteve Windels, Steve Windels Voyageurs National Park, International Falls, MNSearch for more papers by this authorStotra Chakrabarti, Stotra Chakrabarti Macalester College, Saint Paul, MNSearch for more papers by this author First published: 01 June 2022 https://doi.org/10.1002/fee.2512Read the full textAboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat No abstract is available for this article. Volume20, Issue5June 2022Pages 275-277 RelatedInformation
Like many ecological processes, natural disturbances exhibit scale‐dependent dynamics that are largely a function of the magnitude, frequency and scale at which they are assessed. Ecosystem engineers create patch‐scale disturbances that affect ecological processes, yet we know little about how these effects scale across space or vary through time. Here, we investigate how patch disturbances by beavers Castor canadensis, ecosystem engineers renowned for their pond‐creation behavior, affect ecological processes across space and time. We evaluated how beaver population recovery influenced surface water dynamics in relation to population density over 70 years across multiple spatial scales (pond, watershed and regional) in northern Minnesota. Surface water area was positively related to population density at the watershed scale; however, despite variation in beaver densities (and therefore surface water area) at the watershed scale, regional‐scale surface water area was stable through time. This stability appears to have been driven by asynchronous beaver density fluctuations among watersheds, combined with the increasing importance of abandoned ponds. Beavers initially created and occupied larger ponds with greater surface water area, but through time shifted towards occupying smaller ponds. As ponds accumulated on the landscape proportionally more surface water was stored within abandoned ponds, which offset the smaller size of occupied ponds. Beaver engineering – driven by density‐dependent mechanisms and the legacy effects from abandoned ponds – not only follows general patterns of patch disturbance dynamics by creating a spatial mosaic of patches, but the organism‐created mosaic also appears to generate ecological stability at greater spatial scales. We suggest restoring beavers to landscapes is a viable method for increasing surface water storage and will ultimately help advance numerous conservation and rewilding objectives. Our study demonstrates that ecosystem engineering effects can be scale‐dependent, indicating researchers should evaluate the ecological impact of engineers across diverse spatiotemporal scales to fully understand their functional roles in ecosystems.
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.
Comprehensive knowledge of ambush behavior requires an understanding of where a predator expects prey to be, which is generally unknowable because ambush predators often hunt mobile prey that exhibit complex, irregular, or inconspicuous movements. Wolves (Canis lupus) are primarily cursorial predators, but they use ambush strategies to hunt beavers (Castor canadensis). Terrestrial beaver activity is predictable because beavers use well-defined, conspicuous habitat features repeatedly. Thus, studying where wolves wait-in-ambush for beavers provides a unique opportunity to understand how predators choose ambush locations in relation to prey activity. We searched 11 817 clusters of GPS locations from wolves in the Greater Voyageurs Ecosystem, International Falls, MN, and documented 748 ambushing sites and 214 instances where wolves killed beavers. Wolves chose ambush locations: 1) with olfactory concealment to avoid detection from the highly developed olfactory senses of beavers and 2) close (generally <5 m) to beaver habitat features to take advantage of beavers’ inability to visually detect motionless predators. Our work describes in detail the ambush strategies wolves use to hunt beavers and continues to overturn the traditional notion that wolves rely solely on cursorial hunting strategies. We also demonstrate that ambush predators can anticipate the movements and behavior of their prey due to a fundamental understanding of their prey’s sensory abilities. Wolves, therefore, and likely ambush predators in general, appear capable of simultaneously accounting for abiotic and biotic factors when choosing ambush locations, ultimately allowing them to counter and capitalize on the sensory abilities of their prey.
Muskrats Ondatra zibethicus are semiaquatic herbivores experiencing long-term and widespread population declines across North America. Translocation may be a viable tool to bolster or reestablish local populations; however, subsequent effects of translocation on muskrats are unknown. We live-trapped and translocated radiomarked muskrats (n = 65) during the summers of 2018–2019 in Voyageurs National Park, MN, USA and assessed post-translocation effects on weekly survival probabilities and space-use patterns. We did not observe homing behavior, though individuals moved an average of 2.2 km (SE = 0.30 km) from release sites and established home ranges within ∼8 days (SE = 1.16 days) post-translocation. Weekly post-translocation survival probabilities (0.95, SE = 0.001) and average home-range sizes (2.52 ha, SE = 0.44 ha) were similar to other studies of non-translocated muskrats. Our most-supported known-fate survival model revealed muskrats using beaver Castor canadensis lodges had greater weekly survival probabilities. Additionally, weekly muskrat survival varied between years suggesting a positive response to a novel soft-release technique applied in 2019. Our study provides the first empirical assessment of translocation effects on muskrats and suggests translocation may be effective for establishing or enhancing local muskrat populations. Additionally, our study suggests beaver lodges may confer fitness benefits to sympatric muskrats particularly during dispersal.
Abstract 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.
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.
Context Interspecific competition can limit species distributions unless competitors partition niche space to enable coexistence. Landscape heterogeneity can facilitate niche partitioning and enable coexistence, but land-use change is restructuring terrestrial ecosystems globally with unknown consequences for species interactions. Objectives We tested the relationship between landscape heterogeneity and carnivore co-occurrence in natural and human-dominated ecosystems to assess the landscape-mediated impacts of anthropogenic change on coexistence. Methods We used boosted regression trees to model the distributions and co-occurrence of two competing forest carnivores, American martens and fishers, at two contrasting sites in the Great Lakes region, USA: a "natural" site largely devoid of human impacts and a "human-dominated" site with substantial development and a history of land-use change. We assessed the importance of climate and habitat variables for each species, measured spatial niche overlap, and quantified co-occurrence as a function of compositional (patch richness), configurational (landscape shape), and topographical (elevation range) heterogeneity per site. Results We observed significant differences in the effect of heterogeneity on co-occurrence between sites. The natural landscape exhibited little niche overlap and co-occurrence had a significant, positive relationship with heterogeneity. Conversely, the human-dominated site exhibited high niche overlap with variable effects of heterogeneity on co-occurrence. Elevation, snowpack, and development had strong, contrasting effects on marten and fisher distributions, suggesting that differential use of habitat and anthropogenic features facilitates coexistence. Conclusions Heterogeneity can facilitate coexistence, but too much may undermine carnivore coexistence in human-dominated landscapes where habitat and space are limited. Moreover, future climate change will likely erode niche partitioning among martens and fishers, with particularly strong consequences for coexistence in human-dominated landscapes and at range boundaries.