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Status and management of moose in the northeastern United States

Moose (Alces alces) populations have recolonized much of their historic range in the northeastern United States in the past 30 years, with their southern range edge extending to southern New England and northern New York. This southerly expansion occurred when certain other populations in the United States were in decline along the southern range edge, with climate change often cited as a probable cause. The areas that moose have recently occupied in the northeastern United States are some of the most densely human populated in moose range, which has raised concern about human safety and moose-vehicle collisions (MVC). We conducted a literature search about moose in the northeastern United States, and distributed a questionnaire and conducted phone interviews with regional biologists responsible for moose management to determine the status of moose, management activity, and research deficiencies and needs. Moose numbers appear stable throughout much of the region, with slow population growth in northern New York. Management activity ranges from regulated harvest of moose in Maine, New Hampshire, and Vermont, to no active management in southern New England and New York. The combined annual harvest in Maine, New Hampshire, and Vermont is >3,000. MVCs are a widespread regional concern with >1,000 occurring annually involving several human fatalities. Research should address impacts of parasitism by winter tick (Dermacentor albipictus) and brain-worm (Parelaphostrongylus tenuis) on productivity and mortality of moose, influence of climate change on population dynamics and range, and conflicts in areas with high human population density.

Alces

Space use and movements of moose in Massachusetts: implications for conservation of large mammals in a fragmented environment

Moose ( Alces alces ) have recently re-occupied a portion of their range in the temperate deciduous forest of the northeastern United States after a >200 year absence. In southern New England, moose encounter different forest types, more human development, and higher temperatures than in other parts of their geographic range in North America. We analyzed seasonal minimum convex polygon home ranges, utilization distributions, movement rates, and home range composition of GPS-collared moose in Massachusetts. Seasonal home range sizes were not different for males and females and were within the range reported for low latitudes elsewhere in North America. Seasonal movement patterns reflected the seasonal changes in metabolic rate and the influence of the species’ reproductive cycle and weather. Home ranges consisted almost entirely of forested habitat, included large amounts of conservation land, and had lower road densities as compared to the landscape as a whole, indicating that human development may be a limiting factor for moose in the region. The size and configuration of home ranges, seasonal movement patterns, and use relative to human development have implications for conservation of moose and other wide-ranging species in more highly developed portions of their ranges.

Massachusetts

Moose habitat in Massachusetts: Assessing use at the southern edge of the range

Moose (Alces alces) have recently re-occupied a portion of their range in the temperate deciduous forest of the northeastern United States after a more than 200 year absence. In southern New England, moose are exposed to a variety of forest types, increasing development, and higher ambient temperatures as compared to other parts of their geographic range. Additionally, large-scale disturbances that shape forest structure and expansive naturally occurring shrub-willow communities used commonly elsewhere are lacking. We used utilization distributions to determine third order habitat selection (selection within the home range) of GPS-collared moose. In central Massachusetts, forests regenerating from logging were the most heavily used cover type in all seasons (48 - 63% of core area use). Habitat use of moose in western Massachusetts varied more seasonally, with regenerating forests used most heavily in summer and fall (57 and 46%, respectively), conifer and mixed forests in winter (47 - 65%), and deciduous forests in spring (41%). This difference in habitat selection reflected the transition from northern forest types to more southern forest types across the state. The intensive use of patches of regenerating forest emphasizes the importance of sustainable forest harvesting to moose. This study provides the first assessment of habitat requirements in this southern portion of moose range and provides insights into re-establishment of moose in unoccupied portions of its historic range in New York and Pennsylvania.

Massachusetts, New Hampshire, Vermont

Modeling moose habitat use by age, sex, and season in Vermont, USA using high-resolution lidar and national land cover data

Moose ( Alces alces ) populations have experienced unprecedented declines along the southern periphery of their range, including Vermont, USA. Habitat management may be used to improve the status of the population and health of individuals. To date, however, Vermont wildlife managers have been challenged to effectively use this important tool due to the lack of fine-scale information on moose space use and habitat characteristics. To assess habitat use, we combined more than 40,000 moose locations collected from radio-collared individuals (n = 74), recent land cover data, and high resolution, 3-dimensional lidar ( light detection and ranging ) data to develop Resource Utilization Functions (RUF) by age (mature and young adult), season (dormant and growth), and sex. Each RUF linked home range use to average habitat conditions within 400 m or 1 km of each 30 m 2 pixel within the home range. Across analyses, the top RUF models included both composition (as measured through the National Land Cover Database) and structure (as measured through lidar) variables, and significantly outperformed models that excluded lidar variables. These findings support the notion that lidar is an effective tool for improving the ability of models to estimate patterns of habitat use, especially for larger bodied mammals. Generally speaking, female moose actively used areas with proportionally more regenerating forest (i.e., forage < 3.0 m) and more mature forest (i.e., canopy structure > 6.0 m), while males actively used more high elevation, mixed forest types. Further, moose exhibited important seasonal differences in habitat use that likely reflect temporal changes in energetic and nutritional requirements and behavior across the year. Moose used areas with proportionally more regenerating forest (i.e., forage < 3.0 m) during the growth period and female moose had strong positive associations with lidar-derived canopy structure during the growth (but not the dormant) period. Ultimately, the resultant maps of habitat use provide a means of informing management activities (e.g., the restoration or alteration of habitats to benefit moose) and policies around land use that may contribute to population recovery.

Vermont

Extending body condition scoring beyond measurable rump fat to estimate full range of nutritional condition for moose

Moose ( Alces alces ) populations along the southern extent of their range are largely declining, and there is growing evidence that nutritional condition — which influences several vital rates – is a contributing factor. Moose body condition can presently be estimated only when there is measurable subcutaneous rump fat, which equates to animals with >6% ingesta-free body fat (IFBFat). There is need for a technique to allow body fat estimation of animals in poorer body condition (i.e., <6% body fat). We advance current methods for moose, following those used and validated with other ungulate species, by establishing a moose-specific body condition score (BCS) that can be used to estimate IFBFat in the lower range of condition. Our modified BCS was related strongly ( r 2 = 0.89) to IFBFat estimates based on measurable rump fat. By extending the predicted relationship to individuals without measurable fat, the BCS equated severe emaciation with 0.67% IFBFat, supporting the accuracy of the method. The lower end of nutritional condition is important for identifying relationships involving life-history characteristics because most state-dependent changes occur at lower levels of condition. Therefore, until the BCS can be validated with moose carcasses, we believe our method to estimate body fat across the full range of condition should yield better understanding of the drivers underlying declining moose populations.

Alces

Birth rates and neonate survival in a parasite rich moose population in Vermont, USA

Moose ( Alces alces ) populations are declining across much of their southern geographic range in North America. In Vermont and other northeastern states, measurable declines are attributed to low calf survival and reduced productivity associated with persistent winter tick ( Dermacentor albipictus ) parasitism. In 2017–2020, we studied 75 radio-collared female moose (38 calves and 37 adults) in Vermont to examine physiological, spatial, and temporal parameters relative to calf survival and adult productivity. Physiological measures included concentration of fecal glucocorticoid metabolites (fGCM) which reflects stress, and urea nitrogen:creatinine ratios in urine (UN:C) which proxy nutritional state. The pregnancy rate at capture across years was 0.67 (95% CI = 0.50 – 0.80), and was negatively related to presence of lungworm ( Dictyocaulus spp.). The birth rate calculated as the average number of offspring delivered per adult female was <1.0 overall (2017–2020, LCI = 0.22, UCI = 0.86), similar across years, but increased with age. Logistic exposure models indicated that daily calf survival to 60 d increased as Julian birth date and days since birth increased (log odds = 0.0819, SE = 0.0215). The per capita independence rate, or rate that adult females add independent calves to the population, was negatively related to UN:C ratios and positively with fGCM. Further, this rate was related to autumnal habitat use of adult females; it was greater in home ranges characterized by large amounts of mature (canopy) evergreen forests and wetland habitats, and small amounts of mixed forests and elevation than in ranges with abundant levels of mixed forest at high elevation. We conclude that winter ticks can negatively affect moose fecundity, and efforts to reduce host (moose) density through harvest or parasite (host) abundance through habitat manipulation may improve productivity and recruitment in local moose populations.

Vermont

Exploring management options for moose at their southern range limits considering growing disease risk

1. Populations of cold-adapted species are increasingly vulnerable along their low-latitude range limits due to shifting environmental conditions, biotic interactions, and anthropogenic pressures. Managing these populations is particularly challenging because of complex ecological dynamics, conflicting stakeholder interests, and decision-making under uncertainty. 2. We explored population growth (λ) of moose ( Alces alces ) under different hypothetical management scenarios, simulating combinations of five hypothetical harvest levels with three levels of disease impact. 3. Facing current disease, the population could support multiple levels of conservative harvest. Under elevated levels of disease-caused mortality, projections indicated declines for all harvest scenarios. We projected increases across all harvest scenarios when overlap with white-tailed deer ( Odocoileus virginianus ) was hypothetically reduced and disease mortality in moose minimized. We investigated whether uncertainty in moose demographic parameters altered population trajectory, and found changes in adult fecundity and calf survival could alter harvest decisions when λ<1. 4. Practical implication . While moose population trajectories may remain stable under current conditions, management of white-tailed deer that reduces moose exposure to lethal parasites may provide the greatest utility in sustaining moose in New York given the potential for increased disease. Continued monitoring of population size, growth, and disease prevalence would inform sustainable moose harvest levels that balance social and ecological management considerations.

New York

State-dependent behavior alters endocrine–energy relationship: Implications for conservation and management

Glucocorticoids (GC) and triiodothyronine (T3) are two endocrine markers commonly used to quantify resource limitation, yet the relationships between these markers and the energetic state of animals has been studied primarily in small-bodied species in captivity. Free-ranging animals, however, adjust energy intake in accordance with their energy reserves, a behavior known as state-dependent foraging. Further, links between life-history strategies and metabolic allometries cause energy intake and energy reserves to be more strongly coupled in small animals relative to large animals. Because GC and T3 may reflect energy intake or energy reserves, state-dependent foraging and body size may cause endocrine–energy relationships to vary among taxa and environments. To extend the utility of endocrine markers to large-bodied, free-ranging animals, we evaluated how state-dependent foraging, energy reserves, and energy intake influenced fecal GC and fecal T3 concentrations in free-ranging moose ( Alces alces ). Compared with individuals possessing abundant energy reserves, individuals with few energy reserves had higher energy intake and high fecal T3 concentrations, thereby supporting state-dependent foraging. Although fecal GC did not vary strongly with energy reserves, individuals with higher fecal GC tended to have fewer energy reserves and substantially greater energy intake than those with low fecal GC. Consequently, individuals with greater energy intake had both high fecal T3 and high fecal GC concentrations, a pattern inconsistent with previous documentation from captive animal studies. We posit that a positive relationship between GC and T3 may be expected in animals exhibiting state-dependent foraging if GC is associated with increased foraging and energy intake. Thus, we recommend that additional investigations of GC– and T3–energy relationships be conducted in free-ranging animals across a diversity of body size and life-history strategies before these endocrine markers are applied broadly to wildlife conservation and management.

Wyoming

Life-history theory provides a framework for detecting resource limitation: A test of the Nutritional Buffer Hypothesis

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity—a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose ( Alces alces ) populations that varied in calf recruitment (33–69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

Colorado, Utah, Wyoming

Monitoring animal populations with cameras using open, multistate, N-mixture models

Remote cameras have become a mainstream tool for studying wildlife populations. For species whose developmental stages or states are identifiable in photographs, there are opportunities for tracking population changes and estimating demographic rates. Recent developments in hierarchical models allow for the estimation of ecological states and rates over time for unmarked animals whose states are known. However, this powerful class of models has been underutilized because they are computationally intensive, and model outputs can be difficult to interpret. Here, we use simulation to show how camera data can be analyzed with multistate, Dail-Madsen (hereafter multistate DM) models to estimate abundance, survival, and recruitment. We evaluated four commonly encountered scenarios arising from camera trap data (low and high abundance and 25% and 50% missing data) each with 18 different sample size combinations (camera sites = 40, 250; surveys = 4, 8, and 12; and years = 2, 5, 10) and evaluated the bias and precision of abundance, survival, and recruitment estimates. We also analyzed our empirical camera data on moose ( Alces alces ) with multistate DM models and compared inference with telemetry studies from the same time and region to assess the accuracy of camera studies to track moose populations. Most scenarios recovered the known parameters from our simulated data with higher accuracy and increased precision for scenarios with more sites, surveys, and/or years. Large amounts of missing data and fewer camera sites, especially at higher abundances, reduced accuracy, and precision of survival and recruitment. Our empirical analysis provided biologically realistic estimates of moose survival and recruitment and recovered the pattern of moose abundance across the region. Multistate DM models can be used for estimating demographic parameters from camera data when developmental states are clearly identifiable. We discuss several avenues for future research and caveats for using multistate DM models for large-scale population monitoring.

New Hampshire, Vermont

SiMPL wildlife magnets: A camera trap tool for detecting all creatures great and small

We created the SiMPL wildlife magnet—a baited camera trap design that allows passive monitoring of wildlife, particularly small- to medium-sized mammals. The SiMPL wildlife magnet is inexpensive and easy to construct. To evaluate its effectiveness, we conducted a two-year case study using 9 camera stations along an elevation gradient in the White Mountains of the northeastern United States. We examined how the detection probability of mammal species changed with the inclusion of a SiMPL wildlife magnet using data from pre-and post-establishment. We found a significant increase in community-level detection probability with the use of SiMPL wildlife magnets and for individual species, including red squirrels ( Tamiasciurus hudsonicus ), American marten ( Martes americana ), and fisher ( Pekania pennanti ). Moreover, we were able to capture more species with SiMPL wildlife magnets than without, including flying squirrels ( Glaucomys spp.), various rodents ( Cricetidae spp.), black bears ( Ursus americanus ), moose ( Alces alces ), owls, and other birds. The SiMPL wildlife magnet is an effective, low-cost method for surveying wildlife communities, especially rodents and mesocarnivores. It addresses the limited field of view presented by other techniques for capturing small mammals on camera traps and enables efficient collection of phenology data, including vegetation and snowpack. This tool has several applications, including monitoring species' responses to management practices and global change.

Ecology and Evolution

Moose survival and habitat‐associated risk of endoparasites

Parasite-induced morbidity and mortality can alter the trajectories of incidental host populations. Yet, parasites rarely act in isolation and may be one of a multitude of biotic and abiotic stressors that collectively shape mortality risk in vertebrate populations. We quantified sources of mortality in a low-density population of moose ( Alces alces ) in New York State and investigated factors including parasite infection, nutritional limitation, and thermal stress influencing mortality risk in calf moose. We observed high rates of annual survival (0.81–0.92) in adult ( n = 25) and calf ( n = 27) moose monitored 2015–2018 and 2022–2024, respectively. The majority of cause-specific mortality was attributed to disease induced by giant liver fluke ( Fascioloides magna ; 75% in adults, 67% in calves). Calf mortality risk increased by 72% for every unit increase in giant liver fluke infection risk, measured as cumulative monthly proportion of wetlands used by moose, and decreased by 16% with each additional unit of nutritional energy available. The combination of flukes, coinfecting parasites, and available nutritional energy is important to calf survival in this population, highlighting the importance of managing multiple stressors for species conservation, although the effects are hard to disentangle given the high rates of survival observed. Identifying causes of mortality and mechanisms underlying increased mortality risk contributes to the continued conservation of moose in fluctuating populations and highlights the importance of managing parasite-induced disease.

New York

Warm places, warm years, and warm seasons increase parasitizing of moose by winter ticks

Observed links between parasites, such as ticks, and climate change have aroused concern for human health, wildlife population dynamics, and broader ecosystem effects. The one-host life history of the winter tick ( Dermacentor albipictus ) links each annual cohort to environmental conditions during three specific time periods when they are predictably vulnerable: spring detachment from hosts, summer larval stage, and fall questing for hosts. We used mixed-effects generalized linear models to investigate the drivers of tick loads carried by moose ( Alces alces ) relative to these time periods and across 750 moose, 10 years, and 16 study areas in the western United States. We tested for the effects of biotic factors (moose density, shared winter range, vegetation, migratory behavior) and weather conditions (temperature, snow, humidity) during each seasonal period when ticks are vulnerable and off-host. We found that warm climatic regions, warm seasonal periods across multiple partitions of the annual tick life cycle, and warm years relative to long-term averages each contributed to increased tick loads. We also found important effects of snow and other biotic factors such as host density and vegetation. Tick loads in the western United States were, on average, lower than those where tick-related die-offs in moose populations have occurred recently, but loads carried by some individuals may be sufficient to cause mortality. Lastly, we found interannual variation in tick loads to be most correlated with spring snowpack, suggesting this environmental component may have the highest potential to induce change in tick load dynamics in the immediate future of this region.

Ecosphere

Winter tick sharing between ungulates in the Greater Yellowstone Ecosystem and implications for apparent competition

Host species heterogeneity can drive parasite dynamics through variation in host competency as well as host abundance. We explored how elk ( Cervus canadensis ) with apparent subclinical infestations of winter tick ( Dermacentor albipictus ) may be a cryptic reservoir and drive winter tick dynamics, impacting moose ( Alces alces ) populations. We found that winter tick infestation loads did not vary remarkably between both host species and winter ticks sourced from elk and moose produced similar numbers of larvae which activated within 5 days of each other. We also found similar larval densities in habitats predominately used by elk, moose, and both host species. Our analysis of 2793 informative single-nucleotide polymorphisms showed genetic differentiation among tick populations that were only ~75 km apart, but fewer differences among ticks from elk or moose in the same locality, suggesting sharing of winter ticks across host species. Despite the clinical signs of high winter tick infestations being most apparent on moose, elk may be critical drivers of winter tick population dynamics and indirectly compete with moose in areas where they outnumber moose populations, a common characteristic of ungulate communities in western North America. Management interventions aimed at addressing winter tick issues on moose may wish to consider the movement patterns and abundance of cryptic reservoirs like elk.

Wyoming

Antipredator response diminishes during periods of resource deficit for a large herbivore

The starvation-predation hypothesis predicts that, during resource shortages, prey forego antipredator behavior and forage as much as possible to avoid starvation, even when risk of predation is high. We tested this hypothesis using GPS locations collected simultaneously from moose (Alces alces) and wolves (Canis lupus) in the Greater Yellowstone Ecosystem of North America. We assessed shifts in the speed, displacement, and habitat selection of moose 24 hours following encounter with wolves (0–1500 m distance). We examined whether the strength of antipredator behaviors would weaken as winter progressed and the nutritional condition of moose declined. Moose responded to wolf encounters by increasing their rate of movement in early winter, but only within 500 m distance. Importantly, these responses attenuated as winter progressed. Moose did not avoid their preferred foraging habitat (riparian areas) following encounters with wolves at any distance, and instead they more strongly selected riparian areas, especially in early winter. Our findings support theoretical predictions that resource deficits should dampen prey antipredator behavior, and suggest that nutritional condition of prey may buffer against run-away risk effects in food webs involving large mammalian predators and prey.

Wyoming

Estimating occupancy probability of moose using hunter survey data

Monitoring rare species can be difficult, especially across large spatial extents, making conventional methods of population monitoring costly and logistically challenging. Citizen science has the potential to produce observational data across large areas that can be used to monitor wildlife distributions using occupancy models. We used citizen science (i.e., hunter surveys) to facilitate monitoring of moose ( Alces alces ) populations, an especially important endeavor because of their recent apparent declines in the northeastern and upper midwestern regions of the United States. To better understand patterns of occurrence of moose in New York, we used data collected through an annual survey of approximately 11,000 hunters between 2012 and 2014 that recorded detection–non-detection data of moose and other species. We estimated patterns of occurrence of moose in relation to land cover characteristics, climate effects, and interspecific interactions using occupancy models to analyze spatially referenced moose observations. Coniferous and deciduous forest with low prevalence of white-tailed deer ( Odocoileus virginianus ) had the highest probability of moose occurrence. This study highlights the potential of data collected using citizen science for understanding the spatial distribution of low-density species across large spatial extents and providing key information regarding where and when future research and management activities should be focused.

New York

Climate change effects on deer and moose in the midwest

Climate change is an increasing concern for wildlife managers across the United States and Canada. Because climate change may alter populations and harvest dynamics of key species in the region, midwestern states have identified the effects of climate change on ungulates as a priority research area. We conducted a literature review of projected climate change in the Midwest and the potential effects on white‐tailed deer ( Odocoileus virginianus ) and moose ( Alces alces ). Warmer temperatures and decreasing snowpack in the region favor survival of white‐tailed deer. In contrast, moose may become physiologically stressed in response to warming, and increasing deer populations spreading disease will exacerbate the problem. Although there is some uncertainty about exactly how the climate will change, and to what degree, robust projections suggest that deer populations will increase in response to climate change and moose populations will decrease. Managers can begin preparing for these changes by proactively creating management plans that take this into account.

Journal of Wildlife Management

Effects of winter ticks and internal parasites on moose survival in Vermont, USA

Moose ( Alces alces ) have experienced considerable declines along the periphery of their range in the northeastern United States. In Vermont, the population declined 45% from 2010 to 2017 despite minimal hunter harvest and adequate habitat. Similarly, nearby populations recently experienced epizootics characterized by >50% mortality. Declines have largely been associated with the effects of winter ticks ( Dermacentor albipictus ), but uncertainty exists about the effects of environmental and other parasite-related conditions on moose survival. We examined patterns of moose survival among a radio-collared population ( n = 127) in Vermont from 2017 to 2019. Our objectives were to estimate causes of mortality and model survival probability as a function of individual and landscape variables for calves (<1 yr) and adults (≥1 yr). Observed adult survival was 90% in 2017, 84% in 2018, and 86% in 2019, and winter calf survival was 60% in 2017, 50% in 2018, and 37% in 2019. Winter tick infestation was the primary cause of mortality (91% of calves, 25% of adults), and 32% of all mortalities had evidence of meningeal worm ( Parelaphostrongylus tenuis ). Other sources of mortality such as vehicles, harvest, predation, deep snow, and other parasitic infections were negligible. The best supported calf model included sex differences and negative effects of tick engorgement (%/week) and parasite level (roundworm and lungworm). The best supported adult model included the effect of cumulative tick engorgement (cumulative %/week), which negatively affected survival. Our results indicate that winter tick engorgement strongly affects survival, and is probably compounded by the presence of meningeal worm and other parasites. Reduced tick effects may be achieved by decreasing moose density through harvest and managing late winter habitat to minimize tick density. Management of white-tailed deer ( Odocoileus virginianus ) density may also affect the transmission of meningeal worm.

Vermont