Temperature Mediated Moose Survival in Northeastern Minnesota

ABSTRACT The earth is in the midst of a pronounced warming trend and temperatures in Minnesota, USA, as elsewhere, are projected to increase. Northern Minnesota represents the southern edge to the circumpolar distribution of moose (Alces alces), a species intolerant of heat. Moose increase their metabolic rate to regulate their core body temperature as temperatures rise. We hypothesized that moose survival rates would be a function of the frequency and magnitude that ambient temperatures exceeded the upper critical temperature of moose. We compared annual and seasonal moose survival in northeastern Minnesota between 2002 and 2008 with a temperature metric. We found that models based on January temperatures above the critical threshold were inversely correlated with subsequent survival and explained >78% of variability in spring, fall, and annual survival. Models based on late-spring temperatures also explained a high proportion of survival during the subsequent fall. A model based on warm-season temperatures was important in explaining survival during the subsequent winter. Our analyses suggest that temperatures may have a cumulative influence on survival. We expect that continuation or acceleration of current climate trends will result in decreased survival, a decrease in moose density, and ultimately, a retreat of moose northward from their current distribution.     

Survival and cause-specific mortality of moose calves in Northeastern Minnesota

Ungulate reproductive success (calf production and survival) influences population performance. The moose (Alces alces) population in northeastern Minnesota, USA, has declined 65% from 2006 to 2018 but has begun to stabilize. Because causes of this decline were largely unknown, we investigated production, survival, and cause-specific mortality of calves of the global positioning system (GPS)-collared females in this population. In 2013 and 2014, we GPS-collared 74 neonates and monitored them for survival. In 2015 and 2016, we monitored 50 and 35 calving females for signs of neonatal mortality using changes in adult female velocities and assessed seasonal calf survival by aerial surveys. In 2013 and 2014 (pooled), survival to 9 months was 0.34 (95% CI = 0.23–0.52) for collared calves, and in 2015 and 2016 (pooled) survival was 0.35 (95% CI = 0.26–0.48) for uncollared calves. Mortality in all 4 years was high during the first 50 days of life. In 2013 and 2014 (pooled), calving sites were relatively safe for collared neonates; predator-kills occurred a median 17.0 days after departure and a median 1,142 m from calving sites. Predation was the leading cause of death of collared calves (84% of mortalities), with wolves (Canis lupus) accounting for 77% of these. Other forms of mortality for collared and uncollared calves included drowning, infection, vehicle collision, and natural abandonment. We documented higher wolf predation than other recent studies with similar predator communities. Identifying specific causes of calf mortality and understanding their relations to various landscape characteristics and other extrinsic factors should yield insight into mechanisms contributing to the declining moose population in northeastern Minnesota and serve as a basis for ecologically sound management responses. © 2019 The Wildlife Society.     

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. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Predation on Moose Calves by European Brown Bears

Abstract: In North America, brown bears (Ursus arctos) can be a significant predator on moose (Alces alces) calves. Our study in Sweden is the first in which brown bears are the only predator on moose calves. Bears and moose occurred at densities of about 30/1,000 km² and 920/1,000 km², respectively, and bears killed about 26% of the calves. Ninety-two percent of the predation took place when calves were <1 month old. Bear predation was probably additive to other natural mortality, which was about 10% in areas both with and without bears. Females that lost their calves in spring produced more calves the following year (1.54 calves/F) than females that kept their calves (1.11 calves/F), which reduced the net loss of calves due to predation to about 22%.     

Moose Movement Rates Along Highways and Crossing Probability Models

ABSTRACT We developed and validated a density-adjusted spatial model to predict moose (Alces alces) highway-crossing probability to see if the model could be used as an index of moose-vehicle collision risk. We installed Global Positioning System telemetry collars on 47 moose in the north of the Laurentides Wildlife Reserve, Québec, for 2–36 months. We recorded only 84 highway crossings in spring (0.29% of 28,967 2-hr steps) and 122 crossings in summer (0.18% of 68,337 2-hr steps), despite a high sampling effort and having captured moose close to highways. Moose movement rates during movement steps crossing a highway were on average 3 times higher than during the steps preceding or following highway crossing. Paths used by moose when crossing a highway were characterized by a high proportion of food stands, low proportion of lakes and rivers, and topography typical of a valley. Highway-crossing sites were located in valleys with brackish pools and forest stands providing coniferous cover but a low proportion of lakes and rivers. We adjusted moose crossing probability for local variation in moose density using aerial survey data and assessed crossing probability along the highways in the entire Laurentides Wildlife Reserve. We tested the model using moose-vehicle accident data from 1990 to 2002. The relationship between the density-adjusted crossing probability and number of accidents was relatively loose at the 1-km scale but improved markedly when using longer highway sections (5–15 km; r > 0.80). Our results demonstrate that roads and their surroundings are perceived as low-quality habitat by moose. We also conclude that road segments installed along secondary valleys could be a highly strategic site to deploy mitigation measures such as fences and that it could be desirable to increase the width of road shoulders to reduce forest cover and to eliminate brackish pools to reduce cervid-vehicle collisions. We suggest using empirical data such as location of vehicle-wildlife collisions to plan mitigation measures at a fine scale.     

Managing for Elevated Yield of Moose in Interior Alaska

ABSTRACT Given recent actions to increase sustained yield of moose (Alces alces) in Alaska, USA, we examined factors affecting yield and moose demographics and discussed related management. Prior studies concluded that yield and density of moose remain low in much of Interior Alaska and Yukon, Canada, despite high moose reproductive rates, because of predation from lightly harvested grizzly (Ursus arctos) and black bear (U. americanus) and wolf (Canis lupus) populations. Our study area, Game Management Unit (GMU) 20A, was also in Interior Alaska, but we describe elevated yield and density of moose. Prior to our study, a wolf control program (1976–1982) helped reverse a decline in the moose population. Subsequent to 1975, moose numbers continued a 28-year, 7-fold increase through the initial 8 years of our study (λ_B1 = 1.05 during 1996–2004, peak density = 1,299 moose/1,000 km²). During these initial 8 hunting seasons, reported harvest was composed primarily of males ( = 88%). Total harvest averaged 5% of the prehunt population and 57 moose/1,000 km², the highest sustained harvest-density recorded in Interior Alaska for similar-sized areas. In contrast, sustained total harvests of <10 moose/1,000 km² existed among low-density, predator-limited moose populations in Interior Alaska (≤417 moose/1,000 km²). During the final 3 years of our study (2004–2006), moose numbers declined (λ_B2 = 0.96) as intended using liberal harvests of female and male moose ( = 47%) that averaged 7% of the prehunt population and 97 moose/1,000 km². We intentionally reduced high densities in the central half of GMU 20A (up to 1,741 moose/1,000 km² in Nov) because moose were reproducing at the lowest rate measured among wild, noninsular North American populations. Calf survival was uniquely high in GMU 20A compared with 7 similar radiocollaring studies in Alaska and Yukon. Low predation was the proximate factor that allowed moose in GMU 20A to increase in density and sustain elevated yields. Bears killed only 9% of the modeled postcalving moose population annually in GMU 20A during 1996–2004, in contrast to 18–27% in 3 studies of low-density moose populations. Thus, outside GMU 20A, higher bear predation rates can create challenges for those desiring rapid increases in sustained yield of moose. Wolves killed 8–15% of the 4 postcalving moose populations annually (10% in GMU 20A), hunters killed 2–6%, and other factors killed 1–6%. Annually during the increase phase in GMU 20A, calf moose constituted 75% of the predator-killed moose and predators killed 4 times more moose than hunters killed. Wolf predation on calves remained largely additive at the high moose densities studied in GMU 20A. Sustainable harvest-densities of moose can be increased several-fold in most areas of Interior Alaska where moose density and moose: predator ratios are lower than in GMU 20A and nutritional status is higher. Steps include 1) reducing predation sufficient to allow the moose population to grow, and 2) initiating harvest of female moose to halt population growth and maximize harvest after density-dependent moose nutritional indices reach or approach the thresholds we previously published.     

小兴安岭黑河胜山林区冬季驼鹿的生境选择

2002年、2003年和2004年的12月至3月,在小兴安岭黑河胜山林场开展了驼鹿生境选择的研究。研究中选择了9类与驼鹿生境选择相关的生态因子：植被型、离公路距离、离采伐点距离、平均雪深、隐蔽程度、坡向、坡位、坡度、海拔,运用SPSS软件进行交叉汇总定量分析。结果表明,胜山驼鹿冬季以落叶阔叶林、灌丛为主要生境,影响驼鹿分布的主要生态因子为隐蔽程度、坡位,其次为雪深、坡向、离采伐点距离、离公路距离,坡度、海拔对驼鹿分布的影响不明显。     

Moose calf mortality in central Ontario,Canada

Although some populations remain stable, moose (Alces alces) density and distribution have been declining in many areas along the southern edge of their North American distribution. During 2006–2009, we deployed 99 vaginal implant transmitters (VITs) in 86 adult female moose in central Ontario, Canada to assist in locating and radiocollaring neonatal moose calves. We monitored radiocollared calves to estimate calf survival and assess the relative importance of specific causes of death. Calves in the western portion of our study area (WMU49) were exposed to a 6-day general hunting season, whereas calves in the eastern portion of our study area (Algonquin Provincial Park [APP]) were not exposed to hunting. Annual survival for 87 collared calves was greater in the protected area than the harvested area (72.4 ± 6.8% and 55.8 ± 8.3%, respectively) and averaged 63.7 ± 7.1% overall. Predation by wolves (Canis sp.) and American black bears (Ursus americanus) was the dominant cause of death but occurred predominately in APP, whereas other natural mortality agents were 4× more common in WMU49. Only 16% of the collared calves in WMU49 were harvested each year despite a high proportion (approx. 50%) of accessible, public land. Most natural mortality occurred prior to the autumn hunting season such that reductions in natural mortality had little potential to compensate for calf harvest. Overall, calf survival in our study area was moderate to high and our findings suggest predator control or further restrictions of calf hunting in this area is not justified. © The Wildlife Society, 2013     

Population genetic structure and the effect of founder events on the genetic variability of moose, Alces alces, in Canada

Moose, Alces alces, occur naturally throughout most of Canada but successful introductions of known numbers of animals have been made to the islands of Newfoundland and Cape Breton. Five microsatellite loci were used to investigate the population genetic structure and any change in genetic variability due to founder events of moose in Canada. Comparisons of allele frequencies for moose from 11 regions of the country suggested that there are at least seven genetically distinct populations (P < 0.05) in North America, namely Alberta, eastern Ontario, New Brunswick, Cape Breton, Labrador, western Newfoundland, and the Avalon Peninsula of Newfoundland. The average population heterozygosity was approximately 33% (range from 22 to 41%). UPGMA analysis of Nei's genetic distances produced phenograms similar to what would be expected when geographical location and population history are considered. The loss of heterozygosity due to a single founder event (n = 3; two introductions and a natural colonization) ranged from 14 to 30%, and the cumulative loss of heterozygosity due to two successive founder events (an introduction followed by a natural colonization) was 46%. In these examples loss of genetic variability has not been associated with any known phenotypic deviances, suggesting that populations may be established from a small number of founders. However, the viability of these founded populations over evolutionary timescales cannot be determined and is highly dependent upon chance.     

Spending degrees of freedom in a poor economy: A case study of building a sightability model for moose in northeastern Minnesota

Sightability models are binary logistic-regression models used to estimate and adjust for visibility bias in wildlife-population surveys. Like many models in wildlife and ecology, sightability models are typically developed from small observational datasets with many candidate predictors. Aggressive model-selection methods are often employed to choose a best model for prediction and effect estimation, despite evidence that such methods can lead to overfitting (i.e., selected models may describe random error or noise rather than true predictor–response curves) and poor predictive ability. We used moose (Alces alces) sightability data from northeastern Minnesota (2005–2007) as a case study to illustrate an alternative approach, which we refer to as degrees-of-freedom (df) spending: sample-size guidelines are used to determine an acceptable level of model complexity and then a pre-specified model is fit to the data and used for inference. For comparison, we also constructed sightability models using Akaike's Information Criterion (AIC) step-down procedures and model averaging (based on a small set of models developed using df-spending guidelines). We used bootstrap procedures to mimic the process of model fitting and prediction, and to compute an index of overfitting, expected predictive accuracy, and model-selection uncertainty. The index of overfitting increased 13% when the number of candidate predictors was increased from three to eight and a best model was selected using step-down procedures. Likewise, model-selection uncertainty increased when the number of candidate predictors increased. Model averaging (based on R = 30 models with 1–3 predictors) effectively shrunk regression coefficients toward zero and produced similar estimates of precision to our 3-df pre-specified model. As such, model averaging may help to guard against overfitting when too many predictors are considered (relative to available sample size). The set of candidate models will influence the extent to which coefficients are shrunk toward zero, which has implications for how one might apply model averaging to problems traditionally approached using variable-selection methods. We often recommend the df-spending approach in our consulting work because it is easy to implement and it naturally forces investigators to think carefully about their models and predictors. Nonetheless, similar concepts should apply whether one is fitting 1 model or using multi-model inference. For example, model-building decisions should consider the effective sample size, and potential predictors should be screened (without looking at their relationship to the response) for missing data, narrow distributions, collinearity, potentially overly influential observations, and measurement errors (e.g., via logical error checks). © 2011 The Wildlife Society.     

Temporal,Spatial, and Landscape Habitat Characteristics of Moose—Vehicle Collisions in Western Maine

Abstract: We analyzed moose (Alces alces)-vehicle collisions (MVCs) in western Maine, USA, from 1992 to 2005 (n = 8,156) using Geographic Information Systems to identify patterns of temporal and spatial distribution and develop predictive models based on road and landscape characteristics. We used chi-square and correlation analyses to assess temporal characteristics of MVCs, K-function and kernel analyses to identify spatial clusters of MVCs, and logistic regression to relate covariates for traffic, land-cover, land-form, and relative moose abundance to probability of MVC. We evaluated candidate models using Akaike's Information Criterion, area under the receiver operating characteristic curve (AUC), and the percentage of correctly classified observations. Most (81.6%) MVCs occurred from May to October, with peak monthly frequencies in June (18.6%). Moose-vehicle collisions were clustered spatially on roads at local (0–4 km) and regional scales (22–41 km and 45–54 km), but not at intermediate scales. Traffic-related covariates predicting MVCs included traffic volume and speed limit. For each additional 500 vehicles/day, odds of a location being an MVC increased by 57%. For each 8-km/hr increase in speed limit, odds of an MVC increased by 35%. Landscape composition covariates best predicted MVCs within a 2.5-km radius of the collision site. Mean percent cover within 2.5 km of MVCs was comprised of 36% more cutover forest, 10% more coniferous forest, 5% less deciduous-mixed forest, and 10% less nonwoody wetland than for random points. For every 5% increase in percent cutover and coniferous forest within 2.5 km of the road, predicted odds of MVC increased by 36% and 19%, respectively. Landscape configuration covariates best predicted MVCs within the 5.0-km radius. Moose-vehicle collisions were associated with areas of less interspersion of cover types; for each 5% increase in an index of interspersion-juxtaposition, predicted odds of MVC decreased by 11%. Our final model attained high predictive power (AUC = 0.835) and validation accuracy (75.0%). The model also proved robust to physiographic variation, exhibiting high predictive power (AUC = 0.828) and validation accuracy (68.8%). Managers seeking to prioritize resources for reducing MVCs or predicting future areas of high MVC probability should assess land-cover composition and configuration surrounding MVC hotspots at geographic extents out to 2.5–5 km and use this information to plan expensive roadside management practices such as fencing. The importance of traffic and landscape covariates in our modeling suggests that effective management to reduce MVCs will require a complex combination of driving speed reductions and modifications to forest management along roads.     

Genetic Structure and Evidence of a Local Bottleneck in Moose in Sweden

ABSTRACT The moose (Alces alces) is the most intensely managed game species in Sweden. Despite the biological and socioeconomical importance of moose, little is known of its population genetic structure. We analyzed 132 individuals from 4 geographically separate regions in Sweden for genetic variability at 6 microsatellite loci. We found evidence of strong substructuring and restricted levels of gene flow in this potentially mobile mammal. F_ST values were around 10%, and assignment tests indicated 3 genetically distinct populations over the study area. Spatial autocorrelation analysis provided a genetic patch size of approximately 420 km, implying that moose less than this distance apart are genetically more similar than 2 random individuals. Allele and genotype frequency distributions suggested a recent bottleneck in southern Sweden. Results indicate that moose may be more genetically divergent than currently anticipated, and therefore, the strong hunting pressure that is maintained over all of Sweden may have considerable local effects on genetic diversity. Sustainable moose hunting requires identification of spatial genetic structure to ensure that separate, genetically distinct subpopulations are not overharvested.     

A Demographic model for ring-necked ducks breeding in Minnesota

Population models can provide insights into how perturbations of demographic vital rates influence population growth rates (λ) and help prioritize efforts to achieve management goals. Although population models have been developed for numerous duck species, this tool has not been developed for ring-necked ducks (Aythya collaris). Ring-necked ducks are a late-nesting species that may not be surveyed well by the May Waterfowl Survey, which is timed optimally for earlier nesting ducks like the mallard (Anas platyrhynchos). Information specific to ring-necked ducks would help identify important influences on the rate of population growth and the direction of population change. We used data from our own research during 2008–2012, long-term (1975–2016) survey data from northcentral Minnesota, USA, vital rate estimates from the literature, and long-term (1987–2016) banding data to develop a population model for ring-necked ducks. We estimated annual survival and recovery rates of after-hatching-year and hatching-year female ring-necked ducks in Minnesota. Survey results and our matrix models indicated that ring-necked ducks are declining in the forested portion of Minnesota. Thus, we examined a boom-or-bust simulation scenario that might maintain a population through periodic occurrence of exceptional reproductive years in conjunction with high hatching-year survival. Our results indicated that long-term persistence is only expected in this population if reproductive output doubled at the same time that hatching-year survival was at its highest value, or if the population is maintained through immigration. Sensitivity analysis indicated that unit changes in 30-day brood survival will produce the most change in λ in the parameter space observed, although elasticity analysis indicated that proportional changes in annual survival of adult females will produce the largest proportional changes in λ. Management to improve brood survival to increase λ might include improving the habitat quality of brood-rearing lakes, especially those with more open water and less nesting habitat than those used for nesting. Our findings might also help explain disparities between annual breeding waterfowl surveys, which indicate stable or increasing populations, and hunter experiences in the fall. In Minnesota, hunter experiences have not matched expectations based on historical fall numbers and this could occur if regional production declined and fewer young birds were available for harvest. Our findings highlight the need for further study to inform management in this rapidly changing region. © 2019 The Wildlife Society.     

Influence of young black spruce plantations on moose winter distribution

Logging in the boreal forest may benefit moose by increasing food availability. However, the influence of tree plantations on moose behavior, especially on moose spatial ecology, is poorly understood. We assessed the impacts of black spruce plantations on moose winter distribution at a landscape scale in the Bas-Saint-Laurent region (Québec, Canada). We used winter aerial surveys to examine relationships among plantation characteristics and other habitat variables known to affect moose distribution. The total area of plantations positively influenced moose abundance, but highly aggregated plantations resulted in fewer moose. Moose abundance was also positively associated with food availability and the density of edges between stands providing cover and stands offering high food availability, but moose abundance was negatively associated with road density. Although plantation characteristics were less influential than habitat variables related to foraging and predator avoidance, we demonstrate that the area of black spruce plantations and their configuration should be considered in moose management. We conclude that an integrated management strategy is needed to find a balance between overdeveloped road networks (needed to join homogeneously distributed plantations) and agglomerated plantations in order to mitigate impacts on moose winter distribution. © 2012 The Wildlife Society.