Moose (Alces alces) reacts to high summer temperatures by utilizing thermal shelters in boreal forests – an analysis based on airborne laser scanning of the canopy structure at moose locations

The adaptation of different species to warming temperatures has been increasingly studied. Moose (Alces alces) is the largest of the ungulate species occupying the northern latitudes across the globe, and in Finland it is the most important game species. It is very well adapted to severe cold temperatures, but has a relatively low tolerance to warm temperatures. Previous studies have documented changes in habitat use by moose due to high temperatures. In many of these studies, the used areas have been classified according to how much thermal cover they were assumed to offer based on satellite/aerial imagery data. Here, we identified the vegetation structure in the areas used by moose under different thermal conditions. For this purpose, we used airborne laser scanning (ALS) data extracted from the locations of GPS‐collared moose. This provided us with detailed information about the relationships between moose and the structure of forests it uses in different thermal conditions and we were therefore able to determine and differentiate between the canopy structures at locations occupied by moose during different thermal conditions. We also discovered a threshold beyond which moose behaviour began to change significantly: as day temperatures began to reach 20 °C and higher, the search for areas with higher and denser canopies during daytime became evident. The difference was clear when compared to habitat use at lower temperatures, and was so strong that it provides supporting evidence to previous studies, suggesting that moose are able to modify their behaviour to cope with high temperatures, but also that the species is likely to be affected by warming climate.     

Seasonal release from competition explains partial migration in European moose

Partial migration, whereby a proportion of a population migrates between distinct seasonal ranges, is common throughout the animal kingdom. However, studies linking existing theoretical models of migration probability, with empirical data are lacking. The competitive release hypothesis for partial migration predicts that due to density-dependent habitat selection, the proportion of migrants increases as the relative quality and size of the seasonal range increases, but decreases with increasing migration cost and population density. To test this prediction, we developed a quantitative framework to predict the proportion of migrants, using empirical data from 545 individually GPS-marked moose Alces alces from across Fennoscandia, spanning latitudes of 56° to 68°N. Moose contracted their ranges to common and spatially limited winter areas (typically at lower elevation), but expanded them during summer due to an increase in suitable habitat (at highland ranges). As predicted from our model, a better and larger highland range relative to the lowland range corresponded to a higher proportion of migrants in an area. Quantitative predictions coupling the balance of habitat availability of seasonal ranges with the probability of migrating in a large herbivore is a necessary step towards an enhanced understanding of the mechanisms underlying migration at the population level.     

What drives fine‐scale movements of large herbivores? A case study using moose

Understanding animal movements across heterogeneous landscapes is of great interest because it helps explain the dynamic processes influencing the distribution of individuals in space. Research on how animals move relative to short‐range environmental characteristics are scarce. Our objective was to determine the variables influencing movement of a large ungulate, the moose Alces alces, ranging across a boreal landscape, and to link movement behaviour with limiting factors at a fine scale. We assessed 7 candidate models composed of vegetation, solar energy, and topography variables using step selection functions (SSF) for male and female moose across daily and annual periods. We selected and weighted models using the Bayesian Information Criterion. Variables influencing small‐scale movements of moose differed among periods and between sexes, likely in response to corresponding changes in the importance of limiting factors. Best models often combined many types of variables, although simpler models composed of only vegetation or topography variables explained male's movements during rut and early winter. Moose steps were observed in good feeding stands from summer to early winter for females and from spring to early winter for males, supporting other studies of moose habitat selection. From summer to early winter, females alternatively selected and avoided cover stands during day and night, respectively. Solar energy reaching the ground was important, particularly during late winter and spring, likely due to its effect on snow cover, air temperature, or plant phenology. Moose generally moved in gentle slopes and variable elevation, which may have increased their chances of finding high quality forage, or improved their search of suitable calving sites or mates. Our study revealed the great complexity and dynamic aspects of animal movements in a heterogeneous landscape. Analysis of animal movement provides complementary information to more static habitat selection analyses and helps understanding the spatial variations in the distribution of individuals through time.     

Density-dependent habitat selection and partitioning between two sympatric ungulates

Theory on density-dependent habitat selection predicts that as population density of a species increases, use of higher quality (primary) habitat by individuals declines while use of lower quality (secondary) habitat rises. Habitat partitioning is often considered the primary mechanism for coexistence between similar species, but how this process evolves with changes in population density remains to be empirically tested for free-ranging ungulates. We used resource-selection functions to quantify density effects on landscape-scale habitat selection of two sympatric species of ungulates [moose (Alces alces) and elk (Cervus canadensis manitobensis)] in Riding Mountain National Park, Manitoba, Canada (2000–2011). The density of elk was actively reduced from 1.2 to 0.4 elk km^?2 through increased hunting effort during the period of study, while moose density decreased without additional human influence from 1.6–0.7 moose km^?2. Patterns of habitat selection during winter by both species changed in accordance to expectations from density-dependent habitat-selection theory. At low intraspecific density, moose and elk did not partition habitat, as both species selected strongly for mixed forest (primary habitat providing both food and cover), but did so in different areas segregated across an elevational gradient. As intraspecific density increased, selection for primary habitat by both species decreased, while selection for secondary, lower quality habitat such as agricultural fields (for elk) and built-up areas (for moose) increased. We show that habitat-selection strategies during winter for moose and elk, and subsequent effects on habitat partitioning, depend heavily on the position in state space (density) of both species.     

Patterns and causes of demographic variation in a harvested moose population: evidence for the effects of climate and density-dependent drivers

1.?Better understanding of the mechanisms affecting demographic variation in ungulate populations is needed to support sustainable management of harvested populations. While studies of moose Alces alces L. populations have previously explored temporal variation in demographic processes, managers responsible for populations that span large heterogeneous landscapes would benefit from an understanding of how demography varies across biogeographical gradients in climate and other population drivers. Evidence of thresholds in population response to manageable and un-manageable drivers could aid resource managers in identifying limits to the magnitude of sustainable change. 2.?Generalized additive models (GAMs) were used to evaluate the relative importance of population density, habitat abundance, summer and winter climatic conditions, primary production, and harvest intensity in explaining spatial variation in moose vital rates in Ontario, Canada. Tree regression was used to test for thresholds in the magnitudes of environmental predictor variables that significantly affected population vital rates. 3.?Moose population growth rate was negatively related to moose density and positively related to the abundance of mixed deciduous habitat abundant in forage. Calf recruitment was negatively related to a later start of the growing season and calf harvest. The ratio of bulls to cows was related to male harvest and hunter access, and thresholds were evident in predictor variables for all vital rate models. 4.?Findings indicate that the contributions of density-dependent and independent factors can vary depending on the scale of population process. The importance of density dependence and habitat supply to low-density ungulate populations was evident, and management strategies for ungulates may be improved by explicitly linking forest management and harvest. Findings emphasize the importance of considering summer climatic influences to ungulate populations, as recruitment in moose was more sensitive to the timing of vegetation green-up than winter severity. The efficacy of management decisions for harvested ungulates may require regional shifts in targets where populations span bioclimatic gradients. The use of GAMs in combination with recursive partitioning was demonstrated to be an informative analytical framework that captured nonlinear relationships common in natural processes and thresholds that are relevant to population management in diverse systems.     

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.     

Using agent-based models to inform the dynamics of winter tick parasitism of moose

North American moose (Alces alces) populations along the southern extent of their range have been experiencing high levels of calf mortality in recent years. In New England, this phenomenon has been linked to extensive blood loss resulting from extreme winter tick (Dermacentor albipictus) parasitism. Moose are symbolic of the region and generate income through tourism and the auction of hunting permits; thus, successive years of greater than 50% calf mortality (epizootics) are of concern to wildlife managers and others. It is hypothesized that high localized moose density coupled with climate change are the driving forces behind moose-winter tick epizootics; however, the impact that variable combinations of these and other factors have on the occurrence and magnitude of epizootics has not been previously simulated. We, therefore, developed and implemented a spatially explicit agent-based model with two model environments, each representative of a distinct location within an ongoing field study site in northern New Hampshire that differed in the proportional availability of optimal moose habitat. Three experiments were devised to test the sensitivity of the outcome variables, calf infestation level and calf mortality, to 1) winter tick abundance, 2) winter tick aggregation, and 3) moose density, for the length of the winter tick questing period. Each experiment was conducted in both model environments. This model generated similar mortality levels to those measured in the field study under representative moose density and weather conditions. Additionally, the modeled moose agents and the radio-collared moose in the field reflect comparable habitat usage. While the infestation levels reported by calf agents are considered conservative, we believe that future versions of this model, parameterized with more accurate estimates of winter tick abundance and moose density, may be an effective tool for managing moose against winter tick parasitism.     

Environmental variation and moose Alces alces density as determinants of spatio-temporal heterogeneity in browsing

Understanding temporal variation in habitat selection and browsing intensity by large herbivores is fundamental because of their large impact on the ecosystems. I studied the annual variation in winter browsing pressure on young trees and habitat selection by moose Alces alces over a ten year period. Specifically, the relationships between browsing pressure on Scots pine Pinus sylvestris and two birch species ( Betula ssp.) and three explanatory variables – 1) availability of forage, 2) moose density (estimated by pellet group counts) and 3) snow cover was studied. At a larger spatial scale (forest stand level) the relationship between moose habitat selection between three different habitat types (forest <30 yr, forest>30 yr and mire) and two explanatory variables, 1) snow condition and 2) moose density, were studied. Browsing pressure on Scots pine, the dominating food plant, was related to forage availability, moose density and snow condition. No significant relationships between any of the three explanatory variables and browsing pressure on the two birch species were found. Moose selection for certain habitats varied between years and was affected by number of days with >0.10 m of snow.
Habitat selection was not significantly related to moose density and the relationship between overall moose density and habitat specific moose densities was proportional within the studied density range. These findings have implications for understanding varying browsing patterns – and will affect both the ability to predict herbivores' effect on the forest ecosystem. A snow dependent browsing pattern also indicates that one can expect a long term decrease in browsing pressure on the tree and shrub layer as a consequence of the ongoing large-scale climate change.     

Comparative Space Use and Habitat Selection of Moose Around Feeding Stations

ABSTRACT The practice of feeding cervids in winter, either as a supplement to enhance nutritional status or to divert animals away from roads, railways, or vulnerable habitats, is rising noticeably. Moose (Alces alces) densities in Scandinavia are currently at historically high levels, resulting in amplified damage to economically important young Scots pine (Pinus sylvestris) forest stands. Nevertheless, there is limited information as to how diversionary feeding affects herbivore space use and habitat selection. We followed 32 female moose marked with Global Positioning System collars to evaluate 1) if feeding stations serve as attraction points to the extent that habitat-selection patterns resemble those of central-place foragers (i.e., high usage and more uniform selection close to the attraction point), and 2) if moose using feeding sites select young pine stands less than those not using feeding sites. Moose that used diversionary forage concentrated their space use around feeding stations and selected habitats as predicted for a central-place forager with a decreasing probability of using areas away from feeding sites and a low degree of habitat selectivity close to feeding sites. However, moose that used feeding sites continued to select young pine stands to the same extent as moose that did not use feeding sites. Feeding sites were, therefore, not successful in diverting moose away from valuable natural browse, so we recommend wildlife managers establish feeding sites in sacrifice areas where moose browsing is permissible and, if possible, >1 km from young pine plantations.     

Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use.     

Range Expansion of Moose in Arctic Alaska Linked to Warming and Increased Shrub Habitat

Twentieth century warming has increased vegetation productivity and shrub cover across northern tundra and treeline regions, but effects on terrestrial wildlife have not been demonstrated on a comparable scale. During this period, Alaskan moose (Alces alces gigas) extended their range from the boreal forest into tundra riparian shrub habitat; similar extensions have been observed in Canada (A. a. andersoni) and Eurasia (A. a. alces). Northern moose distribution is thought to be limited by forage availability above the snow in late winter, so the observed increase in shrub habitat could be causing the northward moose establishment, but a previous hypothesis suggested that hunting cessation triggered moose establishment. Here, we use recent changes in shrub cover and empirical relationships between shrub height and growing season temperature to estimate available moose habitat in Arctic Alaska c. 1860. We estimate that riparian shrubs were approximately 1.1 m tall c. 1860, greatly reducing the available forage above the snowpack, compared to 2 m tall in 2009. We believe that increases in riparian shrub habitat after 1860 allowed moose to colonize tundra regions of Alaska hundreds of kilometers north and west of previous distribution limits. The northern shift in the distribution of moose, like that of snowshoe hares, has been in response to the spread of their shrub habitat in the Arctic, but at the same time, herbivores have likely had pronounced impacts on the structure and function of these shrub communities. These northward range shifts are a bellwether for other boreal species and their associated predators.     

Lying in wait: Limiting factors on a low-density ungulate population and the latent traits that can facilitate escape from them

Predation, habitat, hunting, and environmental conditions have all been implicated as regulatory mechanisms in ungulate populations. The low-density equilibrium hypothesis predicts that in low-density populations, predators regulate their prey and that the population will not escape unless predation pressure is eased. We evaluated survival of adult and juvenile moose (Alces alces) in north-central Alaska to determine whether or not the population supported the hypothesis. We instrumented adult male and female moose with radiocollars and used aerial observations to track parturition and subsequent survival of juvenile moose. Generalized linear mixed-effects models were used to assess survival. Adult annual survival rates were high (∼89%), but may be negatively influenced by winter conditions. Migratory status did not affect moose survivorship or productivity. Approximately 60% of the calf crop died before 5 months of age. Productivity was significantly lower in the northern section of the study area where there is less high-quality habitat, suggesting that, even in this low-density population, nutrition could be a limiting factor. It appears that predation on young calves, winter weather, and nutritional constraints may be interacting to limit this population. Latent traits, such as overproduction of calves and migratory behavior, which do not currently enhance fitness, may persist within this population so that individuals with these traits can reap benefits when environmental conditions change.     

Spatial and temporal predictions of moose winter distribution

Herbivores are usually distributed unevenly across the landscape often because of variation in resource availability. We used zero-inflated generalised additive models (to account for data with a high number of zeros) that include georeferences to predict winter distribution of a large herbivore (moose Alces alces). Moose distribution was analysed in relation to forage availability and distance to neighbouring sites. Our results showed that the ability to explain moose distribution indexed by pellet count data at a local scale increased when spatial information (longitude and latitude) was added to the model compared to the model only including food availability. By using the relationship between moose and forage distribution, and the spatial information, we predicted patch choice by moose reasonably well in 2 out of 4?years. However, the distribution of moose was also influenced by weather conditions, as it was most clumped in the year with most snow. In conclusion, our study lends support for a non-linear approach using georeferences for a comprehensive understanding of herbivore distribution at a small scale. This result also indicates that the use of a certain patch by moose not only depends on the selected patch itself but is also influenced by the neighbouring patch and factors at a larger spatial scale, such as moose management influencing the density above moose home range level. The relatively high proportion of unexplained variation suggests that the use of a certain patch is also influenced by other factors such as topography, predation, competition, weather conditions, and wildlife management strategies.     

Contrasting responses of two passerine bird species to moose browsing

Large herbivores may modify the ecosystem in a way that affects habitat quality and resource availability for other fauna. The increase in wild ungulate abundance in many areas may therefore lead to ecosystem changes, affecting distribution and reproduction of other species. Moose (Alces alces) in Scandinavia is a good example of a herbivore that has recently increased in abundance and has the potential to affect the ecosystem. In this study, we investigated how different levels of moose winter activity around supplementary feeding stations for moose affect reproduction in two insectivorous passerines: great tits (Parus major) and pied flycatchers (Ficedula hypoleuca). The two bird species showed contrasting responses to high moose activity at feeding stations. Great tits avoided habitats with high moose activity, where fledging success and feeding frequency was lower than at low moose activity habitats. Flycatchers nested more often at high moose activity habitats where fledging weight and feeding frequency were higher than at low moose activity habitats. Filming of nest boxes with great tits showed an increase in adult Lepidoptera in the diet at supplementary feeding stations for moose, and a smaller size of caterpillar prey at intermediate moose activity. The results support the hypothesis that herbivores may affect insectivorous passerines through changed arthropod food availability.     

Small mammal responses to moose supplementary winter feeding

Supplementary feeding of wild large herbivores is a widespread practice in North America and Europe. The presence of feeding stations may have ecological consequences through changes to animal distributions, patterns of herbivory and a net nutrient input into the ecosystem. In Fennoscandia, supplementary feeding of moose in winter (Alces alces) is increasing. Although it has been shown to affect bird communities, its effects on small mammal communities were unknown. Here, we studied the effects of moose supplementary feeding stations on plants and on abundance, reproduction, and biomass of small mammals in years with low and high vole abundance. We sampled small mammals with snap traps and conducted surveys of the field layer vegetation, at varying distances from moose supplemental feeding stations. Due to the vegetation changes induced by feeding stations, abundance of common shrews (Sorex araneus) and Microtus voles were positively affected by long-term moose winter feeding, while bank voles (Myodes glareolus) were not affected. Moose feeding stations did not affect reproduction, individual body mass, or the total biomass of small mammals. Moose winter-feeding stations have impacts on nontarget species, providing islands of preferred grass and forb habitat for Microtus spp. and common shrews, allowing them to penetrate into a matrix of less preferred forest habitat.     

Climate change impacts population dynamics and distribution shift of moose (Alces alces) in Heilongjiang Province of China

The earth is experiencing obvious climate warming, which may impact population dynamics and the distribution of moose (Alces alces). In this study, we examined the effects of density dependence, temperature, snow depth, and the vegetation (NDVI) on the population dynamics of moose in Heilongjiang Province of China using historical data. Our results demonstrated that moose distribution had continued to contract from the 1980s to the 1990s; moose densities and late spring temperatures in the 1980s were negatively correlated to the rate of increase of the moose population; low and high snow depths in the 1990s showed positive and negative effects, respectively, on the rate of population increase; and the effect of NDVI in the 1980s was similar to the effect of snow depth. Therefore, we confirmed that moose population dynamics is influenced both by intrinsic density-dependent and extrinsic habitat factors, especially late spring temperatures. In addition, an increase in late spring temperatures may shift the southern limit of the distribution of the moose northwards, or may isolate the southernmost portion of the moose population because the rate of warming is higher to the north of a present latitudinal constriction in range than it is at the latitude of the southernmost moose.     

Population structure and genetic diversity of moose in Alaska

Moose (Alces alces) are highly mobile mammals that occur across arboreal regions of North America, Europe, and Asia. Alaskan moose (Alces alces gigas) range across much of Alaska and are primary herbivore consumers, exerting a prominent influence on ecosystem structure and functioning. Increased knowledge gained from population genetics provides insights into their population dynamics, history, and dispersal of these unique large herbivores and can aid in conservation efforts. We examined the genetic diversity and population structure of moose (n = 141) with 8 polymorphic microsatellites from 6 regions spanning much of Alaska. Expected heterozygosity was moderate (H(E) = 0.483-0.612), and private alleles ranged from 0 to 6. Both F(ST) and R(ST) indicated significant population structure (P < 0.001) with F(ST) < 0.109 and R(ST) < 0.125. Results of analyses from STRUCTURE indicated 2 prominent population groups, a mix of moose from the Yakutat and Tetlin regions versus all other moose, with slight substructure observed among the second population. Estimates of dispersal differed between analytical approaches, indicating a high level of historical or current gene flow. Mantel tests indicated that isolation-by-distance partially explained observed structure among moose populations (R(2) = 0.45, P < 0.01). Finally, there was no evidence of bottlenecks either at the population level or overall. We conclude that weak population structure occurs among moose in Alaska with population expansion from interior Alaska westward toward the coast.     

Moose herbivory,browse quality,and nutrient cycling in an Alaskan treeline community

Moose (Alces alces) browsing on diamondleaf willow (Salix planifolia pulchra) caused significant increases in subsequent growth of stems and leaves in treeline plant communities in central Alaska, USA. Willows growing in the shade were significantly more palatable for moose than those growing in the sun. Moose density had strong effects on rates of nutrient cycling, ostensibly through effects of browsing and inputs from fecal and urinary nitrogen. Moose are a keystone herbivore that likely mediate rates of nutrient cycling in northern ecosystems.     

Female Moose Prioritize Forage Over Mortality Risk in Harvested Landscapes

Since 2010, several moose (Alces alces) populations have declined across North America. These declines are believed to be broadly related to climate and landscape change. At the western reaches of moose continental range, in the interior of British Columbia, Canada, wildlife managers have reported widespread declines of moose populations. Disturbances to forests from a mountain pine beetle (Dendroctonum ponderosae) outbreak and associated salvage logging infrastructure in British Columbia are suspected as a mechanism manifested in moose behavior and habitat selection. We examined seasonal differences in moose habitat selection in response to landscape change from mountain pine beetle salvage logging infrastructure: dense road networks and large intensive forest harvest cutblocks. We used 157,447 global positioning system locations from 83 adult female moose from 2012 to 2016 on the Bonaparte Plateau at the southern edge of the Interior Plateau of central British Columbia to test whether increased forage availability, landscape features associated with increased mortality risk, or the cumulative effects of salvage logging best explain female moose distribution using resource selection functions in an information-theoretic framework. We tested these hypotheses across biological seasons, defined using a cluster analysis framework. The cumulative effects of forage availability and risk best predicted resource selection of female moose in all seasons; however, the covariates included in the cumulative models varied between seasons. The top forage availability model better explained moose habitat use than the top risk model in all seasons, except for the calving and fall seasons where the top risk model (distance to road) better predicted moose space use. Selection of habitat that provides forage in winter, spring, and summer suggests that moose seasonally trade predation risk for the benefits of foraging in early seral vegetation communities in highly disturbed landscapes. Our results identified the need for intensive landscape-scale management to stem moose population declines. Additional research is needed on predator densities, space use, and calf survival in relation to salvage logging infrastructure. © 2020 The Wildlife Society.     

Testing the risk of predation hypothesis: the influence of recolonizing wolves on habitat use by moose

Considered as absent throughout Scandinavia for >100 years, wolves (Canis lupus) have recently naturally recolonized south-central Sweden. This recolonization has provided an opportunity to study behavioral responses of moose (Alces alces) to wolves. We used satellite telemetry locations from collared moose and wolves to determine whether moose habitat use was affected by predation risk based on wolf use distributions. Moose habitat use was influenced by reproductive status and time of day and showed a different selection pattern between winter and summer, but there was weak evidence that moose habitat use depended on predation risk. The seemingly weak response may have several underlying explanations that are not mutually exclusive from the long term absence of non-human predation pressure: intensive harvest by humans during the last century is more important than wolf predation as an influence on moose behavior; moose have not adapted to recolonizing wolves; and responses may include other behavioral adaptations or occur at finer temporal and spatial levels than investigated.