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.     

What regulate and limit reindeer populations in Norway?

An understanding of how species are affected by top-down and bottom-up processes in food webs, and how these forces interact with climatic conditions is crucial for how ecosystems should be managed. In Norway large carnivores are effectively removed from extensive areas to protect livestock, leaving human harvesting as the only significant top-down force on ungulate populations. We examined the relative role of top-down and bottom-up processes for 58 semi-domesticated reindeer populations in Norway subjected to contrasting climatic regimes over the period 1981–2005. Intensive herding and international agreement have resulted in a situation where some populations are unable to undertake seasonal migration to the interior to escape the unfavourable climatic conditions that rule the coastal region in the winter, a critical season for northern ungulates. We used this natural manipulation to contrast between populations with 'poor' and 'good' winter conditions. For populations with good winter conditions, average body size increased with harvesting, suggesting that some top-down process was necessary to avoid food limitation. Time-series analyses revealed that direct regulation of population size was only evident in populations subjected to intensive harvesting. The lack of direct regulation in populations subjected to low harvesting resulted in high vulnerability to harsh winter weather. The body size and climate vulnerability of populations with poor winter conditions was unaffected of harvesting, but average densities was positively related to overall vegetation productivity as indexed by satellite images (NDVI). Top-down processes appeared to be necessary to dampen the effect of harsh winters in populations with generally good winter conditions. Conversely, populations subjected to generally poor winter conditions appeared to be more influenced by bottom-up processes and buffered climatic perturbations by increasing body size.     

Metrics of predation: perils of predator-prey ratios

We developed an original modeling approach using program Stella® to investigate the usefulness of predator–prey ratios (PPRs) for interpreting top-down and bottom-up forcing on moose Alces alces. We included density-dependent feedbacks for the moose population, allowed K to vary based on amount and quality of available forage for moose, integrated effects of compensatory mortality, and added time lags in wolves Canis lupus tracking the moose population. Modeling scenarios we developed included bottom-up and top-down regulation as predetermined outcomes. We then evaluated whether PPRs would reflect the various combinations of trajectories of predator and prey populations under top-down versus bottom-up regulation. The resulting patterns of PPRs were impossible to disentangle from one another, and did not provide reliable insights into whether top-down or bottom-forcing occurred, especially over short time spans where critical decisions related to management of moose and wolves might be necessary. Only under top-down regulation did PPRs reflect the degree of predation experienced by moose, but in that instance, knowledge of top-down regulation must be known a priori to correctly interpret PPRs. Potential problems with interpreting PPRs include their double-variable nature, which resulted in the failure to reflect patterns of increase and decrease for predators and prey. We suggest that confidence intervals for PPRs be calculated from a binomial, similar to that proposed for sex and age ratios, which should help discourage the inappropriate use of this metric. We caution that the temptation to use PPRs often is irresistible, but their reliability is highly questionable. We provide an alternative method to using PPRs or other predation metrics for determining whether top-down or bottom-up forcing is occurring by adopting an approach based on the physical condition and life-history characteristics of prey.     

Abiotic,bottom-up,and top-down influences on recruitment of Rocky Mountain elk in Oregon: A retrospective analysis

Understanding the relative effects of the many factors that may influence recruitment of ungulates is fundamental to managing their populations. Over the last 4 decades, average recruitment in some populations of elk (Cervus elaphus) in Oregon, USA declined from >50 to <20 juveniles per 100 females, and several competing hypotheses address these declines. We developed a priori models and constructed covariates spanning 1977–2005 from hunter-killed elk, elk population estimates, cougar harvest, and weather statistics to evaluate abiotic, bottom-up, and top-down factors that may explain annual variation and long-term trends of pregnancy, juveniles-at-heel in late autumn, and recruitment of juvenile elk in spring. In models of pregnancy status, August precipitation, age, and cougar index had positive effects, whereas previous year (t − 1) winter severity or winter precipitation_(t−1) and elk density had negative effects. In models of juvenile-at-heel in late autumn, August precipitation, August precipitation_(t−1), cougar index × elk density_(t−1), and age had positive effects, whereas cougar index, elk density_(t−1), and winter precipitation_(t−1) had negative effects. Juvenile recruitment was best explained by positive effects of August precipitation_(t−1), lactation rate, and cougar index × elk density_(t−1) and negative effects of cougar index and elk density_(t−1). Winter severity, precipitation, and temperature were not significant in explaining variation in elk recruitment. Annual variation in pregnancy, juvenile-at-heel, and recruitment was most influenced by August precipitation, whereas long-term trends in recruitment were most influenced by cougar densities with relatively weak effects of elk density. These results provide insight into causes of year-to-year and long-term trends of elk recruitment and provide a basis for more rigorous evaluation of factors affecting recruitment of elk. © 2012 The Wildlife Society.     

Environmental fluctuations restrict eco-evolutionary dynamics in predator–prey system

Environmental fluctuations, species interactions and rapid evolution are all predicted to affect community structure and their temporal dynamics. Although the effects of the abiotic environment and prey evolution on ecological community dynamics have been studied separately, these factors can also have interactive effects. Here we used bacteria–ciliate microcosm experiments to test for eco-evolutionary dynamics in fluctuating environments. Specifically, we followed population dynamics and a prey defence trait over time when populations were exposed to regular changes of bottom-up or top-down stressors, or combinations of these. We found that the rate of evolution of a defence trait was significantly lower in fluctuating compared with stable environments, and that the defence trait evolved to lower levels when two environmental stressors changed recurrently. The latter suggests that top-down and bottom-up changes can have additive effects constraining evolutionary response within populations. The differences in evolutionary trajectories are explained by fluctuations in population sizes of the prey and the predator, which continuously alter the supply of mutations in the prey and strength of selection through predation. Thus, it may be necessary to adopt an eco-evolutionary perspective on studies concerning the evolution of traits mediating species interactions.     

Microhabitat selection in the common lizard: implications of biotic interactions,age, sex,local processes,and model transferability among populations

Modeling species' habitat requirements are crucial to assess impacts of global change, for conservation efforts and to test mechanisms driving species presence. While the influence of abiotic factors has been widely examined, the importance of biotic factors and biotic interactions, and the potential implications of local processes are not well understood. Testing their importance requires additional knowledge and analyses at local habitat scale. Here, we recorded the locations of species presence at the microhabitat scale and measured abiotic and biotic parameters in three different common lizard (Zootoca vivipara) populations using a standardized sampling protocol. Thereafter, space use models and cross‐evaluations among populations were run to infer local processes and estimate the importance of biotic parameters, biotic interactions, sex, and age. Biotic parameters explained more variation than abiotic parameters, and intraspecific interactions significantly predicted the spatial distribution. Significant differences among populations in the relationship between abiotic parameters and lizard distribution, and the greater model transferability within populations than between populations are in line with effects predicted by local adaptation and/or phenotypic plasticity. These results underline the importance of including biotic parameters and biotic interactions in space use models at the population level. There were significant differences in space use between sexes, and between adults and yearlings, the latter showing no association with the measured parameters. Consequently, predictive habitat models at the population level taking into account different sexes and age classes are required to understand a specie's ecological requirements and to allow for precise conservation strategies. Our study therefore stresses that future predictive habitat models at the population level and their transferability should take these parameters into account.     

Top-down and bottom-up community regulation in marine rocky intertidal habitats

Strong top-down control by consumers has been demonstrated in rocky intertidal communities around the world. In contrast, the role of bottom-up effects (nutrients and productivity), known to have important influences in terrestrial and particularly freshwater ecosystems, is poorly known in marine hard-bottom communities. Recent studies in South Africa, New England, Oregon and New Zealand suggest that bottom-up processes can have important effects on rocky intertidal community structure. A significant aspect of all of these studies was the incorporation of processes varying on larger spatial scales than previously considered (10’s to 1000’s of km). In all four regions, variation in oceanographic factors (currents, upwelling, nutrients, rates of particle flux) was associated with different magnitudes of algal and/or phytoplankton abundance, availability of particulate food, and rates of recruitment. These processes led to differences in prey abundance and growth, secondary production, consumer growth, and consumer impact on prey resources. Oceanographic conditions therefore may vary on scales that generate ecologically significant variability in populations at the bottom of the food chain, and through upward-flowing food chain effects, lead to variation in top-down trophic effects. I conclude that top-down and bottom-up processes can be important joint determinants of community structure in rocky intertidal habitats, and predict that such effects will occur generally wherever oceanographic ‘discontinuities’ lie adjacent to rocky coastlines. I further argue that increased attention by researchers and of funding agencies to such benthic–pelagic coupling would dramatically enhance our understanding of the dynamics of marine ecosystems.     

Complex numerical responses to top-down and bottom-up processes in vertebrate populations

Population growth rate is determined in all vertebrate populations by food supplies, and we postulate bottom-up control as the universal primary standard. But this primary control system can be overridden by three secondary controls: top-down processes from predators, social interactions within the species and disturbances. Different combinations of these processes affect population growth rates in different ways. Thus, some relationships between growth rate and density can be hyperbolic or even have multiple nodes. We illustrate some of these in marsupial, ungulate and rabbit populations. Complex interactions between food, predators, environmental disturbance and social behaviour produce the myriad observations of population growth in nature, and we need to develop generalizations to classify populations. Different animal groups differ in the combination of these four processes that affect them, in their growth rates and in their vulnerability to extinction. Because conservation and management of populations depend critically on what factors drive population growth, we need to develop universal generalizations that will relieve us from the need to study every single population before we can make recommendations for management.     

Geographical and latitudinal variation in growth patterns and adult body size of Swedish moose (Alces alces)

We examined the geographical pattern in growth and adult body size among 14 populations of Swedish moose (Alces alces) using data from 4,294 moose (1.5 years old) killed during the hunting season in 1989–1992. In both sexes, adult body mass was significantly positively correlated with latitude. Moose in northern populations had a 15–20% larger adult body mass than moose in the south. Juvenile body mass was correlated with neither latitude nor adult body mass. Thus, variation in time (years) and rate of body growth after the juvenile stage were responsible for most of the variation in adult body mass among populations. Moose in northern populations grew for approximately 2 more years of life than southern moose. In contrast to adult body mass, skeletal size (measured as jawbone length) was not correlated with latitude, suggesting that variation in adult body mass was primarily due to differences in fat reserves. Discrimination between population characteristics, such as moose density, climate, and the amount of browse available to moose, showed climatic harshness to be the most important variable explaining geographical variation in body mass among populations. The results support the notion that in mammals body size increases with latitude in accordance with Bergmann's rule. We conclude that (1) variation in patterns of growth after the juvenile stage is the main cause of the latitudinal trend in adult body size in moose, and (2) climatic conditions are a more important factor than population density and availability of food in explaining geographical variation in growth patterns and adult body mass between populations of Swedish moose.     

Annual variation in maternal age and calving date generate cohort effects in moose (<Emphasis Type="Italic">Alces alces</Emphasis>) body mass

A general feature of the demography of large ungulates is that many demographic traits are dependent on female body mass at early ages. Thus, identifying the factors affecting body mass variation can give important mechanistic understanding of demographic processes. Here we relate individual variation in autumn and winter body mass of moose calves living at low density on an island in northern Norway to characteristics of their mother, and examine how these relationships are affected by annual variation in population density and climate. Body mass increased with increasing age of their mother, was lower for calves born late in the spring, decreased with litter size and was larger for males than for female calves. No residual effects of variation in density and climate were present after controlling for annual variation in mother age and calving date. The annual variation in adult female age structure and calving date explained a large part (71–75%) of the temporal variation in calf body mass. These results support the hypotheses that (a) body mass of moose calves are affected by qualities associated with mother age (e.g. body condition, calving date); and (b) populations living at low densities are partly buffered against temporal fluctuations in the environment.     

Within-Plant Bottom-Up Effects Mediate Non-Consumptive Impacts of Top-Down Control of Soybean Aphids

There is increasing evidence that top-down controls have strong non-consumptive effects on herbivore populations. However, little is known about how these non-consumptive effects relate to bottom-up influences. Using a series of field trials, we tested how changes in top-down and bottom-up controls at the within-plant scale interact to increase herbivore suppression. In the first experiment, we manipulated access of natural populations of predators (primarily lady beetles) to controlled numbers of A. glycines on upper (i.e. vigorous-growing) versus lower (i.e. slow-growing) soybean nodes and under contrasting plant ages. In a second experiment, we measured aphid dispersion in response to predation. Bottom-up and top-down controls had additive effects on A. glycines population growth. Plant age and within-plant quality had significant bottom-up effects on aphid size and population growth. However, top-down control was the dominant force suppressing aphid population growth, and completely counteracted bottom-up effects at the plant and within-plant scales. The intensity of predation was higher on upper than lower soybean nodes, and resulted in a non-consumptive reduction in aphid population growth because most of the surviving aphids were located on lower plant nodes, where rates of increase were reduced. No effects of predation on aphid dispersal among plants were detected, suggesting an absence of predator avoidance behavior by A. glycines. Our results revealed significant non-consumptive predator impacts on aphids due to the asymmetric intensity of predation at the within-plant scale, suggesting that low numbers of predators are highly effective at suppressing aphid populations.     

Relative importance of top-down and bottom-up forces in food webs of Sarracenia pitcher communities at a northern and a southern site

The relative importance of resources (bottom-up forces) and natural enemies (top-down forces) for regulating food web dynamics has been debated, and both forces have been found to be critical for determining food web structure. How the relative importance of top-down and bottom-up forces varies between sites with different abiotic conditions is not well understood. Using the pitcher plant inquiline community as a model system, I examine how the relative importance of top-down and bottom-up effects differs between two disparate sites. Resources (ant carcasses) and top predators (mosquito larvae) were manipulated in two identical 4 × 4 factorial press experiments, conducted at two geographically distant sites (Michigan and Florida) within the range of the purple pitcher plant, Sarracenia purpurea, and the aquatic community that resides in its leaves. Overall, top predators reduced the density of prey populations while additional resources bolstered them, and the relative importance of top-down and bottom-up forces varied between sites and for different trophic levels. Specifically, top-down effects on protozoa were stronger in Florida than in Michigan, while the opposite pattern was found for rotifers. These findings experimentally demonstrate that the strength of predator–prey interactions, even those involving the same species, vary across space. While only two sites are compared in this study, I hypothesize that site differences in temperature, which influences metabolic rate, may be responsible for variation in consumer–resource interactions. These findings warrant further investigation into the specific factors that modify the relative importance of top-down and bottom-up effects.     

Population dynamics of cereal aphids: influence of a shared predator and weather

1 Aphid populations may show strong year-to-year fluctuations, but questions remain regarding the dominance of factors that cause this variation, especially the role of natural enemies. To better understand the dynamics of aphid species that occur as pests in cereals, we investigated the relative influence of top-down control by a predator and weather (temperature and precipitation) on population fluctuations of three cereal aphid species.
2 From 1987 to 2005, populations of Metopolophium dirhodum , Sitobion avenae and Rhopalosiphum padi in insecticide-free stands of winter wheat were monitored in the Praha-Ruzyné region of the Czech Republic. Densities of an aphidophagous predator, the ladybeetle Coccinella septempunctata , were recorded from an overwintering site in the landscape. Weather was quantified using historical records.
3 A significant bottom-up effect of densities of aphids on those of C.   septempunctata was found, but evidence of direct top-down regulation of aphids by C.   septempunctata was only significant in the case of R.   padi . There was no significant periodicity in the dynamics of the aphid or C.   septempunctata , suggesting that there was no clear predator-prey cycle. Combinations of C.   septempunctata and weather variables could be used to explain M.   dirhodum and R.   padi per capita rate of change. There were also indications that weather directly affected peak density of M.   dirhodum .
4 We conclude that regional estimates of C.   septempunctata densities are not sufficient to determine whether aphid population dynamics are driven by predator–prey interactions. Feasibility of time series analysis as an investigative tool in aphid population dynamics studies is discussed.     

Spatial, bottom-up, and top-down effects on the abundance of a leaf miner

Many plant and animal species have higher densities at the centre of their distribution, with a gradual decline in abundance towards the edge of the range, though reasons for this pattern is not well known. We examined the abundance of the leaf miner Cameraria sp. nova over the range of its host plant Quercus myrtifolia in Florida and addressed how bottom-up and top-down factors varied over its whole distribution. Leaf miner densities, plant quality and natural enemy effects on mine survivorship were evaluated in 40 sites and spatially structured models were used to determine the effects of spatial location on the abundance of Cameraria and effects of both bottom-up (tannin concentration, foliar nitrogen, soil nitrogen, and leaf area) and top-down factors (larval parasitism and predation) on abundance and survivorship. Cameraria mines were, on average, three times more abundant on edge/coastal sites compared to centre/inland sites and did not support the hypothesis of higher abundance on the centre of the distribution. Differences in plant quality, larval parasitism and successful emergence of mines on edge versus central sites might be partially responsible for this finding. A trend surface equation with latitude and longitude combined explained almost 52% of the variation in Cameraria density and a trend surface map also revealed peaks of Cameraria abundance on the edges of the plant distribution. Correlograms also indicated a significant spatial structure of Cameraria as mines were positively spatially autocorrelated at small distances (≈122 km). Partial regression analyses indicated that 69% of the variation in Cameraria abundance was explained by the effects of latitude, longitude, elevation and percentage of foliar nitrogen. Our results indicated that variation in Cameraria abundance was mostly explained by spatial position and significant effects of bottom-up and top-down factors were not detected in our large-scale study.     

The influence of prey consumption and demographic stochasticity on population growth rate of Isle Royale wolves Canis lupus

The relationship between the rates of prey capture and predator population growth is a fundamental aspect of predation, yet it is rarely measured for vertebrate predators. For the isolated wolf population on Isle Royale, annual variation in kill rate explains 22% of the variation in wolf population growth rate. From the slope of this relationship, we estimate that the production efficiency (ratio of production to respiration) of wolves is between 0.5% and 1.5%. More generally, we assess the relative extent to which wolf population growth rate is affected by density dependence, prey availability (moose, Alces alces ), winter weather, and demographic stochasticity. Prey availability explains the most variation in wolf growth rate (42%), but this is only recognized after accounting for the influence of a disease-induced population crash and age structure of the prey population (i.e. number of vulnerable moose, >9 years of age). Demographic stochasticity accounts for approximately 30% of the variation in wolf growth rate. This recognition is important, but not surprising, given that the average population size of Isle Royale wolves is 22. Previous work indicates that the effect of winter climate, as mediated through prey vulnerability and kill rates, is substantial. This work indicates that the direct effect of winter climate is weak, and explains only about 4% of the variation in wolf growth rate (P=0.10).     

Moose body mass variation revisited: disentangling effects of environmental conditions and genetics

Large-scale geographical variation in phenotypic traits within species is often correlated to local environmental conditions and population density. Such phenotypic variation has recently been shown to also be influenced by genetic structuring of populations. In ungulates, large-scale geographical variation in phenotypic traits, such as body mass, has been related to environmental conditions and population density, but little is known about the genetic influences. Research on the genetic structure of moose suggests two distinct genetic lineages in Norway, structured along a north-south gradient. This corresponds with many environmental gradients, thus genetic structuring provides an additional factor affecting geographical phenotypic variation in Norwegian moose. We investigated if genetic structure explained geographical variation in body mass in Norwegian moose while accounting for environmental conditions, age and sex, and if it captured some of the variance in body mass that previously was attributed to environmental factors. Genetic structuring of moose was the most important variable in explaining the geographic variation in body mass within age and sex classes. Several environmental variables also had strong explanatory power, related to habitat diversity, environmental seasonality and winter harshness. The results suggest that environmental conditions, landscape characteristics, and genetic structure should be evaluated together when explaining large-scale patterns in phenotypic characters or life history traits. However, to better understand the role of genetic and environmental effects on phenotypic traits in moose, an extended individual-based study of variation in fitness-related characters is needed, preferably in an area of convergence between different genetic lineages.     

Estimates of annual survival,growth, and recruitment of a white-tailed ptarmigan population in Colorado over 43 years

Long-term datasets for high-elevation species are rare, and considerable uncertainty exists in understanding how high-elevation populations have responded to recent climate warming. We present estimates of demographic vital rates from a 43-year population study of white-tailed ptarmigan (Lagopus leucura), a species endemic to alpine habitats in western North America. We used capture-recapture models to estimate annual rates of apparent survival, population growth, and recruitment for breeding-age ptarmigan, and we fit winter weather covariates to models in an attempt to explain annual variation. There were no trends in survival over the study period but there was strong support for age and sex effects. The average rate of annual growth suggests a relatively stable breeding-age population ( \( \bar{\lambda } \)  = 1.036), but there was considerable variation between years for both population growth and recruitment rates. Winter weather covariates only explained a small amount of variation in female survival and were not an important predictor of male survival. Cumulative winter precipitation was found to have a quadratic effect on female survival, with survival being highest during years of average precipitation. Cumulative winter precipitation was positively correlated with population growth and recruitment rates, although this covariate only explained a small amount of annual variation in these rates and there was considerable uncertainty among the models tested. Our results provide evidence for an alpine-endemic population that has not experienced extirpation or drastic declines. However, more information is needed to understand risks and vulnerabilities of warming effects on juveniles as our analysis was confined to determination of vital rates for breeding-age birds.     

Large Impact of Eurasian Lynx Predation on Roe Deer Population Dynamics

The effects of predation on ungulate populations depend on several factors. One of the most important factors is the proportion of predation that is additive or compensatory respectively to other mortality in the prey, i.e., the relative effect of top-down and bottom-up processes. We estimated Eurasian lynx (Lynx lynx) kill rate on roe deer (Capreolus capreolus) using radio-collared lynx. Kill rate was strongly affected by lynx social status. For males it was 4.85 ± 1.30 S.E. roe deer per 30 days, for females with kittens 6.23 ± 0.83 S.E. and for solitary females 2.71 ± 0.47 S.E. We found very weak support for effects of prey density (both for Type I (linear) and Type II (non-linear) functional responses) and of season (winter, summer) on lynx kill rate. Additionally, we analysed the growth rate in a roe deer population from 1985 to 2005 in an area, which lynx naturally re-colonized in 1996. The annual roe deer growth rate was lower after lynx re-colonized the study area, but it was also negatively influenced by roe deer density. Before lynx colonized the area roe deer growth rate was λ = 1.079 (± 0.061 S.E.), while after lynx re-colonization it was λ = 0.94 (± 0.051 S.E.). Thus, the growth rate in the roe deer population decreased by Δλ = 0.14 (± 0.080 S.E.) after lynx re-colonized the study area, which corresponded to the estimated lynx predation rate on roe deer (0.11 ± 0.042 S.E.), suggesting that lynx predation was mainly additive to other mortality in roe deer. To conclude, this study suggests that lynx predation together with density dependent factors both influence the roe deer population dynamics. Thus, both top-down and bottom-up processes operated at the same time in this predator-prey system.     

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.     

A piecewise linear modeling approach for testing competing theories of habitat selection: an example with mule deer in northern winter ranges

Habitat selection fundamentally drives the distribution of organisms across landscapes; density-dependent habitat selection (DDHS) is considered a central component of ecological theories explaining habitat use and population regulation. A preponderance of DDHS theories is based on ideal distributions, such that organisms select habitat according to either the ideal free, despotic, or pre-emptive distributions. Models that can be used to simultaneously test competing DDHS theories are desirable to help improve our understanding of habitat selection. We developed hierarchical, piecewise linear models that allow for simultaneous testing of DDHS theories and accommodate densities from multiple habitats and regional populations, environmental covariates, and random effects. We demonstrate the use of these models with data on mule deer (Odocoileus hemionus) abundance and net energy costs in different snow depths within winter ranges of five regional populations in western Idaho, USA. Regional population density explained 40 % of the variation in population growth, and we found that deer were ideal free in winter ranges. Deer occupied habitats with lowest net energy costs at higher densities and at a higher rate than compared to habitats with intermediate and high energy costs. The proportion of a regional population in low energy cost habitat the previous year accounted for a significant amount of variation in population growth (17 %), demonstrating the importance of winter habitat selection in regulating deer populations. These linear models are most appropriate for empirical data collected from centralized habitat patches within the local range of a species where individuals are either year-round residents or migratory (but have already arrived from migration).