Habitat–performance relationships of a large mammal on a predator‐free island dominated by humans

The demographic consequences of changes in habitat use driven by human modification of landscape, and/or changes in climate, are important for any species. We investigated habitat–performance relationships in a declining island population of a large mammal, the moose (Alces alces), in an environment that is predator‐free but dominated by humans. We used a combination of demographic data, knowledge of habitat selection, and multiannual movement data of female moose (n = 17) to understand how space use patterns affect fecundity and calf survival. The calving rate was 0.64 and was similar to calving rates reported in other populations. Calf survival was 0.22 (annually) and 0.32 (postsummer), which are particularly low compared to other populations where postsummer survival is typically above 0.7. Home ranges were mainly composed of arable land (>40%), and selection for arable land was higher in winter than in summer, which contrasts with previous studies. Females that spent more time in broadleaf forest in the summer prior to the rut had higher fecundity rates, while more time spent in arable land resulted in lower fecundity rates. Females that spent more time in thicket/scrubland habitats during winter had lower calf survival, while females that had higher use of mixed forests tended to have higher calf survival. The dominance, and subsequent use, of suboptimal foraging habitats may lead to poor body condition of females at parturition, which may lower calf body weights and affect the mother's ability to lactate. In addition, our results indicated that the growing season has advanced significantly in recent decades, which may be causing a mismatch between parturition and optimal resource availability. These effects may exacerbate the female's ability to meet the energetic demands of lactation. Therefore, the observed low calf survival appears to be caused by a combination of factors related to current land use and may also be due to changing vegetation phenology. These results have important implications for the management of species in human‐dominated landscapes in the face of climate change, and for an increased understanding of how species may adapt to future land use and climate change.

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title: Mass‐ and Density‐Dependent Reproductive Success and Reproductive Costs in a Capital Breeder
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title: Capital and income breeding as alternative tactics of resource use in reproduction
abstract: In order to compensate for the resource demands of reproduction, organisms usually increase the amount of total food resources available. This may be achieved by different tactics of resource use that also include foraging decisions. Two such general tactics are discussed in this paper under the concepts of capital and income breeding. These are defined mainly from the temporal distribution of resource acquisition relative to resource use. A capital breeder acquires its resources in advance and store them endogenously or exogenously until they are needed to supply aspects of offspring production. An income breeder, on the other hand, adjusts its food intake concurrently with breeding, without reliance on stores. In a perfectly predictable environment without limited resources, income breeding is the best option since capital breeders may have to pay a number of energetic and demographic costs for their stored resources. However, under unpredictable food conditions, food/time limitations, and risky foraging conditions, capital breeding offers many benefits. The costing systems (pre- or postbreeding costs) induced by capital and income breeding will largely influence the opportunity for these energetic tactics to evolve. While capital and income breeders may potentially experience both pre- and postbreeding costs, capital breeders may be more exposed to prebreeding costs due to predation in connection with acquisition and carriage of stores.

title: Capital breeding and income breeding: their meaning, measurement, and worth.
abstract: The life histories of organisms can show pronounced variation in the way that the costs of reproduction are financed. To facilitate discussions of this variation, two terms are increasing in use: "capital breeding" describes the situation in which reproduction is financed using stored capital; "income breeding" refers to the use of concurrent intake to pay for a reproductive attempt. We consider the value of the capital and income typology with reference to three functions that it might serve: description, explanation, and prediction. We find that interpretations of the terms have diversified and lack rigidity, leading to subjectivity in their definition. We recognize that time frames of interest will vary among taxa, but we urge consistency of use within those taxa. We also urge consistency in the use of a single metric designed to measure the reliance of an organism on capital. The concepts of capital and income breeding have served well as proximate explanations for behavioral or physiological diversity, but efforts to explain their adaptive value have been disproportionately focused on individual taxa. Mapping cause to effect is difficult in ecology. Nevertheless, further analyses, based on consistently applied measures of reliance on stored capital, may reveal which of the putative ecological, morphological, and physiological drivers have the most consistent and widespread effects. The capital-income typology has yet to be applied to the question of prediction, and thus, it remains to be seen whether these concepts will be of use in identifying the likely responses of different populations to changes in their environment.

title: Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose
abstract: A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.

title: Life history patterns in female moose (Alces alces): the relationship between age, body size, fecundity and environmental conditions
abstract: I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (≥2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.

title: Body mass and individual fitness in female ungulates: bigger is not always better
abstract: In female vertebrates, differences in fitness often correspond to differences in phenotypic quality, suggesting that larger females have greater fitness. Variation in individual fitness can result from variation in life span and/or variation in yearly reproductive success, but no study has yet assessed the relationships between the components of fitness and phenotypic quality while controlling for life span. We tried to fill this gap using data from long–term monitoring (23 years) of marked roe deer and bighorn sheep, two ungulates with very different life histories. In both species, we found a strong positive relationship between an adult female's mass and her probability of reaching old age: over the long term, bigger is indeed better for ungulate females. On the other hand, we found no evidence in either species that heavier females had higher fitness when differences in life span were accounted for: over the short term, bigger is not necessarily better. Our results indicate that, while broad differences in phenotypic quality affect individual fitness, when differences in life span are accounted for phenotypic quality has no residual effect on fitness. Therefore, within a given range of phenotypic quality, bigger is not always better, for reasons which may differ between species.

title: Factors affecting maternal care in an income breeder, the European roe deer
abstract: 1. Maternal care is a major component of demographic tactics in mammals. In ungulates most work has been done on capital breeders (e.g. bighorn sheep), which rely heavily on body reserves to raise their young. Roe deer, in contrast, are close to the income breeder end of the capital–income breeder continuum, and show high levels of maternal care. 
 
 
 
2. The aim of this study was to explore the factors determining the level of maternal care in roe deer, in particular the effects of maternal body weight, mother's parity, litter size and year of birth on the amount of prenatal care (i.e. the average mass of an offspring multiplied by the number of offspring) and postnatal care (i.e. the average growth rate of the offspring multiplied by their number). The study was carried out on a captive population of roe deer fed ad libitum, and in a wild population. 
 
 
 
3. In both populations prenatal care increased with increasing maternal body weight. In the population fed ad libitum this effect was found in light females only (< 22 kg); in the wild population the positive relationship between maternal body weight and prenatal care was entirely accounted for by variation in litter size (i.e. heaviest females produced larger litters) and density (i.e. females were lightest in years with high population density). Parity did not affect prenatal care. 
 
 
 
4. In 14 females fed ad libitum there was no relationship between postnatal care and maternal body weight. Multiparous females had higher levels of postnatal care. In contrast, 20 wild females showed a positive relationship between postnatal care and maternal body weight, and only litter size affected the level of postnatal care. 
 
 
 
5. Even after accounting for the confounding effects of maternal body weight, parity, litter size and population density, we found no trade-off between pre- and postnatal care in any of the two populations. The conditions under which the pattern of maternal care could impose trade-offs that affect the individual offspring are discussed.

title: Experimental evidence of a risk-sensitive reproductive allocation in a long-lived mammal.
abstract: When reproduction competes with the amount of resources available for survival during an unpredictable nonbreeding season, individuals should adopt a risk-sensitive regulation of their reproductive allocation. We tested this hypothesis on female reindeer (Rangifer tarandus), which face a trade-off between reproduction and acquisition of body reserves during spring and summer, with autumn body mass functioning as insurance against stochastic winter climatic severity. The study was conducted in a population consisting of two herds: one that received supplementary winter feeding for four years while the other utilized natural pastures. The females receiving additional forage allocated more to their calves. Experimental translocation of females between the herds was conducted to simulate two contrasting rapid alterations of winter conditions. When females receiving supplementary feeding were moved to natural pastures, they promptly reduced their reproductive allocation the following summer. However, when winter conditions were improved, females were reluctant to increase their reproductive allocation. This asymmetric response to improved vs. reduced winter conditions is consistent with a risk-averse adjustment in reproductive allocation. The ability of individuals to track their environment and the concordant risk-sensitive adjustment of reproductive allocation may render subarctic reindeer more resilient to climate change than previously supposed.

title: Population dynamics of large herbivores: variable recruitment with constant adult survival.
abstract: Recent long-term studies of population ecology of large herbivorous mammals suggest that survival of prime-aged females varies little from year to year and across populations. Juvenile survival, on the other hand, varies considerably from year to year. The pattern of high and stable adult survival and variable juvenile survival is observed in contrasting environments, independently of the main proximal sources of mortality and regardless of whether mortality is stochastic or density-dependent. High yearly variability in juvenile survival may play a predominant role in population dynamics.

title: Nutrition integrates environmental responses of ungulates.
abstract: 1Nutrition influences most aspects of animal ecology: juvenile growth rates and adult mass gain, body condition, probability of pregnancy, over-winter survival, timing of parturition, and neonatal birth mass and survival. We provide an overview among ungulates of the extent of these influences resulting from interactions among bioenergetics, foraging, and nutritional demands. 
2 Body condition of an animal is the integrator of nutritional intake and demands, affecting both survival and reproduction. The deposition and mobilization of body fat and body protein vary with physiological requirements and environmental conditions as species use dietary income and body stores to integrate the profits of summer and the demands of winter. Results from our simulation model and uncertainty analysis of the influence of body mass and changes in body composition of Rangifer over winter indicate that percent body fat rather than body mass in early winter is most important in determining whether animals die, live without reproducing, or live and reproduce. Animal responses are also sensitive to rates of change in body protein. Depending on timing of calving and maternal reserves, seasonal habitats vary in their nutritional value for the production of offspring. 
3For free-ranging animals, life is a balance among numerous ecological factors, including nutritional requirements, nutritional resources to meet those demands, and intra- and inter-specific interactions. Predation effects on population demography may mask nutritional limitations of habitat. We suggest that over the long term of life histories, ungulates use seasonal strategies that minimize the maximum detriment, and that the basis for most strategies is nutritional.

title: Timing and Synchrony of Parturition in Alaskan Moose: Long-Term Versus Proximal Effects of Climate
abstract: We studied timing and synchrony of parturition in Alaskan moose ( Alces alces ) in Denali National Park and Preserve, Alaska, from 1990 to 1994. Mean date of birth was 25 May ( SD = 5.7 days) and did not differ significantly among years. Although moose did not congregate to give birth, parturition was highly synchronized (95% of births occurred in 16 days) with no significant differences among years. Most young moose were killed by predators, especially grizzly bears ( Ursus arctos ), but timing of reproduction had no effect on survivorship of young, which was low (ca. 0.2 by 16 June). We reject the hypothesis that moose timed births to avoid predation. We also measured depth of snow in winter, and precipitation, cloud cover, and temperature in spring; none of these variables was related to timing or synchrony of births. Likewise, quality of willow ( Salix pulchra ) in 3 springs in which percent nitrogen and in vitro dry-matter digestibility of this important forage varied markedly was unrelated to timing of births. We reject the hypothesis that moose timed parturition in response to proximal changes in their environment. We hypothesize that both timing and synchrony of parturition in moose are adaptations to long-term patterns of climate that provide the most hospitable conditions to bear and rear young. Consequently, moose may be more susceptible to climatic change than other ungulates that are more adapted to climatic variability.

title: Effects of Predator Treatments, Individual Traits, and Environment on Moose Survival in Alaska
abstract: We studied moose (Alces alces) survival, physical condition, and abundance in a 3-predator system in western Interior Alaska, USA, during 2001–2007. Our objective was to quantify the effects of predator treatments on moose population dynamics by investigating changes in survival while evaluating the contribution of potentially confounding covariates. In May 2003 and 2004, we reduced black bear (Ursus americanus) and brown bear (U. arctos) numbers by translocating bears ≥240 km from the study area. Aircraft-assisted take reduced wolf (Canis lupus) numbers markedly in the study area during 2004–2007. We estimated black bears were reduced by approximately 96% by June 2004 and recovered to within 27% of untreated numbers by May 2007. Brown bears were reduced approximately 50% by June 2004. Late-winter wolf numbers were reduced by 75% by 2005 and likely remained at these levels through 2007. In addition to predator treatments, moose hunting closures during 2004–2007 reduced harvests of male moose by 60% in the study area. Predator treatments resulted in increased calf survival rates during summer (primarily from reduced black bear predation) and autumn (primarily from reduced wolf predation). Predator treatments had little influence on survival of moose calves during winter; instead, calf survival was influenced by snow depth and possibly temperature. Increased survival of moose calves during summer and autumn combined with relatively constant winter survival in most years led to a corresponding increase in annual survival of calves following predator treatments. Nonpredation mortalities of calves increased following predator treatments; however, this increase provided little compensation to the decrease in predation mortalities resulting from treatments. Thus, predator-induced calf mortality was primarily additive. Summer survival of moose calves was positively related to calf mass (&bgr; > 0.07, SE = 0.073) during treated years and lower (&bgr; = -0.82, SE = 0.247) for twins than singletons during all years. Following predator treatments, survival of yearling moose increased 8.7% for females and 21.4% for males during summer and 2.2% for females and 15.6% for males during autumn. Annual survival of adult (≥2 yr old) female moose also increased in treated years and was negatively (&bgr; = -0.21, SE = 0.078) related to age. Moose density increased 45%, from 0.38 moose/ km2 in 2001 to 0.55 moose/km2 in 2007, which resulted from annual increases in overall survival of moose, not increases in reproductive rates. Indices of nutritional status remained constant throughout our study despite increased moose density. This information can be used by wildlife managers and policymakers to better understand the outcomes of predator treatments in Alaska and similar environments.

title: Identifying when weather influences life-history traits of grazing herbivores.
abstract: 1. There is increasing evidence that density-independent weather effects influence life-history traits and hence the dynamics of populations of animals. Here, we present a novel statistical approach to estimate when such influences are strongest. The method is demonstrated by analyses investigating the timing of the influence of weather on the birth weight of sheep and deer. 2. The statistical technique allowed for the pattern of temporal correlation in the weather data enabling the effects of weather in many fine-scale time intervals to be investigated simultaneously. Thus, while previous studies have typically considered weather averaged across a single broad time interval during pregnancy, our approach enabled examination simultaneously of the relationships with weekly and fortnightly averages throughout the whole of pregnancy. 3. We detected a positive effect of temperature on the birth weight of deer, which is strongest in late pregnancy (mid-March to mid-April), and a negative effect of rainfall on the birthweight of sheep, which is strongest during mid-pregnancy (late January to early February). The possible mechanisms underlying these weather-birth weight relationships are discussed. 4. This study enhances our insight into the pattern of the timing of influence of weather on early development. The method is of much more general application and could provide valuable insights in other areas of ecology in which sequences of intercorrelated explanatory variables have been collected in space or in time.

title: EFFECTS OF SUMMER‐AUTUMN NUTRITION AND PARTURITION DATE ON REPRODUCTION AND SURVIVAL OF ELK
abstract: : Recent declines in numbers and juvenile recruitment in many elk (Cervus elaphus) herds in the western U.S. has sparked interest in factors that may cause these declines. Inadequate nutrition or delayed parturition, the latter of which may be caused by inadequate numbers of mature bulls (i.e., highly skewed sex ratios), may have separate or synergistic effects on population dynamics and productivity. We evaluated the implications of late parturition and summer-autumn nutrition on reproduction and survival of Rocky Mountain elk (C. e. nelsoni) using a captive herd of 57 cow elk. 
 
 
 
We induced early (Sep) and late breeding (Oct) and 3 levels of summer-autumn nutrition on the cows. Food was offered ad libitum at 3 levels of digestible energy (DE): high = 2.9-3.0 kcal of DE/g of diets, medium = 2.6-3.0 kcal/g, and low = 2.3-3.0 kcal/g. Within these ranges, DE content was gradually reduced from late June through early November to mimic seasonal changes in the wild. During summer and autumn, we measured calf growth; body mass, nutritional condition, and breeding dynamics of cows; and growth and pregnancy of yearlings. We also measured carry-over (i.e., time-lag) responses including over-winter calf and cow survival and parturition date and birth mass, as functions of previous summer-autumn nutrition and previous parturition date. Between autumn 1995 and spring 1998, we conducted 2 years of parturition-date, summer-autumn nutrition experiments, 2 winters of calf survival experiments, and 1 winter of cow survival experiments. 
 
 
 
Early birth provided calves with more time to grow before onset of winter. This “head-start” advantage was maintained through late autumn, but its magnitude was diluted in some instances due to faster growth of some late-born calves. Body mass, body fat, and timing and probability of conception by cows in autumn were little influenced by parturition date the previous spring. 
 
 
 
Summer-autumn nutrition significantly affected calves and their mothers. Growth of calves in the low and medium nutrition groups ceased by mid-September and late October. By December, calves in the high nutrition group were 40% and 70% heavier than calves in the medium and low groups, respectively. Cows in the high nutrition group accumulated about 75% and 300% more fat than cows in the medium and low groups by mid-October. Eighty percent of cows in the low nutrition group failed to conceive, and those in the medium group bred 10–14 days later than cows in the high group. Summer-autumn nutrition of calves influenced their probability of becoming pregnant as yearlings. Probability of pregnancy approached 100% for those yearlings that had high summerautumn nutrition as calves and yearlings, despite near starvation their first winter of life. 
 
 
 
Winter survival of calves was related to their size at the onset of winter. Smaller calves lost more body mass daily than did large calves, and thus they survived fewer days through winter. Summer-autumn nutrition largely determined calf body size at the start of winter and, consequently, determined the proportion of winter survived. Survival of cows over winter was as related to body fat at the onset of winter as it was to nutrition during winter. 
 
 
 
Carry-over effects of summer-autumn nutrition and parturition date on birth characteristics the following spring were minor. We detected no significant carry-over effect of summer-autumn nutrition or autumn condition on birth mass, although reduced condition in autumn delayed subsequent parturition date. Extent of body fat depletion in cows during the winter-survival experiments in 1998 accounted for 45% of the variation in parturition date. Ninety percent depletion delayed parturition an average of 34 days. 
 
 
 
Delayed parturition, of a magnitude expected due to highly skewed sex ratios (3 weeks under extreme conditions), probably has only a weak influence on vital rates of free-ranging elk. In contrast, fat accretion and probability of pregnancy of cows, and growth and overwinter survival of calves, were sensitive to small (10–20%) differences in DE content of food. Digestible energy levels of our 2 lower nutrition levels reflect DE ranges reported for large ungulate herds during summer and autumn in western North America. Thus, our data suggest that limiting effects of summer-autumn nutrition on populations may be greater than often assumed, perhaps greater than those during winter in some ecosystems, and consequently indicate a need for greater understanding of nutrition's influence on population dynamics and how this influence varies across space and time. To enhance future research, we present animal- and vegetation-based guidelines for evaluating nutritional influences on elk populations.

title: Cohort effects and deer population dynamics
abstract: Among-individual variation of life history traits in a given population of deer has most often been accounted for by differences among sex and age classes and by current environmental variation (mostly changes in density and climatic conditions). However, among-individual differences in fitness can also be generated by differences in environmental conditions during the year of birth. Such cohort effects can be divided into two different components. First, among-year differences in environmental conditions at birth may produce large yearly variation in recruitment that generates a direct numerical effect (i.e., a high proportion of newborns are recruited in good years, whereas very low proportions are recruited when environmental conditions encountered by newborns during their first weeks of life are poor). Second, when recruited into the population, individuals born in a good year may reproduce earlier, reach a larger body mass, and have a higher reproductive success than individuals born in a poor year. We call the long-lasting influence on individual fitness of environmental conditions during the year of birth a delayed quality effect. Here, we first review briefly evidence of numerical and quality effects recently accumulated in deer populations. Then, by using long-term monitoring (> 20 y) of two contrasted roe deer (Capreolus capreolus) populations, we assess the influence of the among-cohort differences in reproductive traits, age- and sex-specific survival rates, and population growth rate. Results show that cohort effects (1) are widespread in deer populations and (2) may have a major influence on population dynamics, especially in low-performance populations. Cohort effects should therefore be accounted for in management plans of deer populations.

title: Early development and fitness in birds and mammals.
abstract: Conditions experienced during early development affect survival and reproductive performance in many bird and mammal species. Factors affecting early development can therefore have an important influence both on the optimization of life histories and on population dynamics. The understanding of these evolutionary and dynamic consequences is just starting to emerge.

title: Capital and income breeding traits differentiate trophic match–mismatch dynamics in large herbivores
abstract: For some species, climate change has altered environmental conditions away from those in which life-history strategies evolved. In such cases, if adaptation does not keep pace with these changes, existing life-history strategies may become maladaptive and lead to population declines. We use life-history theory, with a specific emphasis on breeding strategies, in the context of the trophic match–mismatch framework to form generalizable hypotheses about population-level consumer responses to climate-driven perturbations in resource availability. We first characterize the income and breeding traits of sympatric caribou and muskoxen populations in western Greenland, and then test trait-based hypotheses about the expected reproductive performance of each population during a period of high resource variability at that site. The immediate reproductive performance of income breeding caribou decreased with trophic mismatch. In contrast, capital breeding muskoxen were relatively unaffected by current breeding season resource variability, but their reproductive performance was sensitive to resource conditions from previous years. These responses matched our expectations about how capital and income breeding strategies should influence population susceptibility to phenological mismatch. We argue for a taxon-independent assessment of trophic mismatch vulnerability based on a life-history strategy perspective in the context of prevailing environmental conditions.

title: Warming, plant phenology and the spatial dimension of trophic mismatch for large herbivores
abstract: Temporal advancement of resource availability by warming in seasonal environments can reduce reproductive success of vertebrates if their own reproductive phenology does not also advance with warming. Indirect evidence from large-scale analyses suggests, however, that migratory vertebrates might compensate for this by tracking phenological variation across landscapes. Results from our two-year warming experiment combined with seven years of observations of plant phenology and offspring production by caribou (Rangifer tarandus) in Greenland, however, contradict evidence from large-scale analyses. At spatial scales relevant to the foraging horizon of individual herbivores, spatial variability in plant phenology was reduced—not increased—by both experimental and observed warming. Concurrently, offspring production by female caribou declined with reductions in spatial variability in plant phenology. By highlighting the spatial dimension of trophic mismatch, these results reveal heretofore unexpected adverse consequences of climatic warming for herbivore population ecology.

title: 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.

title: Climate and population density induce long‐term cohort variation in a northern ungulate
abstract: Density-dependent and climatic conditions experienced by individuals before and after birth differ considerably between cohorts. Such early environmental variability has the potential to create persistent fitness differences among cohorts. Here we test the hypothesis that conditions experienced by individuals in their early development will have long-term effects on their life history traits. 2. We approached this by analysing and contrasting the effects of climate (the North Atlantic Oscillation, NAO) and population density at year of birth on cohort birth weight, birth date, litter size, age of maturity, survival and fecundity of Soay sheep, Ovies aries L., ewes in the population on the island of Hirta, St Kilda, Scotland. 3. Significant intercohort variations were found in life history traits. Cohorts born after warm, wet and windy (high NAO) winters were lighter at birth, born earlier, less likely to have a twin and matured later than cohorts born following cold and dry (low NAO) winters. High population densities in the winter preceding birth also had a negative effect on birth weight, birth date and litter size, whereas high postnatal densities delayed age of first reproduction. 4. High NAO winters preceding birth depressed juvenile survival but increased adult survival and fecundity. The negative influence of high NAO winters on juvenile survival is likely to be related to mothers’ compromised physical condition while the cohort is in utero , whereas the positive influence on adult survival and fecundity may relate to the improved postnatal forage conditions following high NAO winters. High preand postnatal population densities decreased juvenile (neonatal, yearling) and adult (2‐4 years) survivorship but had no significant effect fecundity.

title: Effects of climate and plant phenology on recruitment of moose at the southern extent of their range
abstract: Climate plays a fundamental role in limiting the range of a species, is a key factor in the dynamics of large herbivores, and is thought to be involved in declines of moose populations in recent decades. We examined effects of climate and growing-season phenology on recruitment (8–9 months old) of young Shiras moose (Alces alces shirasi) over three decades, from 18 herds, across a large geographic area encompassing much of the southern extent of their range. Recruitment declined in 8 of 18 herds during 1980–2009, whereas others did not exhibit a temporal trend (none showed a positive trend). During those three decades, seasonal temperatures increased, spring–summer precipitation decreased, and spring occurred earlier, became shorter in duration, and green-up occurred faster. Recruitment was influenced negatively by warm temperatures during the year before young were born, but only for herds with declining recruitment. Dry spring–summers of the previous year and rapid rates of spring green-up in the year of birth had similar negative influences across declining and stable herds. Those patterns indicate both direct (yeart) and delayed (yeart−1) effects of weather and plant phenology on recruitment of young, which we hypothesize was mediated through effects on maternal nutritional condition. Suppressed nutrition could have been induced by (1) increased thermoregulatory costs associated with warming temperatures and (2) shortened duration of availability of high-quality forage in spring. Progressive reductions in net energetic gain for species that are sensitive to climate may continue to hamper individual fitness and population dynamics.

title: Reproduction and health of moose in southern Sweden
abstract: Moose (Alces alces) is a highly regarded game species in Fennoscandia, where annual harvest numbers in Sweden, Norway, and Finland together exceed 200,000 animals. For successful management, knowledge about male and female reproduction is essential, as well as the extent to which disease and mortality affect the population. In 2006, a sub-normal reproductive output (calf per cow ratio) was reported from the island of Oland, and a pilot study in 2007 revealed embryonic mortality and occurrence of the tick-borne pathogen Anaplasma phagocytophilum. An expansion of the study (including control areas) was conducted due to the need for updated information on moose reproduction 
 
From 2008 to 2011, reproductive organs, blood, spleens, mandibles, and ectoparasites were collected from moose in three areas in southern Sweden. Reproductive organs were inspected macroscopically, weighed and measured, and 
sperm samples were taken. Morphology of spermatozoa, chromatin analyses, histological examinations, and pathogen analyses were performed at SLU or SVA in Uppsala. 
 
Male pubertal age varied from 1.5 to 3.5 years, and the proportion of normal spermatozoa increased significantly with increasing body weight, but decreased temporally over the first month of hunting. Male moose had a low testes:body weight ratio compared with other cervids. Cows showed their first oestrus of the season earlier than heifers, and the hunting period appeared to interfere with oestrus in all females. Onset of puberty in females was positively associated with body weight but not with age. Embryonic mortality and unfertilized oocytes accounted for a significant difference (P<0.01) between ovulation rates and the proportion of viable embryos found in pregnant females. Moose were competent hosts of Anaplasma 
phagocytophilum, and the prevalence of infection, as determined by PCR, varied both temporally and spatially. Moose calf summer survival rates on Oland were significantly 
lower than in the mainland populations. 
 
The studies performed provide updated information on moose reproductive 
characteristics, calf survival and moose health. Some changes in population management could potentially improve the reproductive success of moose in southern Sweden. Not all of these parameters might be affected by a change in management, as the surrounding environment and climate play a considerable role in forage availability, the spread of diseases, and calf survival rates.

title: AGE-RELATED REPRODUCTIVE EFFORT AND SENESCENCE IN FREE-RANGING MOOSE, ALCES ALCES
abstract: A general prediction for aging placental mammals is that they should show a decline in reproductive value, and reproductive effort is therefore predicted to increase with age. To test this, we monitored known-age radio-collared females of a large ungulate, the moose (Alces alces), in a Swedish population. Female moose showed senescence in reproduction (i.e., litter size) from about 12 yr of age. Further evidence of senescence was a decrease in parental care during summer (expressed as increased offspring mortality) with the mother's age. Moreover, aging females facing a declining reproductive value increased their reproductive effort by giving birth to heavier offspring regardless of litter size. A logistic regression model showed that older moose have to give birth to heavier offspring to achieve the same offspring summer survival as that for offspring of younger mothers. This suggests that females increase their reproductive effort (by increasing offspring mass) as they get older. Furthermore, the rather intense harvest in our population may select for an increased reproductive effort in terms of litter size, which furthermore may select for an earlier onset of senescence in reproductive characters and mortality.

title: Life-history consequences of maternal condition in Alaskan moose
abstract: We studied life-history characteristics of Alaskan moose (Alces alces gigas) including the effects of maternal condition of adult females (>33 months old) on sunival and physical condition of young during their first year-of-life. We also examined the relation behveen maternal condition and reproductive parameters of individual adult moose, and tested for effects of those parameters on timing and synchrony of parturition. \t7e radiotracked adult fernales captured in both March 1996 and 1997 throughout the year with intensive monitoring occurring during spring and early summner. That procedure enabled us to capture the offspring of females xire monitored and record other variables related to reproductive success. Females with greater nlmp fat thickness had higher rates of pregnancy, gave birth to more hvins. and produced young with higher birth masses than did females with less rump fat. Time-to-death for indi~idual young increased as birth rnass increased and decreased as birth date and litter size increased; those birth variables, lio\ve\~er. did not act upon time-to-death independently. Our results indicated maternal condition influenced subsequent variables associated wit11 birth, which ultimately affected f i~ t~ t r e sunival of offspring. Further, timing of reproduction varied behveen the 2 years, with births occurring earlier but not more synchronously i11 1996 than in 1997. Time of parturition occurred earlier for individual ferr~ales with the thickest rump fat. That outcolne indicated that timing of parturition was the result of e~nironmental factors acting on fe~nales prior to giving birth rather than effects of attempting to avoici predation.

title: Body condition and adjustments to reproductive effort in female moose (Alces alces)
abstract: We studied condition and reproduction of moose ( Alces alces ) in southcentral Alaska using ultrasonography to determine rump-fat thickness and numbers of corpora lutea and embryos in early gestation, and intensive radio-tracking of the same animals in spring to measure rates of calving and neonatal survival. Rump-fat thickness, pregnancy rate, and embryo size were less among female moose accompanied by a calf in autumn. Fifteen percent of ovulations failed to result in a detectable embryo, and additional reproductive losses occurred between early gestation and birth. Body condition in the autumn was correlated positively with pregnancy and calving rates and negatively with reproductive losses in both early and late gestation and neonatal mortality. Our study documents the extent to which body condition and prior reproductive success affect adjustments to reproductive effort made by female moose within a single reproductive cycle.

title: Moose density and habitat productivity affects reproduction, growth and species composition in field layer vegetation.
abstract: What is the effect of a gradient in moose density on reproduction, growth and functional composition of the field layer vegetation in a boreal forest, and how is this effect modified by habitat productivity? 
 
Location: Northwest of Umea, Vasterbotten, northern Sweden. 
 
 Field layer vegetation was surveyed in an experimental setup with simulation of three different moose densities and a control in eight study sites along a gradient of habitat productivity. 
 
 We found that increased moose density led to decreased cover and reproductive effort of a browsed dwarf shrub (bilberry, Vaccinium myrtillus L.) and increased cover and reproductive effort of a non-browsed graminoid (wavy hair-grass, Avenella flexuosa (L.) Drejer). Increased moose density led to increased light availability and probably reduced competition from V. myrtillus. Total reproductive effort in the field layer vegetation increased, height decreased and cover of light-demanding species and graminoids increased with increasing moose density. The effects of moose density were modified by the productivity gradient, leading to a higher relative increase in light availability and reproductive effort in highly productive areas than in low productive areas. 
 
 Increased light availability was an important indirect effect of moose density, leading to less competition for light and a shift towards early successional species. The effect of moose density on light availability was modified by habitat productivity, leading to stronger relative effects in highly productive areas than in low productive areas.

title: Effects of agri-environment schemes on scrub clearance, livestock grazing and plant diversity in a low-intensity farming system on Oland, Sweden
abstract: Changes in land use during the past century in semi-natural landscape caused bush and tree encroachment in many pastures and hay meadows-here exemplified by alvars (limestone grasslands) in the Baltic island of Oland (Sweden). Positive changes have been recorded after introduction of agri-environment schemes and economic support. Increased figures are shown for areas being grazed, for the number of sheep and young cattle as well as for economic support within Morbylanga (Oland) municipality and in Kalmar county, but not that clear for Sweden in total. Reintroduced cattle grazing in a 15-year ungrazed alvar area increased the number of vascular plant species and reduced canopy height, litter thickness and vascular plant cover. The abundance of 12 species out of 72 increased significantly while that of two species decreased. The presence of small. anthills created by Lasius niger compensated for and even increased the number of vascular species in plots ungrazed for 32 years. After clearings in a dense Juniperus communis scrub at least 3-4 years are needed for regeneration of the field- and bottom layers. For the tong-term clearings should be repeated or cattle grazing re-established.

title: Risk of capture-related mortality in large free-ranging mammals: experiences from Scandinavia
abstract: Chemical capture and anaesthesia of free-ranging mammals will always involve some risk of mortality even in healthy animals. Deaths may be directly or indirectly attributable to the anaesthetic event itself (e.g. drug overdose, drowning during induction and dart trauma) or may be caused by secondary effects from the capture (e.g. stress, myopathy, trauma or instrumentation with radio-transmitters). In long-term research projects on five major wildlife species in Scandinavia, the capture-related mortality rates (number of captures) were: moose Alces alces 0.7% (N = 2,816), brown bears Ursus arctos 0.9% (N = 1,079), wolverines Gulo gulo 2.8% (N = 461), Eurasian lynx Lynx lynx 3.9% (N = 380), and gray wolves Canis lupus 3.4% (N = 89). We suggest that wildlife professionals should strive for a zero mortality rate but adopt the standard that a mortality rate of > 2% probably should not be accepted in any large mammalian species. This can be achieved by: 1) using an experienced professional capture team, 2) developing and following a capture protocol specific to each species, and 3) requiring that a mortality assessment be undertaken after any capture-related death. This assessment should re-evaluate the capture protocol, including how changes in anaesthetics and methodological approaches could have prevented the mortality.

title: Wireless Remote Animal Monitoring (WRAM) - A new international database e-infrastructure for management and sharing of telemetry sensor data from fish and wildlife
abstract: New tracking technologies have become available to ecologists, allowing remote real-time data capture from an increasing number of taxa, species and animals. To realize the potential of the data, researchers must be able to share data and collaborate with the global research community. The Wireless Remote Animal Monitoring (WRAM) database system, started in 2003, contains 96.5 million positions and other sensor data and is used to date by 32 user groups from 8 countries, tracking 16 species and 2 179 individual animals. The infrastructure represents the Swedish national data node and data sharing portal for real-time telemetry sensor data from fish and wildlife. WRAM will be part of Swedish LifeWatch and is cooperating with several other international database initiatives. The infrastructure consists of 2 main parts: 1) The WRAM Data Warehouse (WDW) is a high performance data warehouse for real-time 'big data' as position, acceleration, or heartbeat data from fish and wildlife, hosted by the High Performance Computing Center North (HPC2N) in Umea, Sweden. 2) The WRAM Data Broker (WDB) is the single-sign-on web interface federating the WDW with other similar database systems around the world as Movebank, EuroDeer or CAnMove to enable seamless querying across systems and easy data sharing between data owners, while honoring local authentication and authorization settings. Query results are also accessible through ODBC in a temporary database, enabling users to use local analysis tools for further analyses. Here, we give an overview over the different parts of the e-infrastructure, including automated data capture tools, database models, data sharing approaches, and web-based visualization tools.

title: Accuracy and repeatability of moose (Alces alces) age as estimated from dental cement layers
abstract: Optimal research and management of species with age structure often depends on estimates of age-specific population parameters, which in turn depends on reliable methods for age determination. By counting annuli in the cementum of incisor root tips from 51 known-age moose (Alces alces) between 1 and 12 years old, we examined the variation in accuracy and repeatability of age estimates provided by three research technicians with different experiences of aging moose. The most experienced technician estimated the moose age correctly in up to 90% of the cases, while the technician with no prior experience estimated age correctly in up to 73% of the cases. The medium-experienced technician achieved a lower accuracy (up to 53%), indicating that experience alone is not sufficient if the basic training or lack of routine checks against other colleagues or a known-age material are not undertaken. The percentage moose aged within ±1 year from correct age was significantly higher in all technicians (94–98%). After the generally high accuracy, we also found high repeatability (0.980–0.994) within technicians. We conclude that this method of age-determination provides reliable estimates that can be used by management and research to gain information about age-specific patterns in moose populations. However, to obtain estimates of high accuracy the technicians should be well trained, have gained the necessary experience, and most preferably, have access to a sample of teeth from known-age moose.

title: Quantifying Migration Behaviour Using Net Squared Displacement Approach: Clarifications and Caveats
abstract: Estimating migration parameters of individuals and populations is vital for their conservation and management. Studies on animal movements and migration often depend upon location data from tracked animals and it is important that such data are appropriately analyzed for reliable estimates of migration and effective management of moving animals. The Net Squared Displacement (NSD) approach for modelling animal movement is being increasingly used as it can objectively quantify migration characteristics and separate different types of movements from migration. However, the ability of NSD to properly classify the movement patterns of individuals has been criticized and issues related to study design arise with respect to starting locations of the data/animals, data sampling regime and extent of movement of species. We address the issues raised over NSD using tracking data from 319 moose (Alces alces) in Sweden. Moose is an ideal species to test this approach, as it can be sedentary, nomadic, dispersing or migratory and individuals vary in their extent, timing and duration of migration. We propose a two-step process of using the NSD approach by first classifying movement modes using mean squared displacement (MSD) instead of NSD and then estimating the extent, duration and timing of migration using NSD. We show that the NSD approach is robust to the choice of starting dates except when the start date occurs during the migratory phase. We also show that the starting location of the animal has a marginal influence on the correct quantification of migration characteristics. The number of locations per day (1–48) did not significantly affect the performance of non-linear mixed effects models, which correctly distinguished migration from other movement types, however, high-resolution data had a significant negative influence on estimates for the timing of migrations. The extent of movement, however, had an effect on the classification of movements, and individuals undertaking short- distance migrations can be misclassified as other movements such as sedentary or nomadic. Our study raises important considerations for designing, analysing and interpreting movement ecology studies, and how these should be determined by the biology of the species and the ecological and conservation questions in focus.

title: Dynamic Approach to Space and Habitat Use Based on Biased Random Bridges
abstract: Although habitat use reflects a dynamic process, most studies assess habitat use statically as if an animal's successively recorded locations reflected a point rather than a movement process. By relying on the activity time between successive locations instead of the local density of individual locations, movement-based methods can substantially improve the biological relevance of utilization distribution (UD) estimates (i.e. the relative frequencies with which an animal uses the various areas of its home range, HR). One such method rests on Brownian bridges (BBs). Its theoretical foundation (purely and constantly diffusive movements) is paradoxically inconsistent with both HR settlement and habitat selection. An alternative involves movement-based kernel density estimation (MKDE) through location interpolation, which may be applied to various movement behaviours but lacks a sound theoretical basis.  I introduce the concept of a biased random (advective-diffusive) bridge (BRB) and show that the MKDE method is a practical means to estimate UDs based on simplified (isotropically diffusive) BRBs. The equation governing BRBs is constrained by the maximum delay between successive relocations warranting constant within-bridge advection (allowed to vary between bridges) but remains otherwise similar to the BB equation. Despite its theoretical inconsistencies, the BB method can therefore be applied to animals that regularly reorientate within their HRs and adapt their movements to the habitats crossed, provided that they were relocated with a high enough frequency.  Biased random walks can approximate various movement types at short times from a given relocation. Their simplified form constitutes an effective trade-off between too simple, unrealistic movement models, such as Brownian motion, and more sophisticated and realistic ones, such as biased correlated random walks (BCRWs), which are too complex to yield functional bridges. Relying on simplified BRBs proves to be the most reliable and easily usable way to estimate UDs from serially correlated relocations and raw activity information.

title: The package “adehabitat” for the R software: A tool for the analysis of space and habitat use by animals
abstract: The practical analysis of space use and habitat selection by animals is often a problem due to the lack of well-designed programs. I present here the “adehabitat” package for the R software, which offers basic GIS (Geographic Information System) functions, methods to analyze radio-tracking data and habitat selection by wildlife, and interfaces with other R packages. These tools can be downloaded freely on the internet. Because the functions of this package can be combined with other functions of R, “adehabitat” provides a powerful environment for the analysis of the space and habitat use.

title: QUANTIFYING HOME-RANGE OVERLAP: THE IMPORTANCE OF THE UTILIZATION DISTRIBUTION
abstract: The concept of an animal's home range has evolved over time, as have methods for estimating home-range size and shape. Recently, home-range estimation methods have focused on estimating an animal's utilization distribution (UD; i.e., the probability distribution defining the animal's use of space). We illustrate the importance of the utilization distribution in characterizing the degree of overlap between home ranges (e.g., when assessing site fidelity or space-use sharing among individuals). We compare several different statistics for their ability to accurately rank paired examples in terms of their degree of overlap. These examples illustrate limitations of indices commonly used to quantify home-range overlap and suggest that new overlap indices that are a function of the UD are likely to be more informative. We suggest 2 new statistics for measuring home-range overlap: (1) for a measure of space-use sharing, we suggest a generalization of Hurlbert's (1978) E/Euniform statistic, which we term the utilization distribution overlap index (UDOI), and (2) for a general measure of similarity between UD estimates, we suggest Bhattacharyya's affinity (BA; Bhattacharyya 1943). Using a short simulation study, we found that overlap indices can accurately rank pairs of UDs in terms of the extent of overlap, but estimates of overlap indices are likely to be biased. The extent of the bias depended on sample size and the degree of overlap (UDs with a high degree of overlap resulted in statistics that were more biased [low]), suggesting that comparisons across studies may be problematic. We illustrate the use of overlap indices to quantify the degree of similarity among UD estimates obtained using 2 different data collection methods (Global Positioning Systems [GPS] and very high frequency [VHF] radiotelemetry) for an adult female northern white-tailed deer (Odocoileus virginianus) in north-central Minnesota.

title: Procedure for classification of forests for CORINE land cover in Sweden.
abstract: The EU-CORINE Land Cover (CLC 2000) project for Sweden was carried out by the Swedish National Land Survey. The classification of forested land cover types, covering about 65% percent of the land area in Sweden, was contracted out to the Swedish University of Agricultural Sciences (SLU). The method of forest classification used detailed plot information from the National Forest Inventory (NFI), map masks and Landsat data, and was developed to be automated and robust. Within each Landsat scene an average of 2,000 NFI plots were available to use as field data. One use of the NFI plots was to use them as spectral reference targets for correction of within-scene haze differences and correction of slope effects. The forest classification was made with a self-calibrating maximum likelihood algorithm, using the NFI plots as ground truth. In the classification, prior probabilities were iterated until the frequency of each class in the classification corresponded to the frequency of that class according to the field plots. All pre-processing of the image data and the NFI data, in addition to the classification, was done using the automated production line written in-house called MUNIN. In total three products were produced: the CLC 2000 product with 4 forest classes and 25 ha minimum mapping unit (MMU); and two national products with 7 forest classes and a MMU of 1 ha and 25 m (pixel level), respectively. The seven forest classes in the national product were: clear felled areas; young forests; deciduous forests; mixed forests; coniferous forests 5 to 15 m in height; coniferous forests greater than 15 m in height; and, lichen dominated coniferous forests. Accuracy assessment was carried out using both photo interpretation and cross-validation. The accuracies using cross-validation at the pixel level were on the order of 50-70 %. When aggregated to the four CORINE forest classes, the accuracy increased to about 80 % at the pixel level.

title: Resource selection by animals: statistical design and analysis for field studies.
abstract: to resource selection studies. Examples of the use of resource selectory studies. Examples of the use of resource selection functions. Statistical modelling procedures. Studies with resources defined by several categories. Estimating a resource selection probability function from a census of resource units using logistic regression. Estimating a resource selection probability function from a census of resource units at several points in time using the proportional hazards model. Estimating a resource selection function from samples of resource units using proportional hazards and log-linear models. Estimating a resource selection function from two samples of resource units using logistic regression and discriminant function methods. General log-linear modelling. Analysis of the amount of use. The comparison of selection for different types of resource unit.

title: STUDY DESIGNS AND TESTS FOR COMPARING RESOURCE USE AND AVAILABILITY
abstract: A controlled-release additive composition for use in aqueous systems. The composition comprises a core containing at least one additive component and a polymeric coating encapsulating said core. The controlled released cooling additive composition slowly releases the additive components to aqueous system, thereby delivering an effective concentration level of additive components over an extended period.

title: Scale and heterogeneity in habitat selection by elk in Yellowstone National Park
abstract: Resource selection functions (RSF) can be used to explore the role of scale in determining patterns of habitat use. We estimated RSFs for 93 radiocollared adult female elk (Cervus canadensis) with resource availability defined at four spatial scales and two seasons in Yellowstone National Park. Habitat selection differed markedly among scales and seasonal ranges. During winter elk moved to ranges at lower elevations where snow water equivalents were low and selected landscapes with a mix of forest and open vegetation at all spatial scales. Areas of high vegetation diversity were selected at large spatial scales during summer, whereas elk selected less diverse areas on winter range. During summer elk selected forests that burned 12-14 y earlier, but they used these burns less than expected by chance during winter. Habitat selection by elk occurred at multiple spatial scales; thus, we cannot prescribe a single scale as being best for modelling habitat use by elk. Instead, selection of an appropriate scale will vary depending on the research question or management issue at hand.

title: A MULTISCALE BEHAVIORAL APPROACH TO UNDERSTANDING THE MOVEMENTS OF WOODLAND CARIBOU
abstract: We assessed the response of woodland caribou (Rangifer tarandus caribou) to land-cover type, predation risk, energetic costs of movement, and patch configuration at multiple spatial scales. We applied a nonlinear model to frequent locations collected with Global Positioning System (GPS) collars to identify discontinuities in the scales of movement by caribou found in forested and alpine (above tree line) habitats. We differ- entiated intra- from interpatch movements and identified collections of patches (multiple- patch scale) where caribou concentrated intrapatch movements. On average, intra- and interpatch movements were 450.7 and 1268.8 m, respectively, and multiple-patch move- ments occurred over an area of 182 ha. Intrapatch movements were highly correlated, indicative of a strong relationship between behavior and place. Caribou in the forest selected patches of Pine terrace, whereas caribou in the alpine selected patches of Alpine-little vegetative cover. Predation risk was not a factor influencing movements of caribou at the intrapatch scale. Selection of cover types was more variable during interpatch movements. At that scale, caribou selected patches of Pine terrace, Lakes/rivers, Alpine-little vegetative cover, and Alpine-grass. The routes selected by caribou had lower energetic costs relative to surrounding terrain, and during some winters, caribou were subjected to higher levels of predation risk during those movements. At the multiple-patch scale, selection was more specific and encompassed patches of Alpine-little vegetative cover, Alpine-grass, and Pine terrace. Predation risk was relatively unimportant at the multi-patch scale, but animals that moved from forested to alpine habitats reduced their relative risk of predation. Patch con- figuration was a poor predictor of those areas where caribou concentrated intrapatch move- ments. There was some evidence of caribou selecting patches of Pine terrace within a matrix of Wetlands and Pine-black spruce/black spruce patches. Caribou in the alpine avoided patches of Alpine-little vegetative cover adjacent to forest types. Our results indicate that forest managers should maintain widely distributed patches of Pine terrace and implement silvicultural regimes that do not stimulate predator populations across areas used for in- terpatch movements.

title: Practical guidance on characterizing availability in resource selection functions under a use-availability design.
abstract: Habitat selection is a fundamental aspect of animal ecology, the understanding of which is critical to management and conservation. Global positioning system data from animals allow fine-scale assessments of habitat selection and typically are analyzed in a use availability framework, whereby animal locations are contrasted with random locations (the availability sample). Although most use-availability methods are in fact spatial point process models, they often are fit using logistic regression. This framework offers numerous methodological challenges, for which the literature provides little guidance. Specifically, the size and spatial extent of the availability sample influences coefficient estimates potentially causing interpretational bias. We examined the influence of availability on statistical inference through simulations and analysis of serially correlated mule deer GPS data. Bias in estimates arose from incorrectly assessing and sampling the spatial extent of availability. Spatial autocorrelation in covariates, which is common for landscape characteristics, exacerbated the error in availability sampling leading to increased bias. These results have strong implications for habitat selection analyses using GPS data, which are increasingly prevalent in the literature. We recommend that researchers assess the sensitivity of their results to their availability sample and, where bias is likely, take care with interpretations and use cross validation to assess robustness.

title: THE IMPORTANCE OF INDIVIDUAL VARIATION IN DEFINING HABITAT SELECTION BY MOOSE IN NORTHERN BRITISH COLUMBIA
abstract: Understanding resource use and selection has been central to many studies of ungulate ecology. Global positioning satellite (GPS) collars, remote sensing, and geographic information systems (GIS) now make it easier to examine variation in use and selection by individuals. Resource selection functions, however, are commonly developed for global (all animals pooled) models and important information on individual variability may be lost. Using data from 14 female moose (Alces alces) collared in the Muskwa-Kechika Management Area of northern British Columbia, we examined differences among global and individual resource selection models for 5 seasons (winter, late winter, calving, summer, and fall). The global models indicated that moose selected for mid-elevations, and for deciduous burns and Carex sedge areas in all seasons. Resource selection models for individuals, however, indicated that no individuals selected the same attributes as the global models. We also examined selection ratios among seasons with individual moose as replicates, and within individuals with bootstrapping techniques. We discuss the importance of considering individual variation in defining resource selection and habitat use by moose and contrast the results of selection ratios and resource selection models. We also use these data to illustrate some of the pitfalls that can be encountered using the 2 methodologies.

title: Application of random effects to the study of resource selection by animals.
abstract: 1. Resource selection estimated by logistic regression is used increasingly in studies to identify critical resources for animal populations and to predict species occurrence. 2. Most frequently, individual animals are monitored and pooled to estimate population-level effects without regard to group or individual-level variation. Pooling assumes that both observations and their errors are independent, and resource selection is constant given individual variation in resource availability. 3. Although researchers have identified ways to minimize autocorrelation, variation between individuals caused by differences in selection or available resources, including functional responses in resource selection, have not been well addressed. 4. Here we review random-effects models and their application to resource selection modelling to overcome these common limitations. We present a simple case study of an analysis of resource selection by grizzly bears in the foothills of the Canadian Rocky Mountains with and without random effects. 5. Both categorical and continuous variables in the grizzly bear model differed in interpretation, both in statistical significance and coefficient sign, depending on how a random effect was included. We used a simulation approach to clarify the application of random effects under three common situations for telemetry studies: (a) discrepancies in sample sizes among individuals; (b) differences among individuals in selection where availability is constant; and (c) differences in availability with and without a functional response in resource selection. 6. We found that random intercepts accounted for unbalanced sample designs, and models with random intercepts and coefficients improved model fit given the variation in selection among individuals and functional responses in selection. Our empirical example and simulations demonstrate how including random effects in resource selection models can aid interpretation and address difficult assumptions limiting their generality. This approach will allow researchers to appropriately estimate marginal (population) and conditional (individual) responses, and account for complex grouping, unbalanced sample designs and autocorrelation.

title: Movements of female moose in relation to birth and death of calves.
abstract: We followed the daily movements of24-59 parturient moose every spring from 19941997 in southcentral Alaska, an area with heavy predation of calves. We tested the hypotheses that: ( l) selection of a parturition site involves movements not typical of normal home range use; (2) movements during the neonatal period increase risk to calves; and (3) females are more likely to return to an area for parturition if their previous calf escaped predation. Movements of moose increased significantly in the 2 days prior to parturition, were greatly reduced for at least the next 9 days and did not approach pre-parturition levels until calves were about 26 days old. Daily movements by females that eventually lost a calf(> 48 hours after day observed) were greater than those by females that successfully reared their calf or calves through the first 45 days of life (P = 0.049). Distances between parturition sites in successive years were greater among females that lost their calves the first year ( 4.9 vs 2.6 km, P = 0.02), regardless of the age at which the calf died.  (2000)

title: Nonparametric Estimation from Incomplete Observations
abstract: In lifetesting, medical follow-up, and other fields the observation of the time of occurrence of the event of interest (called a death) may be prevented for some of the items of the sample by the previous occurrence of some other event (called a loss). Losses may be either accidental or controlled, the latter resulting from a decision to terminate certain observations. In either case it is usually assumed in this paper that the lifetime (age at death) is independent of the potential loss time; in practice this assumption deserves careful scrutiny. Despite the resulting incompleteness of the data, it is desired to estimate the proportion P(t) of items in the population whose lifetimes would exceed t (in the absence of such losses), without making any assumption about the form of the function P(t). The observation for each item of a suitable initial event, marking the beginning of its lifetime, is presupposed. For random samples of size N the product-limit (PL) estimate can be defined as follows: L...

title: Costs of reproduction in female moose (Alces alces) as measured by means of phenotypic correlations
abstract: The costs of pregnancy and lactation in terms of subsequent body growth and fecundity were studied by comparing different reproductive categories of Swedish female moose (Alces alces) during 1989-1...

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

title: Food plots as a habitat management tool: forage production and ungulate browsing in adjacent forest
abstract: A key challenge for wildlife management is to handle competing goals. High ungulate densities may be desirable from hunting and recreational perspectives, but may come in conflict with needs to limit or reduce browsing damage. Since browsing intensity is negatively related to forage availability it may be possible to mitigate damage on forest by increasing forage availability within the landscape. A commonly used method to increase the attractiveness of a localized part of the landscape is to establish food plots. In a multiyear setup using enclosures, wildlife observations, field surveys, and controlled biomass removal, we studied food plots to document forage production, utilization by ungulates, and browsing on adjacent forests in southern Sweden. The fenced parts of the food plots produced on average 2230 to 5810 kg ha-1 marrow-stem kale, second-year clover mix or early-sown rapeseed. The biomass of target crops was generally higher within ungrazed (exclosures) compared to grazed (controls) quadrats on the food plots, which demonstrates that the crops were used as forage by ungulates. Browsing on deciduous trees in the adjacent forest was higher within 70–135 m from the food plots compared to areas further away. For wildlife management, our study shows that establishment of food plots provides substantial amounts of forage both during growing season and at the onset of the dormant season, and that a large share of this food is consumed. Finally, our study documents that forage availability for ungulates at the onset of the often-limiting dormant season can be increased by fencing food plots throughout the growing season.

title: Space and habitat use of moose in southwestern Sweden
abstract: Increasingly, wildlife managers and land managers are challenged to maintain the viability of large mammal populations. Although the moose (Alces alces) is an ecologically, economically, and culturally important ungulate species found throughout most of Sweden, little is known about its ecology throughout the southern part of the country. We collected baseline ecological data on spatial and habitat use patterns in urbanized southwestern Sweden by fitting 22 adult moose (13 F, nine M) with global positioning system (GPS) radio collars. Home range size of cows did not differ among seasons (P > 0.10); however, bulls had larger home ranges during fall than all other seasons (P < 0.010). Mean home range size of males during fall and spring was larger than females during any season (P < 0.010). We used a Euclidean distance-based approach to analyze multi-scale habitat selection by moose. Moose of both sexes and during all seasons selected boreal forest and mires when establishing a home range (P < 0.10). Moose had seasonal differences in habitat selection within their home range (P 0.001), and generally selected clear-cuts and early successional forests, mature coniferous forests, and glades, but avoided agricultural areas and open water. Habitat and space use characteristics of moose in our urbanizing study area were similar to characteristics reported for moose in forest-dominated landscapes of Fennoscandia.

title: A note on a general definition of the coefficient of determination
abstract: A generalization of the coefficient of determination R2 to general regression models is discussed. A modification of an earlier definition to allow for discrete models is proposed.

title: Hierarchical partitioning as an interpretative tool in multivariate inference
abstract: Much of biogeography, conservation and evolutionary biology, and ecology involves very large spatial and temporal extents. Direct manipulation to test hypotheses is usually almost impossible at appropriate scales so that multivariate modelling and especially regression are used to draw causal inferences about which ‘independent’ variables influence the distribution and abundances of species. Such inferences clearly are crucial for the successful management of biological resources and for conserving threatened species. A succession of regression approaches has arisen, many of which yield inconsistent implications. The main problem has been the quest for one (the ‘best’ or the ‘optimal’) regression model from which the impacts of independent variables are inferred. This note is to draw the attention of ecologists to a relatively recent method, hierarchical partitioning, that does not aim to identify a best regression model as such but rather uses all models in a regression hierarchy to distinguish those variables that have high independent correlations with the dependent variable. Such variables are likely to be most influential in controlling variation in the dependent variable. Hierarchical partitioning is not to be regarded as a substitute for experimental manipulation when that is appropriate, but it is likely to produce better deductions than common regression approaches in the many ecological situations in which manipulation is impossible or of doubtful value.

title: 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%.

title: Moose (Alces alces) survival in three populations in northern Norway
abstract: We used radio-collared individuals to examine the seasonal, annual, and regional variation in age-specific survival of moose (Alces alces) in three populations (Vega, Beiarn, and Troms) in northern Norway. In the two populations subject to a regular hunt, the annual mortality from hunting was higher, on average, than that from natural mortality for adult cows. In these study areas, the hunting mortality rate was higher for calves than for cows. For both age groups there was significant annual variation in hunting mortality, which was associated with differences in quota size among years. The survival rate of adult cows was high outside the hunting season (96%). There was seasonal variation in survival among calves, the highest significant mortality being found among neonates during summer in Vega and Troms and significantly higher mortality being found during winter in Beiarn. The natural mortality of calves differed significantly among regions during both summer and winter. The combined effects of densit...

title: TEMPORAL PATTERNS IN THE SURVIVAL OF TWIN AND SINGLE MOOSE (ALCES ALCES) CALVES IN SOUTHCENTRAL ALASKA
abstract: We studied survival of 220 calves of radiocollared moose (Alces alces) from parturition to the end of July in southcentral Alaska from 1994 to 1997. Prior studies established that predation by brown bears (Ursus arctos) was the primary cause of mortality of moose calves in the region. Our objectives were to characterize vulnerability of moose calves to predation as influenced by age, date, snow depths, and previous reproductive success of the mother. We also tested the hypothesis that survival of twin moose calves was independent and identical to that of single calves. Survival of moose calves from parturition through July was 0.27 ± 0.03 SE, and their daily rate of mortality declined at a near constant rate with age in that period. Mean annual survival was 0.22 ± 0.03 SE. Previous winter's snow depths or survival of the mother's previous calf was not related to neonatal survival. Selection for early parturition was evidenced in the 4 years of study by a 6.3% increase in the hazard of death with each daily increase in parturition date. Although there was no significant difference in survival of twin and single moose calves, most twins that died disappeared together during the first 15 days after birth and independently thereafter, suggesting that predators usually killed both when encountered up to that age.

title: Living on the Edge: Viability of Moose in Northeastern Minnesota
abstract: North temperate species on the southern edge of their distribution are especially at risk to climate-induced changes. One such species is the moose (Alces alces), whose continental United States distribution is restricted to northern states or northern portions of the Rocky Mountain cordillera. We used a series of matrix models to evaluate the demographic implications of estimated survival and reproduction schedules for a moose population in northeastern Minnesota, USA, between 2002 and 2008. We used data from a telemetry study to calculate adult survival rates and estimated calf survival and fertility of adult females by using results of helicopter surveys. Estimated age- and year-specific survival rates showed a sinusoidal temporal pattern during our study and were lower for younger and old-aged animals. Estimates of annual adult survival (when assumed to be constant for ages >1.7 yr old) ranged from 0.74 to 0.85. Annual calf survival averaged 0.40, and the annual ratio of calves born to radiocollared females averaged 0.78. Point estimates for the finite rate of increase (λ) from yearly matrices ranged from 0.67 to 0.98 during our 6-year study, indicative of a long-term declining population. Assuming each matrix to be equally likely to occur in the future, we estimated a long-term stochastic growth rate of 0.85. Even if heat stress is not responsible for current levels of survival, continuation of this growth rate will ultimately result in a northward shift of the southern edge of moose distribution. Population growth rate, and its uncertainty, was most sensitive to changes in estimated adult survival rates. The relative importance of adult survival to population viability has important implications for harvest of large herbivores and the collection of information on wildlife fertility.

title: Risk-sensitive allocation in seasonal dynamics of fat and protein reserves in a long-lived mammal.
abstract: Body reserves of numerous taxa follow seasonal rhythms that are a function of temporal patterns in food availability and life-history events; however, tests of the theory underlying the allocation of somatic reserves for long-lived organisms are rare, especially for free-ranging mammals. We evaluated the hypothesis that allocation of somatic reserves to survival (i.e., metabolic processes) and reproduction should be sensitive to current nutritional state relative to seasonal thresholds in those reserves. Our goal was to reveal the linkages between nutrition and life-history traits to understand how long-lived, iteroparous organisms balance the allocation of somatic reserves to reproduction, while retaining reserves as insurance for survival in unpredictable environments. Our evaluation was based on seasonal dynamics in fat (measured as ingesta-free body fat; IFBFat) and protein reserves (measured as ingesta-free, fat-free body mass; IFFFBMass) of 136 female mule deer (Odocoileus hemionus) over 8 years. Although mean changes in fat and protein reserves were positive over summer and negative over winter, accretion and catabolism of those reserves was not consistent among individuals. Over winter, both lipid and protein stores available in autumn were catabolized in proportion to their availability above a post-winter threshold (5·8% IFBFat, 33 kg IFFFBMass); however, lean body tissue was spared at the expense of lipid reserves. Female deer mostly synthesized lean body tissue over summer and committed post-winter fat reserves to reproduction relative to their availability above an autumn threshold (>8·6% IFBFat), which was lowered by 2·8 percentage points (pp) for each additional young recruited. Mothers reduced their autumn fat threshold to secure current reproductive investment and, thereby, endured a cost of reproduction at the expense of fat accumulation. Allocation of somatic reserves occurred in a risk-sensitive framework; females allocated reserves relative to their availability above seasonal thresholds. In contrast to current notions of summer accretion and winter catabolism of body reserves, some individuals deposited reserves over winter and catabolized reserves over summer, mainly because regulation of individual condition was state-dependent. Consequently, behaviour and life-history strategies may be as much a function of nutritional contributions of the previous season as of the current one.

title: Opposite responses of body condition and fertility in adjacent moose populations
abstract: Moose (Alces alces) populations exceed 3 individuals/km2 in some wildlife reserves and parks of northeastern Canada. Heavy browsing pressure at such densities is potentially altering the ecological integrity of forests with eventual negative consequences for moose. We hypothesized that regulation by resources would be limited by the capacity of female moose to modulate their reproductive strategies to maintain high fertility despite a decline in body condition. We observed a 20–33% decline in rump fat thickness in males and females, respectively, from the high density Matane Wildlife Reserve in Eastern Quebec in October compared to an adjacent population. We also observed a lower mass of the peroneus group of muscles for males (−3%) and prime-aged females (−8%) in the Matane Wildlife Reserve than in the adjacent population. Females from the Matane Wildlife Reserve population had a lower twinning ovulation rate (1 out of 20 vs. 7 out of 21 ovulating females in the adjacent population) but >15% higher overall ovulation rate. Our results suggest that female moose can maintain high fertility despite a decline in body condition, by reducing their litter size at ovulation and conserving energy to increase the probability of annual reproduction. The adjustment of female reproductive strategies illustrates the plasticity of moose in response to decreasing habitat quality. We conclude that in the absence of specialist predators, equilibrium with forage is unlikely for large herbivores in the short or mid-term. Active management of moose populations is likely required to maintain the ecological integrity of boreal forests.

title: POPULATION DYNAMICS AND LIFE HISTORY TRADE‐OFFS OF MOOSE (ALCES ALCES) IN SOUTH‐CENTRAL ALASKA
abstract: A common framework toward understanding the ecological forces affecting herbivore population dynamics is to partition those forces according to their origin higher (“top-down”) or lower (“bottom-up”) on the trophic chain. I employed that framework to describe phenotypic life history trade-offs in a large herbivore, moose (Alces alces). The relative importance of top-down and bottom-up ecological constraints to female moose in south-central Alaska was appraised at the population level by analyzing population size, growth, and winter habitat of moose, and the size and impact of their principal predator populations (wolves [Canis lupus] and brown bears [Ursus arctos]). At the individual level, longitudinal mortality and reproductive patterns of radio-collared individuals were studied to estimate vital rates and life history trade-offs that could be attributed to nutrition and predation, respectively. The population was essentially stable through the 1980s and early 1990s, and began declining in the period of...

title: Moose Alces alces habitat use at multiple temporal scales in a human-altered landscape
abstract: Habitat alteration by humans may change the supply of food and cover for wild ungulates, but few studies have examined how these resources are utilised over time by individuals of different sex and reproductive status. We examined circadian and seasonal variation in habitat utilisation within a moose Alces alces population in central Norway. Our study area covers forests and open habitats, both influenced by human alterations (e.g. forestry and agriculture). We expected moose to select habitats with good forage and cover in all seasons, but to select open foraging habitats mainly during night-time. Moose selected good foraging habitats, such as young forest stands and cultivated land during night, whereas the utilisation of older forest stands providing cover increased during daytime. This circadian pattern changed throughout the year, seemingly related to variation in hours of daylight and provision of forage. Young forest stands provided higher density of preferred food plants compared to older stands and were highly selected from spring until autumn. Relative to young forest, the selection for older forest stands increased towards winter, likely due to provision of higher plant quality late in the growing season, and to reduced accumulation of movement-impeding snow during winter. Selection of cultivated land varied among seasons, being highest when crop biomass was high. We also found some indications of state-dependent habitat selection as reproducing females avoided open, food rich areas in the first months after their calves were born, whereas males and females without young selected these areas in spring and summer. Our results clearly show that moose exploit the variations in cover and food caused by forestry and agriculture. This is particularly relevant for moose in Norway as current changes in forestry practice lead to a reduction in young, food-rich forest stands, possibly aggravating the already declining body conditions and recruitment rates of moose.

title: Quantitative estimates of tree species selectivity by moose (Alces alces) in a forest landscape
abstract: An extensive literature is available on browsing preference for certain tree species. However, useful predictive tools for estimating the impact of deer on forests production and biodiversity can still be improved. A step in that direction is not only to rank preference among tree species but also to quantify the relative risk of being browsed. The foraging selectivity of moose was evaluated using three different statistical methods developed to study habitat utilization. The general pattern for the three methods was consistent. From the results, groups of forage species were clustered and a quantitative index of selectivity was calculated for the groups. The selectivity index showed that rowan (Sorbus aucuparia), willow (Salix ssp.) and aspen (Populus tremula) had a 14 times higher probability of being browsed than a group consisting of Scots pine (Pinus sylvestris) and downy birch (Betula pubescens), while juniper (Juniperus communis) and silver birch (Betula pendula) had a 3.5 times higher probability than Scots pine and downy birch. Since the most preferred species were the least abundant, one should be cautious about the generality of the index between areas, as it may indicate that preference depends on plant species composition. The method used can easily be applied in forest management. Information on quantitative selectivity indices may improve the possibility of managing moose in accordance with acceptable browsing damage.

title: Diet choices made by free-ranging moose in northern Sweden in relation to plant distribution, chemistry, and morphology
abstract: Understanding how large herbivores select their diet requires accurate measurements of the nutritional costs and benefits of food items and the composition of plants in the diet relative to the composition of plants in the habitat. We followed moose (Alces alces) tracks in the snow and measured bites of browse plants taken and the number and mass of twigs of the 10 browse species available within the moose's reach and measured morphological and chemical characteristics of browse plants. The diets of 41 moose in northern coastal Sweden contained 75% Scots pine (Pinus sylvestris) and willow (Salix spp.), with the remaining proportion equally divided among five deciduous browse species and common juniper (Juniperus communis). The moose used approximately 20% of the bites available along the foraging patch, usually taking only one or two bites on each plant selected for browsing. Moose selected rowan (Sorbus aucaparia), willow species, and aspen (Populus tremula) in winter, used gray alder (Alnus incana), sil...

title: Mineral diversity in wild plants: benefits and bane for moose
abstract: We studied the mineral nutrition ecology of moose, Alces alces, by a comprehensive examination of concentration levels of 18 elements in browsing plants, and in rumen and distal colon contents, respectively. The plants, in total 14 species, included deciduous and evergreen trees, shrubs, herbs, grasses and aquatic macrophytes. Our study was done in four sites in southern Norway and the plants were sampled in spring, summer, autumn and winter to document seasonal trends. The plants were generally characterized by low concentrations of major elements, very low phosphorus: calcium ratios, and the willow Salix aurita - a preferred browse - had high cadmium concentrations, up to 9 ppm (dry mass). Sodium concentrations in terrestrial plants were significantly under the dietary requirement, while levels were sufficient in aquatic plants, which, however, had high concentrations of toxic heavy metals. We suggest that the moose experience a nutritional dilemma as a strong physiological need for sodium results in an overintake of heavy metals. A selective intake of plants rich in heavy metals was corroborated by high heavy metal concentrations in rumen and distal colon. Over all, the plants were characterized by a marked mineral diversity due to element concentration levels that were highly variable between species and sites. Thus, there is a strong link between plant species diversity and mineral diversity, which has a pivotal role for herbivores. It is concluded that a diversity of plant species is a prerequisite for the mineral diversity that the moose needs to obtain essential minerals, not only in sufficient amounts, but also in physiologically balanced proportions.

title: A MULTI-SCALE TEST OF THE FORAGE MATURATION HYPOTHESIS IN A PARTIALLY MIGRATORY UNGULATE POPULATION
abstract: The forage maturation hypothesis (FMH) proposes that ungulate migration is driven by selection for high forage quality. Because quality declines with plant maturation, but intake declines at low biomass, ungulates are predicted to select for intermediate forage biomass to maximize energy intake by following phenological gradients during the growing season. We tested the FMH in the Canadian Rocky Mountains by comparing forage availability and selection by both migrant and nonmigratory resident elk (Cervus elaphus) during three growing seasons from 2002-2004. First, we confirmed that the expected trade-off between forage quality and quantity occurred across vegetation communities. Next, we modeled forage biomass and phenology during the growing season by combining ground and remote-sensing approaches. The growing season started 2.2 days earlier every 1 km east of the continental divide, was delayed by 50 days for every 1000-m increase in elevation, and occurred 8 days earlier on south aspects. Migrant and resident selection for forage biomass was then compared across three spatial scales (across the study area, within summer home ranges, and along movement paths) using VHF and GPS telemetry locations from 119 female elk. Migrant home ranges occurred closer to the continental divide in areas of higher topographical diversity, resulting in migrants consistently selecting for intermediate biomass at the two largest scales, but not at the finest scale along movement paths. In contrast, residents selected maximum forage biomass across all spatial scales. To evaluate the consequences of selection, we compared exposure at telemetry locations of migrant and resident elk to expected forage biomass and digestibility. The expected digestibility for migrant elk in summer was 6.5% higher than for residents, which was corroborated with higher fecal nitrogen levels for migrants. The observed differences in digestibility should increase migrant elk body mass, pregnancy rates, and adult and calf survival rates. Whether bottom-up effects of improved forage quality are realized will ultimately depend on trade-offs between forage and predation. Nevertheless, this study provides comprehensive evidence that montane ungulate migration leads to greater access to higher-quality forage relative to nonmigratory congeners, as predicted by the forage maturation hypothesis, resulting primarily from large-scale selection patterns.

title: Transient nutritional peak in browse foliage after forest clearing advocates cohort management of ungulates
abstract: It is well established that forest clearing produces an intense upsurge of new plant material with assumingly improved nutrient availability to large herbivores. One aspect that is surprisingly unexplored is how the nutrient contents of plant tissue progress with time after clearing. We show that large-scale time-series of forest clearing and moose (Alces alces L.) fitness were positively related, with a temporal lag conforming to the nutritional peak in regrowth of birch (Betula pubescens). Foliage from 5-year-old clearcuts had higher contents of crude protein, and lower contents of indigestible lignin compared to 10- and 15-year-old clearcuts. Several mineral ratios also became less favourable with clearcut age, predominantly due to a rise in calcium content. The significant, but transient improvement of browse quality adds to the accumulating evidence emphasizing the need to consider the population ecology of large herbivores also in terms of cohorts. Furthermore, it underlines the applied importance of incorporating ungulate management into forestry planning.

title: Population Dynamics of Newfoundland Moose Using Cohort Analysis
abstract: Changes in abundance, recruitment rates, and age-specific harvest rates of 3 populations of moose (Alces alces) in Newfoundland were estimated with cohort analysis.  due to overharvesting, but recovered in all areas following reduction of license quotas in the mid-1970's. Age-specific vulnerability to hunting was highest for young adults and lowest for calves and mid-aged adults. Differences in vulnerability were possibly related to hunter preferences, previous experience of moose with hunters, and frequency of movements by moose. Data on kills/day and moose seen/day obtained from hunter questionaires were positively correlated with moose abundance but negatively correlated with the number of hunters. The latter suggested that interference had a strong influence on hunter success. 21 A challenging problem facing wildlife managers is how to measure changes in population abundance with sufficient accuracy to make reasonable management decisions. In general, 2 approaches have been employed. Where sufficient funding and manpower are available, many agencies attempt direct estimation of population abundance using aerial surveys, drive counts, or mark-recapture techniques. These methods are costly, time consuming, and often imprecise. Where resources are more limited, wildlife managers are often forced to rely on indirect indices of abundance such as pellet counts, track counts, or hunter success statistics. Such indices can show conflicting trends, and often no other information is available to evaluate which indices are meaningful. Fisheries scientists are faced with similar problems. Their predicament is perhaps even more acute because direct enumeration is impossible for most fish populations. In response, fisheries scientists have developed a wide variety of methods for assessing stock abundance based largely on catch returns from survey vessels or commercial fleets. A powerful technique to reconstruct historical changes in stock abundance from catch-at-age data is the cohort or virtual population analysis (Ricker 1948, Fry 1949, Pope 1972). We used cohort analysis to reconstruct historical changes in moose abundance in 3 areas of Newfoundland from kill-at-age data gathered during 1966-85. Population estimates for 1966-73 (corresponding to cohorts that have completely passed through the population) were used to estimate rates of recruitment and agespec fic vulnerability, and to calibrate indirect indices that have been used in the past to manage moose populations in Newfoundland. Both cohort analysis and indirect indices were then used to evaluate more recent trends in abundance. Funding for this work was provided by the Wildlife Division, Department of Culture, Recreation and Youth, Newfoundland. The Biological Data Centre, University of British Columbia, kindly provided computer facilities. Data were gathered with the assistance of W. A. Collins, S. M. Oosenbrug, and M. J. Strapp. We thank J. S. Collie, P. R. Krausman, M. F. Lapointe, C. J. Walters, G. H. Winters, and an anonymous reviewer for helpful discussions and comments on the manuscript.

title: Diet overlap among ruminants in Fennoscandia
abstract: Information on overlap in resource use is central to understanding of interspecific exploitation competition and resource partitioning. Despite this, measures of diet overlap among northern ruminants in Fennoscandia is limited to one earlier study (reindeer and sheep). Diet overlap between sympatric moose and roe deer calculated with Schoener’s index was 20.7% and 33.6% during summer (data from one area) and winter (data from two areas), respectively, whereas average diet overlap between moose and red deer was 32.0% during winter (data from four areas). Diet overlap between a coastal island population of red deer and sheep was 59.3% during summer and 63.9% during winter. Summer diet overlap between a sheep and a goat population and a sheep and a reindeer population calculated with data on main types of forage plants was 77.0% and 55.1%, respectively. However, overlap calculated with main plant groups was sometimes considerably higher than when calculated for individual forage species. Neither difference in feeding type nor body mass successfully predicted diet overlap between species pairs (n=9), although there tended to be negative correlation (rp=–0.586, P=0.098) between diet overlap of main plant groups (calculated across studies) and difference in feeding type.

title: INBREEDING DEPRESSION IN CONSERVATION BIOLOGY
abstract: ▪ Abstract Inbreeding depression is of major concern in the management and conservation of endangered species. Inbreeding appears universally to reduce fitness, but its magnitude and specific effects are highly variable because they depend on the genetic constitution of the species or populations and on how these genotypes interact with the environment. Recent natural experiments are consistent with greater inbreeding depression in more stressful environments. In small populations of randomly mating individuals, such as are characteristic of many endangered species, all individuals may suffer from inbreeding depression because of the cumulative effects of genetic drift that decrease the fitness of all individuals in the population. In three recent cases, introductions into populations with low fitness appeared to restore fitness to levels similar to those before the effects of genetic drift. Inbreeding depression may potentially be reduced, or purged, by breeding related individuals. However, the Speke's ...

title: Population properties affect inbreeding avoidance in moose
abstract: Mechanisms reducing inbreeding are thought to have evolved owing to fitness costs of breeding with close relatives. In small and isolated populations, or populations with skewed age- or sex distributions, mate choice becomes limited, and inbreeding avoidance mechanisms ineffective. We used a unique individual-based dataset on moose from a small island in Norway to assess whether inbreeding avoidance was related to population structure and size, expecting inbreeding avoidance to be greater in years with larger populations and even adult sex ratios. The probability that a potential mating event was realized was negatively related to the inbreeding coefficient of the potential offspring, with a stronger relationship in years with a higher proportion or number of males in the population. Thus, adult sex ratio and population size affect the degree of inbreeding avoidance. Consequently, conservation managers should aim for sex ratios that facilitate inbreeding avoidance, especially in small and isolated populations.

title: Effects of inbreeding on fitness-related traits in a small isolated moose population
abstract: Inbreeding can affect fitness-related traits at different life history stages and may interact with environmental variation to induce even larger effects. We used genetic parentage assignment based on 22 microsatellite loci to determine a 25 year long pedigree for a newly established island population of moose with 20–40 reproducing individuals annually. We used the pedigree to calculate individual inbreeding coefficients and examined for effects of individual inbreeding (f) and heterozygosity on fitness-related traits. We found negative effects of f on birth date, calf body mass and twinning rate. The relationship between f and calf body mass and twinning rate were found to be separate but weaker after accounting for birth date. We found no support for an inbreeding effect on the age-specific lifetime reproductive success of females. The influence of f on birth date was related to climatic conditions during the spring prior to birth, indicating that calves with a low f were born earlier after a cold spring than calves with high f. In years with a warm spring, calf f did not affect birth date. The results suggest that severe inbreeding in moose has both indirect effects on fitness through delayed birth and lower juvenile body mass, as well as separate direct effects, as there still was a significant relationship between f and twinning rate after accounting for birth date and body mass as calf. Consequently, severe inbreeding as found in the study population may have consequences for population growth and extinction risk.

title: Applications of next-generation sequencing to phylogeography and phylogenetics.
abstract: This is a time of unprecedented transition in DNA sequencing technologies. Next-generation sequencing (NGS) clearly holds promise for fast and cost-effective generation of multilocus sequence data for phylogeography and phylogenetics. However, the focus on non-model organisms, in addition to uncertainty about which sample preparation methods and analyses are appropriate for different research questions and evolutionary timescales, have contributed to a lag in the application of NGS to these fields. Here, we outline some of the major obstacles specific to the application of NGS to phylogeography and phylogenetics, including the focus on non-model organisms, the necessity of obtaining orthologous loci in a cost-effective manner, and the predominate use of gene trees in these fields. We describe the most promising methods of sample preparation that address these challenges. Methods that reduce the genome by restriction digest and manual size selection are most appropriate for studies at the intraspecific level, whereas methods that target specific genomic regions (i.e., target enrichment or sequence capture) have wider applicability from the population level to deep-level phylogenomics. Additionally, we give an overview of how to analyze NGS data to arrive at data sets applicable to the standard toolkit of phylogeography and phylogenetics, including initial data processing to alignment and genotype calling (both SNPs and loci involving many SNPs). Even though whole-genome sequencing is likely to become affordable rather soon, because phylogeography and phylogenetics rely on analysis of hundreds of individuals in many cases, methods that reduce the genome to a subset of loci should remain more cost-effective for some time to come.

title: Early onset of vegetation growth vs. rapid green-up: impacts on juvenile mountain ungulates.
abstract: Seasonal patterns of climate and vegetation growth are expected to be altered by global warming. In alpine environments, the reproduction of birds and mammals is tightly linked to seasonality; therefore such alterations may have strong repercussions on recruitment. We used the normalized difference vegetation index (NDVI), a satellite-based measurement that correlates strongly with aboveground net primary productivity, to explore how annual variations in the timing of vegetation onset and in the rate of change in primary production during green-up affected juvenile growth and survival of bighorn sheep (Ovis canadensis), Alpine ibex (Capra ibex), and mountain goats (Oreamnos americanus) in four different populations in two continents. We indexed timing of onset of vegetation growth by the integrated NDVI (INDVI) in May. The rate of change in primary production during green-up (early May to early July) was estimated as (1) the maximal slope between any two successive bimonthly NDVI values during this period and (2) the slope in NDVI between early May and early July. The maximal slope in NDVI was negatively correlated with lamb growth and survival in both populations of bighorn sheep, growth of mountain goat kids, and survival of Alpine ibex kids, but not with survival of mountain goat kids. There was no effect of INDVI in May and of the slope in NDVI between early May and early July on juvenile growth and survival for any species. Although rapid changes in NDVI during the green-up period could translate into higher plant productivity, they may also lead to a shorter period of availability of high-quality forage over a large spatial scale, decreasing the opportunity for mountain ungulates to exploit high-quality forage. Our results suggest that attempts to forecast how warmer winters and springs will affect animal population dynamics and life histories in alpine environments should consider factors influencing the rate of changes in primary production during green-up and the timing of vegetation onset.

title: 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.
abstract: 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.

title: To feed or not to feed? Evidence of the intended and unintended effects of feeding wild ungulates
abstract: Ungulate populations are important natural resources, associated with both costs and benefits. Conflicts have arisen between stakeholders who benefit from high ungulate numbers and those faced with the costs. Supplementary or diversionary feeding may potentially mitigate conflicts while maintaining harvest yields but can have conservation implications. We quantified the empirical evidence for whether the intended effects, and hence management goals, of feeding are met. We also examined whether any potential unintended consequences of feeding occur and under what conditions. We found clear evidence that supplementary feeding enhanced reproduction and population growth under certain conditions. By contrast, we found limited evidence of the effectiveness of diversionary feeding to protect crops, forestry, and natural habitats, with positive effects often undermined by increases in ungulate density. However, the use of diversionary feeding to reduce traffic collisions seems promising but requires further investigation. The unintended effects of feeding are typically complex, involving changes to demography, behavior, and vegetation with consequent cascading effects on other trophic levels, as well as exacerbated risks of disease transmission. Increased ungulate density is the primary driver behind these unintended effects, the consequences of which tend to increase with longevity of feeding and affect a range of stakeholders. We urge managers to take seriously the risks as well as the economic and ethical issues before deciding to feed ungulates.