Impact of simulated moose densities on abundance and richness of vegetation,herbivorous and predatory arthropods along a productivity gradient

Large herbivores can affect vegetation structure and species composition as well as material and energy flows in the ecosystem through their selective feeding, defecation, urination and trampling. These changes have a large potential to indirectly affect other trophic levels, but the mechanisms are poorly known. We studied the impacts of moose Alces alces browsing along a gradient of site productivity by experimentally simulating four different moose densities. Here we show that moose can affect the richness and abundance of three trophic levels in Swedish boreal forests through complex direct and indirect impacts, but in qualitatively different ways depending on how the physical habitat or food resources of a trophic level are affected. Vegetation richness had a hump‐shaped (unimodal) response to increased moose density. Leaf litter production decreased when browsing increased, which in turn depressed the abundance of flying prey for spiders. Consequently, spider abundance and richness declined monotonically. The responses of spider richness to moose density were further conditioned by site productivity: the response was positive at productive and negative at unproductive sites. In contrast, herbivorous Hemiptera were not affected by moose, most likely because the abundance of their food plants was not affected. The highest simulated moose density had an impact on all variables responding to moose even after a few years of treatment and can be considered as overabundance. We also show that the impacts of low or moderate moose density can be positive to some of the organisms negatively affected by high density. The level of herbivore population density that leads to substantial community impacts also depends on site factors, such as productivity.     

Effect of Roadside Vegetation Cutting on Moose Browsing

Moose (Alces americanus ) vehicle collisions (MVCs) are an issue throughout the distribution of moose. Many mitigation strategies have been tested and implemented to reduce the number of MVCs, but there have been few empirical analyses of the effectiveness of roadside vegetation cutting. The goal of this study was to determine if roadside vegetation cutting attracted moose into roadside areas to browse on the vegetation regrowth. We hypothesized that moose would be attracted to roadside areas with cut vegetation. Consequently, we predicted that there would be higher levels of browsing in cut areas compared to uncut areas. To determine if moose were browsing more in cut or uncut areas, we measured the number of plants browsed by moose in paired treatment (cut on or after 2008) and control (not cut since at least 2008) sites, along with a suite of potential environmental covariates. Using a model selection approach, we fit generalized linear mixed-effects models to determine the most parsimonious set of environmental variables to explain variation in the proportion of moose browse among sites. In contrast to our hypothesis, our results show that the proportion of moose browse in the uncut control areas was significantly higher than in the cut treatment areas. The results of this study suggest that recently cut roadside areas (7 years or less based on our work) may create a less attractive foraging habitat for moose. The majority of the variance in the proportion of moose browse among sites was explained by treatment type and nested plot number within site identification (34.16%), with additional variance explained by traffic region (5.00%) and moose density (4.35%). Based on our study, we recommend that vegetation cutting be continued in roadside areas in Newfoundland as recently cut areas may be less attractive browsing sites for moose.     

Moose (Alces alces) calf survival rates in the presence of wolves (Canis lupus) in southeast Norway

Wolves (Canis lupus) are recolonizing Scandinavia and becoming a new limiting factor that should be taken into account in the management of moose (Alces alces). However, there is a lack of empirical estimates of moose survival after wolf recolonization. We investigated the effects of wolf abundance, moose litter size (single/twin calves), and climatic factors on annual and seasonal calf survival rates in a moose population in southeast Norway. We used data that were obtained over 7?years by radio-tracking and regular visual observations of 68 moose cows to determine the presence or absence of calves at heel. Annual and winter calf survival rates were 20–40 percentage points lower in the wolf territory compared with previous estimates of moose calf survival in similar areas that lacked wolves. Cause-specific studies of mortality would further enhance our ability to determine the relative role of various limiting factors. Our study suggests that moose managers should regulate quotas to buffer the lower survival rates after wolf recolonization.     

Synchrony in short-term fluctuations of moose calf body mass and bank vole population density supports the mast depression hypothesis

Inter-annual variations in body mass of moose, Alces alces , in Norway and Sweden have been considered as most likely due to direct or indirect effects of weather, but so far predictions of autumn body mass of moose calves on the basis of weather data have given a poor fit to data. A striking, but hitherto unnoticed, feature of several time series on body mass of moose calves from south-eastern Norway is an apparently regular 3–4-year fluctuation pattern. This short-term fluctuation could be due to regular variations in forage quality, e.g. caused by a cyclic seed production of some important food plants, as envisaged by the "mast depression" hypothesis. One plant species important as food for moose calves in autumn is bilberry, Vaccinium myrtillus , which usually produces high seed crops (masts) at intervals of 3–4 years. Populations of the bank vole, Clethrionomys glareolus, which feeds on bilberry shoots in winter, are known to peak in bilberry post-mast years. In two study areas in Norway, there was a positive correlation between the autumn body mass of moose calves and the autumn population index of bank vole in the succeeding year. In the northern area there was an additional positive effect of summer precipitation, whereas in the southern area there was an additional negative effect of summer temperature. In both areas, however, the effect of weather was less pronounced than that of the bank vole index. These results support the mast depression hypothesis.     

The non-impact of hunting on moose <Emphasis Type="Italic">Alces alces</Emphasis> movement,diurnal activity,and activity range

Previous studies on moose Alces alces have suggested that interactions with humans may trigger anti-predator behaviors and generate a demographical cost. Therefore, we hypothesized that disturbances from small and big game hunting may have negative effects on moose movements, diurnal activity, and activity range. Using location data from 64 moose equipped with GPS collars from three populations (Low Alpine, Inland, Coastal) with different temporal human presence and spatial accessibility, we evaluated the impact of hunting on moose activity rhythms. On average, female moose in the low human population density (Low Alpine) area (<0.5/km²) had significantly lower movement rates during moose hunting season, but variation in movement rates among individuals were higher compared with female moose in regions with denser human populations (6–24/km²). We found no evidence that reproductive status influenced female moose sensitivity to disturbance. As expected, females used smaller activity ranges and were less active nocturnally than males. The high within-group variation suggests that current hunting disturbance levels do not alter moose population behavior in general. Our data indicate that alterations in movement were related to rutting activity, not human disturbance induced by hunting. In line with behavioral theory, our study suggests that some individuals were more sensitive to hunting disturbance than the general population. Our work suggests that individual moose may perceive human predation risk to be similar to other predation risks.     

Body condition and population regulation by food resources in moose

Summary In southwestern Québec, non-harvested moose populations stabilize at a density of 0.40 animal·km^-2. In an attempt to test population regulation by food resources, we investigated moose body condition near this equilibrium density (0.37) and at 2 lower densities (0.22 and 0.17). Annual population growth rates were evaluated at 4, 18, and 24% respectively. We predicted that moose in the high density area would exhibit growth retardation and poorer body condition, compared to moose in the lower density areas. Measurements of head length, cranial breadth, heart weight, and kidney weight were collected from 443 moose killed during the regular autumn harvesting seasons of 1981 and 1982. There was no indication that body condition was poorer at high moose density, and hence no evidence that foraging conditions were deteriorated. We concluded that food limitation was not sufficient to explain the differences in population growth rates. Predation by wolves and/or black bears is presented as an alternative and testable hypothesis.     

Diet optimization in a generalist herbivore: The moose

In an attempt to understand the foraging of a generalist herbivore, a linear programming optimization model was constructed to describe moose feeding in summer. The model attempts to predict the amounts of aquatic vegetation, deciduous leaves, and forbs a moose should consume each day; and to determine whether or not its feeding is constrained by the maximum feeding time available each day, its daily rumen processing capacity, its sodium requirements, and its energy metabolism. The model can be solved for two alternative strategies: time minimization and energy maximization. The energy-maximizing strategy appears to predict the observed diet chosen by an average moose very well. Also, the diets selected by moose of each sex and various reproductive states appear to fit the energy-maximizing strategy. In addition, it is demonstrated that a moose's body size at weaning, size at first reproduction, and maximum size are related to foraging efficiency. Furthermore, there appears to exist an optimum adult body size for feeding. The general conclusion arrived at is that the foraging of a generalist herbivore can be predicted in a quantitative manner, at least in this case, as has been shown for other types of consumers (carnivores and granivores).     

Linking Moose Population and Plant Growth Models with a Moose Energetics Model

Selective foraging by large mammals can change ecosystem properties such as plant species composition, nutrient cycling rates, and soil fertility. These changes, in turn, alter the availability of forage and could affect the relative efficiencies of foraging strategies used by these animals. We used a simulation model to predict how alternate foraging strategies affected the net annual energy balance of moose (Alces alces), moose density, and distribution of browse across the landscape. The model simulates the spatial distribution of vegetation in an 8-ha landscape of 1-m² cells with seasonal changes in the energetic needs of free-ranging moose and plant phenology. The energetics model was integrated with a moose population model and a plant-growth model for long-term simulations. Changes in bite density in each feeding station are predicted with height and biomass logistic curves modified by a quadratic response to browsing. We tested foraging strategies using random, fractional, and marginal value theorem (MVT) algorithms on landscapes with a range of bite densities and differing spatial distributions. Small-scale disturbances (that is, tree-fall gaps) were required to maintain browse supply and prevent moose population extinction under all foraging strategies. Populations using a fractional stopping rule survived the 100-year simulations because moose browsed across much of the landscape and did not overbrowse patches with high bite density. Populations using random and MVT stopping rules became extinct in about 25 and about 50 years, respectively. Moose using a random stopping rule were in negative energy balance because travel time was high and the net energy intake rate was low on an annual basis. Moose using the MVT stopping rule were initially in positive energy balance, but as the high-density browse patches were overbrowsed and low-density unbrowsed patches grew out of reach, bite density decreased, and energy balance became negative in subsequent years. Thus, the foraging strategy used by individual moose resulted in creation of landscapes that strongly affected browse density, browse distribution, moose population density, and moose survival. Received 30 April 1997; accepted 5 August 1997.     

Evaluating if Energy and Protein Limit Abundance of Hawaiian Moorhen

Abstract: Food abundance can affect a species' distribution. In many studies of potential food limitation, researchers focus on carrying capacity estimates during the nonbreeding season for temperate species consuming a fixed food source. Estimates of energetic carrying capacity for year-round breeders feeding on a replenishing resource would be more difficult and require much data. To determine whether gathering detailed information on year-round carrying capacity would be an important investment, we conducted an assessment to determine whether there was evidence that energy or protein might limit numbers of the tropical, endangered Hawaiian moorhen (Gallinula chloropus sandvicensis). We compared observed numbers of moorhen at 15 Oahu, Hawaii, USA, wetlands with predicted numbers based on measured energy and protein in food plants and abundance of these food plants in each wetland and on estimates of energy expenditure of moorhen. We made comparisons assuming moorhen are limited by their ability to metabolize food plants, by competition for food, and by estimated costs associated with reproduction. We also compared ranked moorhen abundance and density with ranked energy and protein under different wetland management regimes. Energy values consistently overestimated expected numbers of Hawaiian moorhen at wetlands except for one wetland location (predicted, 3803 ± 4856; observed, 6.2 ± 10.8). In addition, we detected no significant relationship between moorhen abundance and measures of energy (all r² = 0.02–0.73, all P > 0.1) or protein abundance (all r² = 0.08–0.50, all P > 0.3). This lack of relationship held once we controlled for wetland area or when we considered whether wetlands were managed for waterbirds. Hawaiian moorhen on Oahu did not appear to be limited by energy, nor did they appear to select sites based on energy or protein, in contrast to many studies relating animal numbers to energy in nonbreeding situations. Consequently, we suggest that researchers and managers explore other potentially limiting factors for Hawaiian moorhen.     

Reconstructing landscapes of ungulate parturition and predation using vegetation phenology

Enhanced vegetation index (EVI) data can be used to identify and define the space in which ungulates practice parturition and encounter predation. This study explores the use of EVI data to identify landscapes linked to ungulate parturition and predation events across space, time, and environmental conditions. As a case study, we used the moose population (Alces alces) of northern Minnesota in the USA. Using remotely sensed EVI data rasters and global positioning system collar data, we quantified how vegetation phenology and moose movement shaped the births and predation of 52 moose calves from 2013 to 2020 on or adjacent to the Grand Portage Indian Reservation. The known sources of predation were American black bears (Ursus americanus, n = 22) and gray wolves (Canis lupus, n = 28). Satellite-derived data summarizing seasonal landscape features at the local level revealed that landscape heterogeneity use by moose can help to quantitatively identify landscapes of parturition and predation in space and time across large areas. Vegetation phenology proved to be differentiable between adult moose ranges, sites of cow parturition, and sites of calf predation. Landscape characteristics of each moose group were consistent and tractable based on environment, suggesting that sites of parturition and predation of moose are predictable in space and time. It is possible that moose selected specific landscapes for parturition despite risk of increased predation of their calves, which could be an example of an "ecological trap." This analytical framework can be employed to identify areas for future ungulate research on the impacts of landscape on parturition and predation dynamics.     

Selectivity by moose vs the spatial distribution of aspen: a natural experiment

Patch use theory predicts that herbivores perceive food as patches and spend more time in high quality patches, i.e. feeding sites providing a high net rate of intake of energy and/or limiting nutrients. The herbivores should accordingly not discriminate among food items in such high quality patches, and food items should thus be eaten in proportion to availability. In contrast, classical diet theory assumes food selection to take place at the level of individual plants, and predicts that the forager should concentrate on the most profitable item until availability drops below some critical threshold.
Here we address how the spatial distribution of European aspen Populus tremula , a highly preferred browse species, affects the selectivity by moose Alces alces at the patch and the tree level. The study was performed in a managed boreal forest landscape in coastal northern Sweden, where aspen occurs highly aggregated almost exclusively in discrete patches. We compared moose' selectivity for aspen and browsing intensity on aspen ramets and other browse species in aspen patches versus at randomly located sites.
Random sites and aspen stands were utilised equally by moose in terms of overall use of forage. There was no difference in total coverage of forage species and relative moose density. Selectivity for aspen was stronger at random sites than at aspen sites, and the browsing intensity on aspen was similar. We conclude that moose did not perceive aspen stands as discrete patches, and used aspen ramets more in accordance with diet theory. These findings agree with the idea that large generalist herbivores strive to maintain a mixed and balanced diet, causing rare species to be over-utilised (negative frequency-dependent food selection). By such selective feeding, moose may reinforce the aggregated distribution of aspen in the managed boreal forest landscape.     

Moose selecting for specific nutritional composition of birch places limits on food acceptability

Despite decades of intense research, it remains largely unsolved which nutritional factors underpin food selection by large herbivores in the wild. We measured nutritional composition of birch foliage (Betula pubescens) available to, and used by, moose (Alces alces) in natural settings in two neighboring regions with contrasting animal body mass. This readily available food source is a staple food item in the diet of moose in the high‐fitness region, but apparently underutilized by moose in the low‐fitness region. Available birch foliage in the two regions had similar concentrations of macronutrients (crude protein [CP], fiber fractions, and water‐soluble carbohydrates [WSC]), although a notably lower variation of WSC in the low‐fitness region. For minerals, there were several area differences: available birch foliage in the low‐fitness region had less Mg (depending on year) and P, but more Ca, Zn, Cu, and Mn. It also had higher concentrations of some plant secondary metabolites: chlorogenic acids, quercetins, and especially MeOH‐soluble condensed tannins. Despite the area differences in available foliage, we found the same nutritional composition of birch foliage used in the two regions. Compared to available birch foliage, moose consistently used birch foliage with more CP, more structural fiber (mainly hemicellulose), less WSC, higher concentrations of several minerals (Ca, Zn, K, Mn, Cu), and lower concentrations of some secondary metabolites (most importantly, MeOH‐soluble condensed tannins). Our study conceptually supports the nutrient‐balancing hypothesis for a large herbivore: within a given temporal frame, moose select for plant material that matches a specific nutritional composition. As our data illustrate, different moose populations may select for the same composition even when the nutritional composition available in a given food source varies between their living areas. Such fastidiousness limits the proportion of available food that is acceptable to the animal and has bearings on our understanding and application of the concept of carrying capacity.     

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

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

The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L)

The nutrient balancing hypothesis proposes that, when sufficient food is available, the primary goal of animal diet selection is to obtain a nutritionally balanced diet. This hypothesis can be tested using the Geometric Framework for nutrition (GF). The GF enables researchers to study patterns of nutrient intake (e.g. macronutrients; protein, carbohydrates, fat), interactions between the different nutrients, and how an animal resolves the potential conflict between over-eating one or more nutrients and under-eating others during periods of dietary imbalance. Using the moose (Alces alces L.), a model species in the development of herbivore foraging theory, we conducted a feeding experiment guided by the GF, combining continuous observations of six captive moose with analysis of the macronutritional composition of foods. We identified the moose’s self-selected macronutrient target by allowing them to compose a diet by mixing two nutritionally complementary pellet types plus limited access to Salix browse. Such periods of free choice were intermixed with periods when they were restricted to one of the two pellet types plus Salix browse. Our observations of food intake by moose given free choice lend support to the nutrient balancing hypothesis, as the moose combined the foods in specific proportions that provided a particular ratio and amount of macronutrients. When restricted to either of two diets comprising a single pellet type, the moose i) maintained a relatively stable intake of non-protein energy while allowing protein intakes to vary with food composition, and ii) increased their intake of the food item that most closely resembled the self-selected macronutrient intake from the free choice periods, namely Salix browse. We place our results in the context of the nutritional strategy of the moose, ruminant physiology and the categorization of food quality.     

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

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

Ranking Alaska Moose Nutrition: Signals to Begin Liberal Antlerless Harvests

Abstract: We focused on describing low nutritional status in an increasing moose (Alces alces gigas) population with reduced predation in Game Management Unit (GMU) 20A near Fairbanks, Alaska, USA. A skeptical public disallowed liberal antlerless harvests of this moose population until we provided convincing data on low nutritional status. We ranked nutritional status in 15 Alaska moose populations (in boreal forests and coastal tundra) based on multiyear twinning rates. Data on age-of-first-reproduction and parturition rates provided a ranking consistent with twinning rates in the 6 areas where comparative data were available. Also, short-yearling mass provided a ranking consistent with twinning rates in 5 of the 6 areas where data were available. Data from 5 areas implied an inverse relationship between twinning rate and browse removal rate. Only in GMU 20A did nutritional indices reach low levels where justification for halting population growth was apparent, which supports prior findings that nutrition is a minor factor limiting most Alaska moose populations compared to predation. With predator reductions, the GMU 20A moose population increased from 1976 until liberal antlerless harvests in 2004. During 1997-2005, GMU 20A moose exhibited the lowest nutritional status reported to date for wild, noninsular, North American populations, including 1) delayed reproduction until moose reached 36 months of age and the lowest parturition rate among 36-month-old moose (29%, n = 147); 2) the lowest average multiyear twinning rates from late-May aerial surveys (x = 7%, SE = 0.9%, n = 9 yr, range = 3-10%) and delayed twinning until moose reached 60 months of age; 3) the lowest average mass of female short-yearlings in Alaska (x̄ = 155 ± 1.6 [SE] kg in the Tanana Flats subpopulation, up to 58 kg below average masses found elsewhere); and 4) high removal (42%) of current annual browse biomass compared to 9-26% elsewhere in boreal forests. When average multiyear twinning rates in GMU 20A (sampled during 1960-2005) declined to <10% in the mid- to late 1990s, we began encouraging liberal antlerless harvests, but only conservative annual harvests of 61-76 antlerless moose were achieved during 1996-2001. Using data in the context of our broader ranking system, we convinced skeptical citizen advisory committees to allow liberal antlerless harvests of 600-690 moose in 2004 and 2005, with the objective of halting population growth of the 16,000-17,000 moose; total harvests were 7-8% of total prehunt numbers. The resulting liberal antlerless harvests served to protect the moose population's health and habitat and to fulfill a mandate for elevated yield. Liberal antlerless harvests appear justified to halt population growth when multiyear twinning rates average ≤10% and ≥1 of the following signals substantiate low nutritional status: <50% of 36-month-old moose are parturient, average multiyear short-yearling mass is <175 kg, or >35% of annual browse biomass is removed by moose.     

Effects of winter recreation on northern ungulates with focus on moose (Alces alces) and snowmobiles

Winter recreation can displace ungulates to poor habitats, which may raise their energy expenditure and lower individual survivorship, causing population declines. Winter recreation could be benign, however, if animals habituate. Moreover, recreation creates trails. Traveling on them could reduce energy expenditure, thereby increasing ungulate survivorship and generating population benefits. Balancing recreation use with wildlife stewardship requires identifying when these effects occur. This task would be simpler if guidelines existed to inform assessments. We developed and tested such guidelines using two approaches. First, we synthesized literature describing the effects of winter recreation—motorized and nonmotorized—on northern ungulates. This synthesis enabled formulating six guidelines, while exposing two requiring further attention (ungulate habituation and displacement). Second, we tested these two guidelines and evaluated the others by quantifying the behavioral responses of moose to snowmobiles, in two areas of south-central Alaska, differing by snowmobile predictability. For each location, we modeled moose preferences during the snowmobile period using different combinations of eight variables—static (elevation and slope), biotic (habitat and cover), and anthropogenic (distance to roads, railroads, snowmobile trails, and trail density). We identified the model with the most support and used it to estimate parameter coefficients for pre- and post-recreation periods. Changes in coefficients between periods indicated snowmobile effects on moose. Overall, we produced and evaluated six guidelines describing when winter recreation is potentially detrimental to ungulates as follows: (1) when unpredictable, (2) spanning large areas, (3) long in duration, (4) large spatial footprint, (5) nonmotorized, and (6) when animals are displaced to poor quality habitats.     

A noninvasive and integrative approach for improving density and abundance estimates of moose

Acquiring demographic data for moose (Alces alces) can be difficult because they are solitary in nature, they prefer densely vegetated and mountainous habitats, and they often occur at low density. Such data, however, are essential for long-term population monitoring, evaluating management practices, and effective conservation. Winter aerial surveys are the standard method for estimating moose population parameters, but they can be logistically challenging, expensive, and subject to sightability correction, which necessitates the capture of study animals for initial model development. Herein, we demonstrate a noninvasive alternative approach for estimating population parameters of moose in northern Yellowstone National Park, where aerial surveys were attempted but proved ineffective. We determined individual moose genotype and sex using microsatellite polymerase chain reaction amplification of DNA extracted from fecal pellets, integrated ancillary pellet sample data (i.e., metadata) in genotype analysis to aid in the identification of matching genotypes, and used spatially explicit capture-recapture (SECR) modeling to estimate sex-specific density and abundance. We collected 616 samples over 3 consecutive winters (Dec 2013–Apr 2016) and within 2 sampling occasions each winter. We recorded 514 captures of 142 individual moose (69 males, 73 females). Overall density ranged between 0.062 moose/km² and 0.076 moose/km² and averaged 0.034/km² for females and 0.033/km² for males. Abundance estimates were 150 moose in 2013 (female = 76, 95% CI = 55–105; male = 74, 95% CI = 54–103), 186 in 2014 (female = 95, 95% CI = 63–142; male = 91, 95% CI = 60–138), and 160 in 2015 (female = 79, 95% CI = 58–108; male = 81, 95% CI = 59–110). Average population sex ratio was 0.99 males/female. We demonstrate that SECR analysis of fecal DNA genotypes, using metadata in genotype analysis to help identify matching moose genotypes, is a promising alternative method for estimating sex-specific density and abundance of a low-density moose population in a mountainous and forested landscape.     

Large-scale climatic fluctuation and population dynamics of moose and white-tailed deer

1. Mech et al . (1987) documented cumulative, negative effects of previous winters' snow on rates of population increase in moose ( Alces alces ) and white-tailed deer ( Odocoileus virginianus ), but noted no effect of predation by wolves ( Canis lupus ). Those results were contested by Messier (1991), who analysed smoothed versions of the original abundance data and reported no effect of snow accumulation on population dynamics of either species, but strong effects of wolf predation and food competition.
2. McRoberts, Mech & Peterson (1995) contended that the conclusions reached by Messier (1991) were an artefact of the use of smoothed data. In a subsequent re-analysis of the smoothed data, Messier (1995) argued that the lack of an effect of snow after one year precluded the potential for a cumulative effect beyond one year.
3. We re-analysed original and smoothed data on dynamics of moose and white-tailed deer densities using the same methods as Mech et al . (1987) and Messier (1991), but we used a measure of global climatic fluctuation, the North Atlantic Oscillation (NAO) index. The NAO is the atmospheric process determining most interannual variation in snowfall and winter temperatures in northern latitudes, and its phases drive decadal trends in wintertime precipitation.
4. We observed that rates of increase of moose and white-tailed deer in both the original and smoothed data were influenced by global climatic fluctuation at 2- and 3-year lags, as well as by delayed density-dependent feedback and wolf predation.     

Simulated responses of moose populations to browsing‐induced changes in plant architecture and forage production

We and others have previously shown that browsing by large mammalian herbivores can alter the fractal dimension and other architectural properties of plant crowns. Using data from an experiment in northern Sweden which manipulated moose population densities from 0 to 50 moose per 1000 ha and measured architectural responses of birch, Betula pendula and B. pubescens, and Scots pine, Pinus sylvestris, we constructed a simulation model to examine how these changes in plant crown architecture affect moose population dynamics. Architectural changes in birch tree crowns caused forage availability to increase from low to moderate moose population densities, then decline at higher densities. In contrast, forage availability of pine decreased monotonically with increasing moose population density. The model, which incorporated equations fitted to these experimental results, predicted realistic moose population densities for northern Sweden. More importantly, the model also predicted that the quadratic responses of birch crowns to moose browsing results in moose population oscillations on high productivity sites because the moose population density overshoots the maximum birch forage availability then declines. Changes in the geometry of plant canopies caused by mammalian browsers and soil fertility may feed back on the population dynamics of the browsers themselves and even result in complex dynamics such as population oscillations.