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USGS2015

Population genetic structure of moose ( Alces alces ) of South-central Alaska

The location of a population can influence its genetic structure and diversity by impacting the degree of isolation and connectivity to other populations. Populations at range margins are often thought to have less genetic variation and increased genetic structure, and a reduction in genetic diversity can have negative impacts on the health of a population. We explored the genetic diversity and connectivity between 3 peripheral populations of moose ( Alces alces ) with differing potential for connectivity to other areas within interior Alaska. Populations on the Kenai Peninsula and from the Anchorage region were found to be significantly differentiated (F ST = 0.071, P < 0.0001) with lower levels of genetic diversity observed within the Kenai population. Bayesian analyses employing assignment methodologies uncovered little evidence of contemporary gene flow between Anchorage and Kenai, suggesting regional isolation. Although gene flow outside the peninsula is restricted, high levels of gene flow were detected within the Kenai that is explained by male-biased dispersal. Furthermore, gene flow estimates differed across time scales on the Kenai Peninsula which may have been influenced by demographic fluctuations correlated, at least in part, with habitat change.

Alces↗

USGS2002

Mitochondrial phylogeography of moose ( Alces alces ): Late Pleistocene divergence and population expansion

We examined phylogeographic relationships of moose ( Alces alces ) worldwide to test the proposed existence of two geographic races and to infer the timing and extent of demographic processes underpinning the expansion of this species across the Northern Hemisphere in the late Pleistocene. Sequence variation within the left hypervariable domain of the control region occurred at low or moderate levels worldwide and was structured geographically. Partitioning of genetic variance among regions indicated that isolation by distance was the primary agent for differentiation of moose populations but does not support the existence of distinct eastern and western races. Levels of genetic variation and structure of phylogenetic trees identify Asia as the origin of all extant mitochondrial lineages. A recent coalescence is indicated, with the most recent common ancestor dating to the last ice age. Moose have undergone two episodes of population expansion, likely corresponding to the final interstade of the most recent ice age and the onset of the current interglacial. Timing of expansion for the population in the Yakutia–Manchuria region of eastern Asia indicates that it is one of the oldest populations of moose and may represent the source of founders of extant populations in North America, which were colonized within the last 15,000 years. Our data suggest an extended period of low population size or a severe bottleneck prior to the divergence and expansion of extant lineages and a recent, less-severe bottleneck among European lineages. Climate change during the last ice age, acting through contraction and expansion of moose habitat and the flooding of the Bering land bridge, undoubtedly was a key factor influencing the divergence and expansion of moose populations.

Alaska, Colorado↗

USGS2015

A genetic discontinuity in moose ( Alces alces ) in Alaska corresponds with fenced transportation infrastructure

The strength and arrangement of movement barriers can impact the connectivity among habitat patches. Anthropogenic barriers (e.g. roads) are a source of habitat fragmentation that can disrupt these resource networks and can have an influence on the spatial genetic structure of populations. Using microsatellite data, we evaluated whether observed genetic structure of moose ( Alces alces ) populations were associated with human activities (e.g. roads) in the urban habitat of Anchorage and rural habitat on the Kenai Peninsula, Alaska. We found evidence of a recent genetic subdivision among moose in Anchorage that corresponds to a major highway and associated infrastructure. This subdivision is most likely due to restrictions in gene flow due to alterations to the highway (e.g. moose-resistant fencing with one-way gates) and a significant increase in traffic volume over the past 30 years; genetic subdivision was not detected on the Kenai Peninsula in an area not bisected by a major highway. This study illustrates that anthropogenic barriers can substructure wildlife populations within a few generations and highlights the value of genetic assessments to determine the effects on connectivity among habitat patches in conjunction with behavioral and ecological data. .

Alaska↗

USGS2004

Trace elements in moose (Alces alces) found dead in Northwestern Minnesota, USA

The moose ( Alces alces ) population in bog and forest areas of Northwestern Minnesota has declined for more than 25 years, and more recently the decline is throughout Northwestern Minnesota. Both deficiencies and elevations in trace elements have been linked to the health of moose worldwide. The objective of this study was to evaluate whether trace element toxicity or deficiency may have contributed to the decline of moose in Northwestern Minnesota. Livers of 81 moose found dead in Northwestern Minnesota in 1998 and 1999 were analyzed for trace elements. With the exception of selenium (Se) and copper (Cu), trace elements were not at toxic or deficient levels based on criteria set for cattle. Selenium concentrations in moose livers based on criteria set for cattle were deficient in 3.7% of livers and at a chronic toxicity level in 16% of livers. Copper concentrations based on criteria set for cattle were deficient in 39.5% of livers, marginally deficient in 29.5% of livers and adequate in 31% of livers. Moose from agricultural areas had higher concentrations, on average, of Cd, Cu, Mo and Se in their livers than moose from bog and forest areas. Older moose had higher concentrations of Cd and Zn, and lower concentrations of Cu than younger moose. Copper deficiency, which has been associated with population declines of moose in Alaska and Sweden, may be a factor contributing to the decline of moose in Northwestern Minnesota. (C) 2004 Elsevier B.V. All rights reserved.

Minnesota↗

USGS2019

Negative frequency-dependent foraging behaviour in a generalist herbivore (Alces alces) and its stabilizing influence on food-web dynamics

1. Resource selection is widely appreciated to be context‐dependent and shaped by both biological and abiotic factors. However, few studies have empirically assessed the extent to which selective foraging behaviour is dynamic and varies in response to environmental conditions for free‐ranging animal populations. 2. Here, we assessed the extent that forage selection fluctuated in response to different environmental conditions for a free‐ranging herbivore, moose (Alces alces), in Isle Royale National Park, over a 10‐year period. More precisely, we assessed how moose selection for coniferous versus deciduous forage in winter varied between geographic regions and in relation to (a) the relative frequency of forage types in the environment (e.g. frequency‐dependent foraging behaviour), (b) moose abundance, (c) predation rate (by grey wolves) and (d) snow depth. These factors are potentially important for their influence on the energetics of foraging. We also built a series of food‐chain models to assess the influence of dynamic foraging strategies on the stability of food webs. 3. Our analysis indicates that moose exhibited negative frequency dependence, by selectively exploiting rare resources. Frequency‐dependent foraging was further mediated by density‐dependent processes, which are likely to be predation, moose abundance or some combination of both. In particular, frequency dependence was weaker in years when predation risk was high (i.e. when the ratio of moose to wolves was relatively low). Selection for conifers was also slightly weaker during deep snow years. 4. The food‐chain analysis indicates that the type of frequency‐dependent foraging strategy exhibited by herbivores had important consequences for the stability of ecological communities. In particular, the dynamic foraging strategy that we observed in the empirical analysis (i.e. negative frequency dependence being mediated by density‐dependent processes) was associated with more stable food web dynamics compared to fixed foraging strategies. 5. The results of this study indicated that forage selection is a complex ecological process, varying in response to both biological (predation and moose density) and abiotic factors (snow depth) and over relatively small spatial scales (between regions). This study also provides a useful framework for assessing the influence of other aspects of foraging behaviour on the stability of food web dynamics.

Michigan↗

USGS2003

Mitochondrial phylogeography of moose (Alces alces) in North America

Nucleotide variation was assessed from the mitochondrial control region of North American moose ( Alces alces ) to test predictions of a model of range expansion by stepping-stone dispersal and to determine whether patterns of genetic variation support the current recognition of 4 subspecies. Haplotypes formed a star phylogeny indicative of a recent expansion of populations. Values of nucleotide and haplotype diversity were low continentwide but were greatest in the central part of the continent and lowest in peripheral populations. Despite low mitochondrial diversity, moose exhibited a high degree of differentiation regionally, which was not explained by isolation by distance. Our data indicate a pattern of colonization consistent with a large central population that supplied founders to peripheral populations (other than Alaska), perhaps through rare, long-distance dispersal events (leptokurtic dispersal) rather than mass dispersal by a stepping-stone model. The colonization scenario does not account for the low haplotype diversity observed in Alaska, which may be derived from a postcolonization bottleneck. Establishment of peripheral populations by leptokurtic dispersal and subsequent local adaptation may have been sufficient for development of morphological differentiation among extant subspecies.

Journal of Mammalogy↗

USGS2013

Age structure of moose (Alces alces) killed by gray wolves (Canis lupus) in northeastern Minnesota, 1967-2011

The ages of 77 adult Moose ( Alces alces ) killed by Gray Wolves ( Canis lupus ) during the period 1967–2011 in northeastern Minnesota were significantly older than those of a sample of 17,585 Moose killed by hunters in nearby Ontario. Our findings support those of earlier studies of protected Moose populations in national parks that found that Gray Wolves tend to kill disproportionately more older Moose.

Minnesota↗

USGS2010

Effects of simulated moose Alces alces browsing on the morphology of rowan Sorbus aucuparia

In much of northern Sweden moose Alces alces browse rowan Sorbus aucuparia heavily and commonly revisit previously browsed plants. Repeated browsing of rowan by moose has created some concern for its long-term survival in heavily browsed areas. We therefore measured how four years of simulated moose browsing at four population densities (0, 10, 30 and 50 moose/1,000 ha) changed plant height, crown width, available bite mass, the number of bites per plant and per plant forage biomass of rowan saplings. Increased biomass removal led to a significant decline in plant height (P < 0.001), but a significant increase in the number of bites per plant (P = 0.012). Increases in the number of bites per plant more than compensated for weak decreases in bite mass, leading to a weak increase in per plant forage biomass (P = 0.072). With the decline in plant height and increase in the number of stems per plant, a greater number of bites remain within the height reach of moose relative to unbrowsed controls. Moose therefore stand to benefit from revisiting previously browsed plants, which may result in feeding loops between moose and previously browsed rowan saplings. ?? 2010 Wildlife Biology, NKV.

Wildlife Biology↗

USGS2025

Observations of tear-drinking by lepidopterans on moose ( Alces alces americana ) in northeastern North America

Lepidoptera have long been known to feed on the tears of vertebrates as a presumed source of minerals or nutrients. While this unusual behavior has been observed in a variety of species, only a single previous record has been documented outside of the tropics. Here, we present the first documentation of moths visiting the eyes of a bull moose ( Alces americanus americanus ), captured via trail camera in Green Mountain National Forest, Vermont, United States. We discuss the biogeography of this behavior, how it may differ between tropical and temperate climates, and its potential impact on moose health.

Vermont↗

USGS2018

Incorporating road crossing data into vehicle collision risk models for moose (Alces americanus) in Massachusetts, USA

Wildlife–vehicle collisions are a human safety issue and may negatively impact wildlife populations. Most wildlife–vehicle collision studies predict high-risk road segments using only collision data. However, these data lack biologically relevant information such as wildlife population densities and successful road-crossing locations. We overcome this shortcoming with a new method that combines successful road crossings with vehicle collision data, to identify road segments that have both high biological relevance and high risk. We used moose ( Alces americanus ) road-crossing locations from 20 moose collared with Global Positioning Systems as well as moose–vehicle collision (MVC) data in the state of Massachusetts, USA, to create multi-scale resource selection functions. We predicted the probability of moose road crossings and MVCs across the road network and combined these surfaces to identify road segments that met the dual criteria of having high biological relevance and high risk for MVCs. These road segments occurred mostly on larger roadways in natural areas and were surrounded by forests, wetlands, and a heterogenous mix of land cover types. We found MVCs resulted in the mortality of 3% of the moose population in Massachusetts annually. Although there have been only three human fatalities related to MVCs in Massachusetts since 2003, the human fatality rate was one of the highest reported in the literature. The rate of MVCs relative to the size of the moose population and the risk to human safety suggest a need for road mitigation measures, such as fencing, animal detection systems, and large mammal-crossing structures on roadways in Massachusetts.

Massachusetts↗

USGS2023

Genetic diversity and connectivity of moose (Alces americanus americanus) in eastern North America

Genetic diversity is critical to a population’s ability to overcome gradual environment change. Large-bodied wildlife existing in regions with relatively high human population density are vulnerable to isolation-induced genetic drift, population bottlenecks, and loss of genetic diversity. Moose ( Alces americanus americanus ) in eastern North America have a complex history of drastic population changes. Current and potential threats to moose populations in this region could be exacerbated by loss of genetic diversity and connectivity among subpopulations. Existing genetic diversity, gene flow, and population clustering and fragmentation of eastern North American moose are not well quantified, while physical and anthropogenic barriers to population connectivity already exist. Here, single nucleotide polymorphism (SNP) genotyping of 507 moose spanning five northeastern U.S. states and one southeastern Canadian province indicated low diversity, with a high proportion of the genomes sharing identity-by-state, with no consistent evidence of non-random mating. Gene flow estimates indicated bidirectionality between all pairs of sampled areas, with magnitudes reflecting clustering and differentiation patterns. A Discriminant Analysis of Principal Components analysis indicated that these genotypic data were best described with four clusters and indicated connectivity across the Saint Lawrence River and Seaway, a potential physical barrier to gene flow. Tests for genetic differentiation indicated restricted gene flow between populations across the Saint Lawrence River and Seaway, and between many sampled areas facing expanding human activity. These results document current genetic variation and connectivity of moose populations in eastern North America, highlight potential challenges to current population connectivity, and identify areas for future research and conservation.

New York, Vermont, New Hampshire, Maine, and Massa↗

USGS2018

Range expansion in unfavorable environments through behavioral responses to microclimatic conditions: Moose (Alces americanus) as the model

Wildlife populations occurring at the edge of their range boundaries are thought to be the most sensitive to climate change due to temperatures being at or near the limit of a species’ thermal envelope. Moose ( Alces americanus ) are a cold adapted species that are showing population declines in some portions of the southern edge of their range. However, other moose populations are actively expanding southward into thermally stressful areas. The direct effects of temperature on moose have not yet been studied in these southwardly expanding populations and may offer insights into how moose are successfully establishing in areas at the edge of their thermal envelope. We used ambient temperature and GPScollar data from moose to quantify the direct effect of temperature on moose habitat use in Massachusetts, USA, which is one of these southwardly expanding populations. The mean daily temperature in our study area exceeded the reported physiological tolerances of moose in over 90% of daytime and 75% of nighttime locations in summer and in over 80% of daytime and 67% of nighttime locations in winter. Across seasons and times of day, moose preferred regenerating forest, but as ambient air temperatures increased, selection for regenerating forest declined and selection for forested wetlands and coniferous forestincreased. This response indicates moose are altering their behavior to utilize thermal shelters when temperatures are high. We observed higher temperatures and stronger behavioral responses than other studies at the southern edge of moose range. We found habitat for moose in Massachusetts is climatically marginal and loss of habitat, increase in parasites, and further climatic warming may cause population declines in the future.

Mammalian Biology↗

USGS2017

White-tailed deer (Odocoileus virginianus) subsidize gray wolves (Canis lupus) during a moose (Alces americanus) decline: A case of apparent competition?

Moose ( Alces americanus ) in northeastern Minnesota have declined by 55% since 2006. Although the cause is unresolved, some studies have suggested that Gray Wolves ( Canis lupus ) contributed to the decline. After the Moose decline, wolves could either decline or switch prey. To determine which occurred in our study area, we compared winter wolf counts and summer diet before and after the Moose decline. While wolf numbers in our study area nearly doubled from 23 in winter 2002 to an average of 41 during winters 2011–2013, calf:cow ratios (the number of calves per cow observed during winter surveys) in the wider Moose range more than halved from 0.93 in 2002 to an average of 0.31 during 2011–2013. Compared to summer 2002, wolves in summers 2011–2013 consumed fewer Moose and more White-tailed Deer ( Odocoileus virginianus ). While deer densities were similar during each period, average vulnerability, as reflected by winter severity, was greater during 2011–2013 than 2002, probably explaining the wolf increase. During the wolf increase Moose calves remained a summer food item. These findings suggest that in part of the Moose range, deer subsidized wolf numbers while wolves also preyed on Moose calves. This contributed to a Moose decline and is a possible case of apparent competition and inverse-density-dependent predation.

Canadian Field-Naturalist↗

USGS2016

Range expansion of moose in arctic Alaska linked to warming and increased shrub habitat

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

Alaska↗

USGS2006

Habitat-based adaptive management at Mount Haggin Wildlife Management Area

The 22,743-hectare Mount Haggin Wildlife Management Area was purchased in 1976, in part for moose (Alces alces) winter range. Observed moose populations climbed from a low of 7 in 1976 to a high of 56 in 2000. A 4-step management program was initiated in 2000 consisting of definition of management objective, monitoring to determine if the objective was attained, developing a management strategy, and implementing the strategy. The management objective for browse was defined to be: browsing will not preventyoung plants from attaining their potential stature, their growth being primarily limited by local environmental conditions. Asurvey of Geyer willow (Salix geyeriana) in critical moose habitat indicated that browse plants were 100% intensely browsed, suggesting that browsing could prevent willowheightgrowth. Beginning in 2000, willow trend was monitored annually at 4 sites using an index based on the height of the tallest live stem and the height of the tallest, dead intensely browsed stem (LD Index). Low LD Index values indicated that browsing did prevent height growth. In 2000 moose harvest quotas were increased by 40%; in 2002 harvest quotas were increased an additional 7%. From 2000 to 2002, willow growth increased at all 4 locations. From 2002 to 2004, growth indicators changed relatively little at Sullivan Creek, Deep Creek, and French Creek; at these sites willow condition in 2004 had improved compared to willow condition in 2000. From 2002 to 2004, growth indicators declined markedly at American Creek; in 2004, growth indicators at American Creek were lower compared to measurements made in 2000. The improvement of willow condition at 3 sites was likely due to a combination of reduced moose numbers (due to an increase in harvest) and increased dispersal (due to low snow-cover conditions). Over the study period, the sporting public complained of reduced moose sightability; harvest quotas were lowered substantially in 2003.

Alaska↗

USGS2004

Relative spatial distributions and habitat use patterns of sympatric moose and white-tailed deer in Voyageurs National Park, Minnesota

We examined the distribution and home range characteristics of moose ( Alces alces ) and white-tailed deer ( Odocoileus virginianus ) at Voyageurs National Park, Minnesota. Pellet count transects revealed low densities of moose and higher densities of white-tailed deer, and provided evidence of partial spatial segregation between moose and white-tailed deer possibly due to habitat heterogeneity. There was limited interspecific overlap in the relatively large annual home ranges of radio-collared moose and white-tailed deer. Both moose and white-tailed deer exhibited significant selection for spruce ( Picea spp.) and balsam fir ( Abies balsamea ) vegetation types at the home range scale. White-tailed deer significantly selected a 12-20 m canopy height over all others while moose significantly selected 5-11 m and 21-30 m canopy heights over the 12-20 m canopy height. Moose significantly selected open/discontinuous canopy cover and white-tailed deer selected both closed/continuous and open/discontinuous canopy covers over dispersed/ sparse canopy cover. Differential habitat selection between moose and white-tailed deer at Voyageurs National Park might be related to the differences between these species' abilities to cope with a northern mid-continental climate. Spatial segregation between moose and white-tailed deer at Voyageurs National Park may allow moose to persist despite the presence of meningeal worm (P arelaphostrongylus tenuis ) in white-tailed deer.

Minnesota↗

USGS2001

Utility of stable isotope analysis in studying foraging ecology of herbivores: Examples from moose and caribou

Recently, researchers emphasized that patterns of stable isotope ratios observed at the individual level are a result of an interaction between ecological, physiological, and biochemical processes. Isotopic models for herbivores provide additional complications because those mammals consume foods that have high variability in nitrogen concentrations. In addition, distribution of amino acids in plants may differ greatly from that required by a herbivore. At northern latitudes, where the growing season of vegetation is short, isotope ratios in herbivore tissues are expected to differ between seasons. Summer ratios likely reflect diet composition, whereas winter ratios would reflect diet and nutrient recycling by the animals. We tested this hypothesis using data collected from blood samples of caribou ( Rangifer tarandus ) and moose ( Alces alces ) in Denali National Park and Preserve, Alaska, USA. Stable isotope ratios of moose and caribou were significantly different from each other in late summer-autumn and winter. Also, late summer-autumn and winter ratios differed significantly between seasons in both species. Nonetheless, we were unable to evaluate whether differences in seasonal isotopic ratios were a result of diet selection or a response to nutrient recycling. We believe that additional studies on plant isotopic ratios as related to ecological factors in conjunction with investigations of diet selection by the herbivores will enhance our understanding of those interactions. Also, controlled studies investigating the relation between diet and physiological responses in herbivores will increase the utility of isotopic analysis in studying foraging ecology of herbivores.

Alaska↗

USGS1991

Application and partial validation of a habitat model for moose in the Lake Superior region

A modified version of the dormant-season portion of a Habitat Suitability Index (HSI) model developed for assessing moose ( Alces alces ) habitat in the Lake Superior Region was incorporated in a Geographic Information System (GIS) for 490 km 2 of Minnesota's Superior National Forest. Moose locations (n=235) were plotted during aerial surveys conducted in December 1988 and January 1990-1991. Dormant-season forage and cover quality for 1,000-m, 500-m, and 200-m radii plots around random points and moose locations were compared using U.S. Forest Service stand examination data. Cover quality indices were lower than forage quality indices within all plots. The median value for the average cover quality index was greater (P=0.003) within 200-m plots around cow moose locations than for plots around random points for the most severe winter of the study. The proportion of highest-quality winter cover, such as mixed stands dominated by mid-age class white spruce ( Picea glauca ) and balsam fir ( Abies balsanea ), was greater within 500-m and 200-m plots around cow moose than within similar plots around random points during the two most severe winters. These results indicate that suboptimum ratings of winter habitat quality used in the GIS for dormant-season forage >100 m from cover, as suggested in the original HSI model, are reasonable. Integrating the habitat model with forest stand data using a GIS permitted analysis of moose habitat within a relatively large geographic area. Simulation of habitat quality indicated a potential shortage of late-winter cover in the study area. The effects of forest management actions on moose habitat quality can be simulated without collecting additional data.

Lake Superior region↗